SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC.

The Effects of Cross & Self-Fertilisation in the Vegetable Kingdom by Charles Darwin, is part of the HackerNoon Books Series. You can jump to any chapter in this book here. SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC.

SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC.

In the family of the Scrophulariaceae I experimented on species in the six following genera: Mimulus, Digitalis, Calceolaria, Linaria, Verbascum, and Vandellia.

[3/2. SCROPHULARIACEAE.—Mimulus luteus.

The plants which I raised from purchased seed varied greatly in the colour of their flowers, so that hardly two individuals were quite alike; the corolla being of all shades of yellow, with the most diversified blotches of purple, crimson, orange, and coppery brown. But these plants differed in no other respect. (3/1. I sent several specimens with variously coloured flowers to Kew, and Dr. Hooker informs me that they all consisted of Mimulus luteus. The flowers with much red have been named by horticulturists as var. Youngiana.) The flowers are evidently well adapted for fertilisation by the agency of insects; and in the case of a closely allied species, Mimulus rosea, I have watched bees entering the flowers, thus getting their backs well dusted with pollen; and when they entered another flower the pollen was licked off their backs by the two-lipped stigma, the lips of which are irritable and close like a forceps on the pollen-grains. If no pollen is enclosed between the lips, these open again after a time. Mr. Kitchener has ingeniously explained the use of these movements, namely, to prevent the self-fertilisation of the flower. (3/2. ‘A Year’s Botany’ 1874 page 118.) If a bee with no pollen on its back enters a flower it touches the stigma, which quickly closes, and when the bee retires dusted with pollen, it can leave none on the stigma of the same flower. But as soon as it enters any other flower, plenty of pollen is left on the stigma, which will be thus cross-fertilised. Nevertheless, if insects are excluded, the flowers fertilise themselves perfectly and produce plenty of seed; but I did not ascertain whether this is effected by the stamens increasing in length with advancing age, or by the bending down of the pistil. The chief interest in my experiments on the present species, lies in the appearance in the fourth self-fertilised generation of a variety which bore large peculiarly-coloured flowers, and grew to a greater height than the other varieties; it likewise became more highly self-fertile, so that this variety resembles the plant named Hero, which appeared in the sixth self-fertilised generation of Ipomoea.

Some flowers on one of the plants raised from the purchased seeds were fertilised with their own pollen; and others on the same plant were crossed with pollen from a distinct plant. The seeds from twelve capsules thus produced were placed in separate watch-glasses for comparison; and those from the six crossed capsules appeared to the eye hardly more numerous than those from the six self-fertilised capsules. But when the seeds were weighed, those from the crossed capsules amounted to 1.02 grain, whilst those from the self-fertilised capsules were only .81 grain; so that the former were either heavier or more numerous than the latter, in the ratio of 100 to 79.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIRST GENERATION.

Having ascertained, by leaving crossed and self-fertilised seed on damp sand, that they germinated simultaneously, both kinds were thickly sown on opposite sides of a broad and rather shallow pan; so that the two sets of seedlings, which came up at the same time, were subjected to the same unfavourable conditions. This was a bad method of treatment, but this species was one of the first on which I experimented. When the crossed seedlings were on an average half an inch high, the self-fertilised ones were only a quarter of an inch high. When grown to their full height under the above unfavourable conditions, the four tallest crossed plants averaged 7.62, and the four tallest self-fertilised 5.87 inches in height; or as 100 to 77. Ten flowers on the crossed plants were fully expanded before one on the self-fertilised plants. A few of these plants of both lots were transplanted into a large pot with plenty of good earth, and the self-fertilised plants, not now being subjected to severe competition, grew during the following year as tall as the crossed plants; but from a case which follows it is doubtful whether they would have long continued equal. Some flowers on the crossed plants were crossed with pollen from another plant, and the capsules thus produced contained a rather greater weight of seed than those on the self-fertilised plants again self-fertilised.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

Seeds from the foregoing plants, fertilised in the manner just stated, were sown on the opposite sides of a small pot (1) and came up crowded. The four tallest crossed seedlings, at the time of flowering, averaged 8 inches in height, whilst the four tallest self-fertilised plants averaged only 4 inches. Crossed seeds were sown by themselves in a second small pot, and self-fertilised seeds were sown by themselves in a third small pot so that there was no competition whatever between these two lots. Nevertheless the crossed plants grew from 1 to 2 inches higher on an average than the self-fertilised. Both lots looked equally vigorous, but the crossed plants flowered earlier and more profusely than the self-fertilised. In Pot 1, in which the two lots competed with each other, the crossed plants flowered first and produced a large number of capsules, whilst the self-fertilised produced only nineteen. The contents of twelve capsules from the crossed flowers on the crossed plants, and of twelve capsules from self-fertilised flowers on the self-fertilised plants, were placed in separate watch-glasses for comparison; and the crossed seeds seemed more numerous by half than the self-fertilised.

The plants on both sides of Pot 1, after they had seeded, were cut down and transplanted into a large pot with plenty of good earth, and on the following spring, when they had grown to a height of between 5 and 6 inches, the two lots were equal, as occurred in a similar experiment in the last generation. But after some weeks the crossed plants exceeded the self-fertilised ones on the opposite side of the same pot, though not nearly to so great a degree as before, when they were subjected to very severe competition.

CROSSED AND SELF-FERTILISED PLANTS OF THE THIRD GENERATION.

Crossed seeds from the crossed plants, and self-fertilised seeds from the self-fertilised plants of the last generation, were sown thickly on opposite sides of a small pot, Number 1. The two tallest plants on each side were measured after they had flowered, and the two crossed ones were 12 and 7 1/2 inches, and the two self-fertilised ones 8 and 5 1/2 inches in height; that is, in the ratio of 100 to 69. Twenty flowers on the crossed plants were again crossed and produced twenty capsules; ten of which contained 1.33 grain weight of seeds. Thirty flowers on the self-fertilised plants were again self-fertilised and produced twenty-six capsules; ten of the best of which (many being very poor) contained only .87 grain weight of seeds; that is, in the ratio of 100 to 65 by weight.

The superiority of the crossed over the self-fertilised plants was proved in various ways. Self-fertilised seeds were sown on one side of a pot, and two days afterwards crossed seeds on the opposite side. The two lots of seedlings were equal until they were above half an inch high; but when fully grown the two tallest crossed plants attained a height of 12 1/2 and 8 3/4 inches, whilst the two tallest self-fertilised plants were only 8 and 5 1/2 inches high.

In a third pot, crossed seeds were sown four days after the self-fertilised, and the seedlings from the latter had at first, as might have been expected, an advantage; but when the two lots were between 5 and 6 inches in height, they were equal, and ultimately the three tallest crossed plants were 11, 10, and 8 inches, whilst the three tallest self-fertilised were 12, 8 1/2, and 7 1/2 inches in height. So that there was not much difference between them, the crossed plants having an average advantage of only the third of an inch. The plants were cut down, and without being disturbed were transplanted into a larger pot. Thus the two lots started fair on the following spring, and now the crossed plants showed their inherent superiority, for the two tallest were 13 inches, whilst the two tallest self-fertilised plants were only 11 and 8 1/2 inches in height; or as 100 to 75. The two lots were allowed to fertilise themselves spontaneously: the crossed plants produced a large number of capsules, whilst the self-fertilised produced very few and poor ones. The seeds from eight of the capsules on the crossed plants weighed .65 grain, whilst those from eight of the capsules on the self-fertilised plants weighed only .22 grain; or as 100 to 34.

The crossed plants in the above three pots, as in almost all the previous experiments, flowered before the self-fertilised. This occurred even in the third pot in which the crossed seeds were sown four days after the self-fertilised seeds.

Lastly, seeds of both lots were sown on opposite sides of a large pot in which a Fuchsia had long been growing, so that the earth was full of roots. Both lots grew miserably; but the crossed seedlings had an advantage at all times, and ultimately attained to a height of 3 1/2 inches, whilst the self-fertilised seedlings never exceeded 1 inch. The several foregoing experiments prove in a decisive manner the superiority in constitutional vigour of the crossed over the self-fertilised plants.

In the three generations now described and taken together, the average height of the ten tallest crossed plants was 8.19 inches, and that of the ten tallest self-fertilised plants 5.29 inches (the plants having been grown in small pots), or as 100 to 65.

In the next or fourth self-fertilised generation, several plants of a new and tall variety appeared, which increased in the later self-fertilised generations, owing to its great self-fertility, to the complete exclusion of the original kinds. The same variety also appeared amongst the crossed plants, but as it was not at first regarded with any particular attention, I know not how far it was used for raising the intercrossed plants; and in the later crossed generations it was rarely present. Owing to the appearance of this tall variety, the comparison of the crossed and self-fertilised plants of the fifth and succeeding generations was rendered unfair, as all the self-fertilised and only a few or none of the crossed plants consisted of it. Nevertheless, the results of the later experiments are in some respects well worth giving.

CROSSED AND SELF-FERTILISED PLANTS OF THE FOURTH GENERATION.

Seeds of the two kinds, produced in the usual way from the two sets of plants of the third generation, were sown on opposite sides of two pots (1 and 2); but the seedlings were not thinned enough and did not grow well. Many of the self-fertilised plants, especially in one of the pots, consisted of the new and tall variety above referred to, which bore large and almost white flowers marked with crimson blotches. I will call it the WHITE VARIETY. I believe that it first appeared amongst both the crossed and self-fertilised plants of the last generation; but neither my gardener nor myself could remember any such variety in the seedlings raised from the purchased seed. It must therefore have arisen either through ordinary variation, or, judging from its appearance amongst both the crossed and self-fertilised plants, more probably through reversion to a formerly existing variety.

In Pot 1 the tallest crossed plant was 8 1/2 inches, and the tallest self-fertilised 5 inches in height. In Pot 2, the tallest crossed plant was 6 1/2 inches, and the tallest self-fertilised plant, which consisted of the white variety, 7 inches in height; and this was the first instance in my experiments on Mimulus in which the tallest self-fertilised plant exceeded the tallest crossed. Nevertheless, the two tallest crossed plants taken together were to the two tallest self-fertilised plants in height as 100 to 80. As yet the crossed plants were superior to the self-fertilised in fertility; for twelve flowers on the crossed plants were crossed and yielded ten capsules, the seeds of which weighed 1.71 grain. Twenty flowers on the self-fertilised plants were self-fertilised, and produced fifteen capsules, all appearing poor; and the seeds from ten of them weighed only .68 grain, so that from an equal number of capsules the crossed seeds were to the self-fertilised in weight as 100 to 40.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIFTH GENERATION.

Seeds from both lots of the fourth generation, fertilised in the usual manner, were sown on opposite sides of three pots. When the seedlings flowered, most of the self-fertilised plants were found to consist of the tall white variety. Several of the crossed plants in Pot 1 likewise belonged to this variety, as did a very few in Pots 2 and 3. The tallest crossed plant in Pot 1 was 7 inches, and the tallest self-fertilised plant on the opposite side 8 inches; in Pots 2 and 3 the tallest crossed were 4 1/2 and 5 1/2, and the tallest self-fertilised 7 and 6 1/2 inches in height; so that the average height of the tallest plants in the two lots was as 100 for the crossed to 126 for the self-fertilised; and thus we have a complete reversal of what occurred in the four previous generations. Nevertheless, in all three pots the crossed plants retained their habit of flowering before the self-fertilised. The plants were unhealthy from being crowded and from the extreme heat of the season, and were in consequence more or less sterile; but the crossed plants were somewhat less sterile than the self-fertilised plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SIXTH GENERATION.

Seeds from plants of the fifth generation crossed and self-fertilised in the usual manner were sown on opposite sides of several pots. On the self-fertilised side every single plant belonged to the tall white variety. On the crossed side some plants belonged to this variety, but the greater number approached in character to the old and shorter kinds with smaller yellowish flowers blotched with coppery brown. When the plants on both sides were from 2 to 3 inches in height they were equal, but when fully grown the self-fertilised were decidedly the tallest and finest plants, but, from want of time, they were not actually measured. In half the pots the first plant which flowered was a self-fertilised one, and in the other half a crossed one. And now another remarkable change was clearly perceived, namely, that the self-fertilised plants had become more self-fertile than the crossed. The pots were all put under a net to exclude insects, and the crossed plants produced spontaneously only fifty-five capsules, whilst the self-fertilised plants produced eighty-one capsules, or as 100 to 147. The seeds from nine capsules of both lots were placed in separate watch-glasses for comparison, and the self-fertilised appeared rather the more numerous. Besides these spontaneously self-fertilised capsules, twenty flowers on the crossed plants again crossed yielded sixteen capsules; twenty-five flowers on the self-fertilised plants again self-fertilised yielded seventeen capsules, and this is a larger proportional number of capsules than was produced by the self-fertilised flowers on the self-fertilised plants in the previous generations. The contents of ten capsules of both these lots were compared in separate watch-glasses, and the seeds from the self-fertilised appeared decidedly more numerous than those from the crossed plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SEVENTH GENERATION.

Crossed and self-fertilised seeds from the crossed and self-fertilised plants of the sixth generation were sown in the usual manner on opposite sides of three pots, and the seedlings were well and equally thinned. Every one of the self-fertilised plants (and many were raised) in this, as well as in the eighth and ninth generations, belonged to the tall white variety. Their uniformity of character, in comparison with the seedlings first raised from the purchased seed, was quite remarkable. On the other hand, the crossed plants differed much in the tints of their flowers, but not, I think, to so great a degree as those first raised. I determined this time to measure the plants on both sides carefully. The self-fertilised seedlings came up rather before the crossed, but both lots were for a time of equal height. When first measured, the average height of the six tallest crossed plants in the three pots was 7.02, and that of the six tallest self-fertilised plants 8.97 inches, or as 100 to 128. When fully grown the same plants were again measured, with the result shown in Table 3/18.

TABLE 3/18. Mimulus luteus (Seventh Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 11 2/8 : 19 1/8. Pot 1 : 11 7/8 : 18.

Pot 2 : 12 6/8 : 18 2/8. Pot 2 : 11 2/8 : 14 6/8.

Pot 3 : 9 6/8 : 12 6/8. Pot 3 : 11 6/8 : 11.

Total : 68.63 : 93.88.

The average height of the six crossed is here 11.43, and that of the six self-fertilised 15.64, or as 100 to 137.

As it is now evident that the tall white variety transmitted its characters faithfully, and as the self-fertilised plants consisted exclusively of this variety, it was manifest that they would always exceed in height the crossed plants which belonged chiefly to the original shorter varieties. This line of experiment was therefore discontinued, and I tried whether intercrossing two self-fertilised plants of the sixth generation, growing in distinct pots, would give their offspring any advantage over the offspring of flowers on one of the same plants fertilised with their own pollen. These latter seedlings formed the seventh generation of self-fertilised plants, like those in the right hand column in Table 3/18; the crossed plants were the product of six previous self-fertilised generations with an intercross in the last generation. The seeds were allowed to germinate on sand, and were planted in pairs on opposite sides of four pots, all the remaining seeds being sown crowded on opposite sides of Pot 5 in Table 3/19; the three tallest on each side in this latter pot being alone measured. All the plants were twice measured—the first time whilst young, and the average height of the crossed plants to that of the self-fertilised was then as 100 to 122. When fully grown they were again measured, as in Table 3/19.

TABLE 3/19. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Intercrossed Plants from Self-fertilised Plants of the Sixth Generation.

Column 3: Self-fertilised Plants of the Seventh Generation.

Pot 1 : 12 6/8 : 15 2/8. Pot 1 : 10 4/8 : 11 5/8. Pot 1 : 10 : 11. Pot 1 : 14 5/8 : 11.

Pot 2 : 10 2/8 : 11 3/8. Pot 2 : 7 6/8 : 11 4/8. Pot 2 : 12 1/8 : 8 5/8. Pot 2 : 7 : 14 3/8.

Pot 3 : 13 5/8 : 10 3/8. Pot 3 : 12 2/8 : 11 6/8.

Pot 4 : 7 1/8 : 14 6/8. Pot 4 : 8 2/8 : 7. Pot 4 : 7 2/8 : 8.

Pot 5 : 8 5/8 : 10 2/8 Pot 5 : 9 : 9 3/8. Pot 5 : 8 2/8 : 9 2/8. Crowded.

Total : 159.38 : 175.50.

The average height of the sixteen intercrossed plants is here 9.96 inches, and that of the sixteen self-fertilised plants 10.96, or as 100 to 110; so that the intercrossed plants, the progenitors of which had been self-fertilised for the six previous generations, and had been exposed during the whole time to remarkably uniform conditions, were somewhat inferior in height to the plants of the seventh self-fertilised generation. But as we shall presently see that a similar experiment made after two additional generations of self-fertilisation gave a different result, I know not how far to trust the present one. In three of the five pots in Table 3/19 a self-fertilised plant flowered first, and in the other two a crossed plant. These self-fertilised plants were remarkably fertile, for twenty flowers fertilised with their own pollen produced no less than nineteen very fine capsules!

THE EFFECTS OF A CROSS WITH A DISTINCT STOCK.

Some flowers on the self-fertilised plants in Pot 4 in Table 3/19 were fertilised with their own pollen, and plants of the eighth self-fertilised generation were thus raised, merely to serve as parents in the following experiment. Several flowers on these plants were allowed to fertilise themselves spontaneously (insects being of course excluded), and the plants raised from these seeds formed the ninth self-fertilised generation; they consisted wholly of the tall white variety with crimson blotches. Other flowers on the same plants of the eighth self-fertilised generation were crossed with pollen taken from another plant of the same lot; so that the seedlings thus raised were the offspring of eight previous generations of self-fertilisation with an intercross in the last generation; these I will call the INTERCROSSED PLANTS. Lastly, other flowers on the same plants of the eighth self-fertilised generation were crossed with pollen taken from plants which had been raised from seed procured from a garden at Chelsea. The Chelsea plants bore yellow flowers blotched with red, but differed in no other respect. They had been grown out of doors, whilst mine had been cultivated in pots in the greenhouse for the last eight generations, and in a different kind of soil. The seedlings raised from this cross with a wholly different stock may be called the CHELSEA-CROSSED. The three lots of seeds thus obtained were allowed to germinate on bare sand; and whenever a seed in all three lots, or in only two, germinated at the same time, they were planted in pots superficially divided into three or two compartments. The remaining seeds, whether or not in a state of germination, were thickly sown in three divisions in a large pot, 10, in Table 3/20. When the plants had grown to their full height they were measured, as shown in Table 3/20; but only the three tallest plants in each of the three divisions in Pot 10 were measured.

TABLE 3/20. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants from Self-fertilised Plants of the Eighth Generation crossed by Chelsea Plants.

Column 3: Plants from an intercross between the Plants of the Eighth Self-fertilised Generation.

Column 4: Self-fertilised Plants of the Ninth Generation from Plants of the Eighth Self-fertilised Generation.

Pot 1 : 30 7/8 : 14 : 9 4/8. Pot 1 : 28 3/8 : 13 6/8 : 10 5/8. Pot 1 : — : 13 7/8 : 10.

Pot 2 : 20 6/8 : 11 4/8 : 11 6/8. Pot 2 : 22 2/8 : 12 : 12 3/8. Pot 2 : — : 9 1/8 : —.

Pot 3 : 23 6/8 : 12 2/8 : 8 5/8. Pot 3 : 24 1/8 : — : 11 4/8. Pot 3 : 25 6/8 : — : 6 7/8.

Pot 4 : 22 5/8 : 9 2/8 : 4. Pot 4 : 22 : 8 1/8 : 13 3/8. Pot 4 : 17 : — : 11.

Pot 5 : 22 3/8 : 9 : 4 4/8. Pot 5 : 19 5/8 : 11 : 13. Pot 5 : 23 4/8 : — : 13 4/8.

Pot 6 : 28 2/8 : 18 6/8 : 12. Pot 6 : 22 : 7 : 16 1/8. Pot 6 : — : 12 4/8 : —.

Pot 7 : 12 4/8 : 15 : —. Pot 7 : 24 3/8 : 12 3/8 : —. Pot 7 : 20 4/8 : 11 2/8 : —. Pot 7 : 26 4/8 : 15 2/8 : —.

Pot 8 : 17 2/8 : 13 3/8 : —. Pot 8 : 22 6/8 : 14 5/8 : —. Pot 8 : 27 : 14 3/8 : —.

Pot 9 : 22 6/8 : 11 6/8 : —. Pot 9 : 6 : 17 : —. Pot 9 : 20 2/8 : 14 7/8 : —.

Pot 10 : 18 1/8 : 9 2/8 : 10 3/8. Pot 10 : 16 5/8 : 8 2/8 : 8 1/8. Pot 10 : 17 4/8 : 10 : 11 2/8. Crowded plants.

Total : 605.38 : 329.50 : 198.50.

In this table the average height of the twenty-eight Chelsea-crossed plants is 21.62 inches; that of the twenty-seven intercrossed plants 12.2; and that of the nineteen self-fertilised 10.44. But with respect to the latter it will be the fairest plan to strike out two dwarfed ones (only 4 inches in height), so as not to exaggerate the inferiority of the self-fertilised plants; and this will raise the average height of the seventeen remaining self-fertilised plants to 11.2 inches. Therefore the Chelsea-crossed are to the intercrossed in height as 100 to 56; the Chelsea-crossed to the self-fertilised as 100 to 52; and the intercrossed to the self-fertilised as 100 to 92. We thus see how immensely superior in height the Chelsea-crossed are to the intercrossed and to the self-fertilised plants. They began to show their superiority when only one inch high. They were also, when fully grown, much more branched with larger leaves and somewhat larger flowers than the plants of the other two lots, so that if they had been weighed, the ratio would certainly have been much higher than that of 100 to 56 and 52.

The intercrossed plants are here to the self-fertilised in height as 100 to 92; whereas in the analogous experiment given in Table 3/19 the intercrossed plants from the self-fertilised plants of the sixth generation were inferior in height to the self-fertilised plants in the ratio of 100 to 110. I doubt whether this discordance in the results of the two experiments can be explained by the self-fertilised plants in the present case having been raised from spontaneously self-fertilised seeds, whereas in the former case they were raised from artificially self-fertilised seeds; nor by the present plants having been self-fertilised during two additional generations, though this is a more probable explanation.

With respect to fertility, the twenty-eight Chelsea-crossed plants produced 272 capsules; the twenty-seven intercrossed plants produced 24; and the seventeen self-fertilised plants 17 capsules. All the plants were left uncovered so as to be naturally fertilised, and empty capsules were rejected.

Therefore 20 Chelsea-crossed plants would have produced 194.29 capsules.

Therefore 20 Intercrossed plants would have produced 17.77 capsules.

Therefore 20 Self-fertilised plants would have produced 20.00 capsules.

The seeds contained in 8 capsules from the Chelsea-crossed plants weighed 1.1 grains.

The seeds contained in 8 capsules from the Intercrossed plants weighed 0.51 grains.

The seeds contained in 8 capsules from the Self-fertilised plants weighed 0.33 grains.

If we combine the number of capsules produced together with the average weight of contained seeds, we get the following extraordinary ratios:

Weight of seed produced by the same number of Chelsea-crossed and intercrossed plants as 100 to 4.

Weight of seed produced by the same number of Chelsea-crossed and self-fertilised plants as 100 to 3.

Weight of seeds produced by the same number of intercrossed and self-fertilised plants as 100 to 73.

It is also a remarkable fact that the Chelsea-crossed plants exceeded the two other lots in hardiness, as greatly as they did in height, luxuriance, and fertility. In the early autumn most of the pots were bedded out in the open ground; and this always injures plants which have been long kept in a warm greenhouse. All three lots consequently suffered greatly, but the Chelsea-crossed plants much less than the other two lots. On the 3rd of October the Chelsea-crossed plants began to flower again, and continued to do so for some time; whilst not a single flower was produced by the plants of the other two lots, the stems of which were cut almost down to the ground and seemed half dead. Early in December there was a sharp frost, and the stems of Chelsea-crossed were now cut down; but on the 23rd of December they began to shoot up again from the roots, whilst all the plants of the other two lots were quite dead.

Although several of the self-fertilised seeds, from which the plants in the right hand column in Table 3/20 were raised, germinated (and were of course rejected) before any of those of the other two lots, yet in only one of the ten pots did a self-fertilised plant flower before the Chelsea-crossed or the intercrossed plants growing in the same pots. The plants of these two latter lots flowered at the same time, though the Chelsea-crossed grew so much taller and more vigorously than the intercrossed.

As already stated, the flowers of the plants originally raised from the Chelsea seeds were yellow; and it deserves notice that every one of the twenty-eight seedlings raised from the tall white variety fertilised, without being castrated, with pollen from the Chelsea plants, produced yellow flowers; and this shows how prepotent this colour, which is the natural one of the species, is over the white colour.

THE EFFECTS ON THE OFFSPRING OF INTERCROSSING FLOWERS ON THE SAME PLANT, INSTEAD OF CROSSING DISTINCT INDIVIDUALS.

In all the foregoing experiments the crossed plants were the product of a cross between distinct plants. I now selected a very vigorous plant in Table 3/20, raised by fertilising a plant of the eighth self-fertilised generation with pollen from the Chelsea stock. Several flowers on this plant were crossed with pollen from other flowers on the same plant, and several other flowers were fertilised with their own pollen. The seed thus produced was allowed to germinate on bare sand; and the seedlings were planted in the usual manner on the opposite sides of six pots. All the remaining seeds, whether or not in a state of germination, were sown thickly in Pot 7; the three tallest plants on each side of this latter pot being alone measured. As I was in a hurry to learn the result, some of these seeds were sown late in the autumn, but the plants grew so irregularly during the winter, that one crossed plant was 28 1/2 inches, and two others only 4, or less than 4 inches in height, as may be seen in Table 3/21. Under such circumstances, as I have observed in many other cases, the result is not in the least trustworthy; nevertheless I feel bound to give the measurements.

TABLE 3/21. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 17 : 17. Pot 1 : 9 : 3 1/8.

Pot 2 : 28 2/8 : 19 1/8. Pot 2 : 16 4/8 : 6. Pot 2 : 13 5/8 : 2.

Pot 3 : 4 : 15 6/8. Pot 3 : 2 2/8 : 10.

Pot 4 : 23 4/8 : 6 2/8. Pot 4 : 15 4/8 : 7 1/8.

Pot 5 : 7 : 13 4/8.

Pot 6 : 18 3/8 : 1 4/8. Pot 6 : 11 : 2.

Pot 7 : 21 : 15 1/8. Pot 7 : 11 6/8 : 11. Pot 7 : 12 1/8 : 11 2/8. Crowded.

Total : 210.88 : 140.75.

The fifteen crossed plants here average 14.05, and the fifteen self-fertilised plants 9.38 in height, or as 100 to 67. But if all the plants under ten inches in height are struck out, the ratio of the eleven crossed plants to the eight self-fertilised plants is as 100 to 82.

On the following spring, some remaining seeds of the two lots were treated in exactly the same manner; and the measurements of the seedlings are given in Table 3/22.

TABLE 3/22. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 15 1/8 : 19 1/8. Pot 1 : 12 : 20 5/8. Pot 1 : 10 1/8 : 12 6/8.

Pot 2 : 16 2/8 : 11 2/8. Pot 2 : 13 5/8 : 19 3/8. Pot 2 : 20 1/8 : 17 4/8.

Pot 3 : 18 7/8 : 12 6/8. Pot 3 : 15 : 15 6/8. Pot 3 : 13 7/8 : 17.

Pot 4 : 19 2/8 : 16 2/8. Pot 4 : 19 6/8 : 21 5/8.

Pot 5 : 25 3/8 : 22 5/8.

Pot 6 : 15 : 19 5/8. Pot 6 : 20 2/8 : 16 2/8. Pot 6 : 27 2/8 : 19 5/8.

Pot 7 : 7 6/8 : 7 6/8. Pot 7 : 14 : 8. Pot 7 : 13 4/8 : 7.

Pot 8 : 18 2/8 : 20 3/8. Pot 8 : 18 6/8 : 17 6/8. Pot 8 : 18 3/8 : 15 4/8. Pot 8 : 18 3/8 : 15 1/8. Crowded.

Total : 370.88 : 353.63.

Here the average height of the twenty-two crossed plants is 16.85, and that of the twenty-two self-fertilised plants 16.07; or as 100 to 95. But if four of the plants in Pot 7, which are much shorter than any of the others, are struck out (and this would be the fairest plan), the twenty-one crossed are to the nineteen self-fertilised plants in height as 100 to 100.6—that is, are equal. All the plants, except the crowded ones in Pot 8, after being measured were cut down, and the eighteen crossed plants weighed 10 ounces, whilst the same number of self-fertilised plants weighed 10 1/4 ounces, or as 100 to 102.5; but if the dwarfed plants in Pot 7 had been excluded, the self-fertilised would have exceeded the crossed in weight in a higher ratio. In all the previous experiments in which seedlings were raised from a cross between distinct plants, and were put into competition with self-fertilised plants, the former generally flowered first; but in the present case, in seven out of the eight pots a self-fertilised plant flowered before a crossed one on the opposite side. Considering all the evidence with respect to the plants in Table3/ 22, a cross between two flowers on the same plant seems to give no advantage to the offspring thus produced, the self-fertilised plants being in weight superior. But this conclusion cannot be absolutely trusted, owing to the measurements given in Table 3/21, though these latter, from the cause already assigned, are very much less trustworthy than the present ones.]

A SUMMARY OF OBSERVATIONS ON Mimulus luteus.

In the three first generations of crossed and self-fertilised plants, the tallest plants alone on each side of the several pots were measured; and the average height of the ten crossed to that of the ten self-fertilised plants was as 100 to 64. The crossed were also much more fertile than the self-fertilised, and so much more vigorous that they exceeded them in height, even when sown on the opposite side of the same pot after an interval of four days. The same superiority was likewise shown in a remarkable manner when both kinds of seeds were sown on the opposite sides of a pot with very poor earth full of the roots of another plant. In one instance crossed and self-fertilised seedlings, grown in rich soil and not put into competition with each other, attained to an equal height. When we come to the fourth generation the two tallest crossed plants taken together exceeded by only a little the two tallest self-fertilised plants, and one of the latter beat its crossed opponent,—a circumstance which had not occurred in the previous generations. This victorious self-fertilised plant consisted of a new white-flowered variety, which grew taller than the old yellowish varieties. From the first it seemed to be rather more fertile, when self-fertilised, than the old varieties, and in the succeeding self-fertilised generations became more and more self-fertile. In the sixth generation the self-fertilised plants of this variety compared with the crossed plants produced capsules in the proportion of 147 to 100, both lots being allowed to fertilise themselves spontaneously. In the seventh generation twenty flowers on one of these plants artificially self-fertilised yielded no less than nineteen very fine capsules!

This variety transmitted its characters so faithfully to all the succeeding self-fertilised generations, up to the last or ninth, that all the many plants which were raised presented a complete uniformity of character; thus offering a remarkable contrast with the seedlings raised from the purchased seeds. Yet this variety retained to the last a latent tendency to produce yellow flowers; for when a plant of the eighth self-fertilised generation was crossed with pollen from a yellow-flowered plant of the Chelsea stock, every single seedling bore yellow flowers. A similar variety, at least in the colour of its flowers, also appeared amongst the crossed plants of the third generation. No attention was at first paid to it, and I know not how far it was at first used either for crossing or self-fertilisation. In the fifth generation most of the self-fertilised plants, and in the sixth and all the succeeding generations every single plant consisted of this variety; and this no doubt was partly due to its great and increasing self-fertility. On the other hand, it disappeared from amongst the crossed plants in the later generations; and this was probably due to the continued intercrossing of the several plants. From the tallness of this variety, the self-fertilised plants exceeded the crossed plants in height in all the generations from the fifth to the seventh inclusive; and no doubt would have done so in the later generations, had they been grown in competition with one another. In the fifth generation the crossed plants were in height to the self-fertilised, as 100 to 126; in the sixth, as 100 to 147; and in the seventh generation, as 100 to 137. This excess of height may be attributed not only to this variety naturally growing taller than the other plants, but to its possessing a peculiar constitution, so that it did not suffer from continued self-fertilisation.

This variety presents a strikingly analogous case to that of the plant called the Hero, which appeared in the sixth self-fertilised generation of Ipomoea. If the seeds produced by Hero had been as greatly in excess of those produced by the other plants, as was the case with Mimulus, and if all the seeds had been mingled together, the offspring of Hero would have increased to the entire exclusion of the ordinary plants in the later self-fertilised generations, and from naturally growing taller would have exceeded the crossed plants in height in each succeeding generation.

Some of the self-fertilised plants of the sixth generation were intercrossed, as were some in the eighth generation; and the seedlings from these crosses were grown in competition with self-fertilised plants of the two corresponding generations. In the first trial the intercrossed plants were less fertile than the self-fertilised, and less tall in the ratio of 100 to 110. In the second trial, the intercrossed plants were more fertile than the self-fertilised in the ratio of 100 to 73, and taller in the ratio of 100 to 92. Notwithstanding that the self-fertilised plants in the second trial were the product of two additional generations of self-fertilisation, I cannot understand this discordance in the results of the two analogous experiments.

The most important of all the experiments on Mimulus are those in which flowers on plants of the eighth self-fertilised generation were again self-fertilised; other flowers on distinct plants of the same lot were intercrossed; and others were crossed with a new stock of plants from Chelsea. The Chelsea-crossed seedlings were to the intercrossed in height as 100 to 56, and in fertility as 100 to 4; and they were to the self-fertilised plants, in height as 100 to 52, and in fertility as 100 to 3. These Chelsea-crossed plants were also much more hardy than the plants of the other two lots; so that altogether the gain from the cross with a fresh stock was wonderfully great.

Lastly, seedlings raised from a cross between flowers on the same plant were not superior to those from flowers fertilised with their own pollen; but this result cannot be absolutely trusted, owing to some previous observations, which, however, were made under very unfavourable circumstances.

[Digitalis purpurea.

The flowers of the common Foxglove are proterandrous; that is, the pollen is mature and mostly shed before the stigma of the same flower is ready for fertilisation. This is effected by the larger humble-bees, which, whilst in search of nectar, carry pollen from flower to flower. The two upper and longer stamens shed their pollen before the two lower and shorter ones. The meaning of this fact probably is, as Dr. Ogle remarks, that the anthers of the longer stamens stand near to the stigma, so that they would be the most likely to fertilise it (3/3. ‘Popular Science Review’ January 1870 page 50.); and as it is an advantage to avoid self-fertilisation, they shed their pollen first, thus lessening the chance. There is, however, but little danger of self-fertilisation until the bifid stigma opens; for Hildebrand found that pollen placed on the stigma before it had opened produced no effect. (3/4. ‘Geschlechter-Vertheilung bei den Pflanzen’ 1867 page 20.) The anthers, which are large, stand at first transversely with respect to the tubular corolla, and if they were to dehisce in this position they would, as Dr. Ogle also remarks, smear with pollen the whole back and sides of an entering humble-bee in a useless manner; but the anthers twist round and place themselves longitudinally before they dehisce. The lower and inner side of the mouth of the corolla is thickly clothed with hairs, and these collect so much of the fallen pollen that I have seen the under surface of a humble-bee thickly dusted with it; but this can never be applied to the stigma, as the bees in retreating do not turn their under surfaces upwards. I was therefore puzzled whether these hairs were of any use; but Mr. Belt has, I think, explained their use: the smaller kinds of bees are not fitted to fertilise the flowers, and if they were allowed to enter easily they would steal much nectar, and fewer large bees would haunt the flowers. Humble-bees can crawl into the dependent flowers with the greatest ease, using the “hairs as footholds while sucking the honey; but the smaller bees are impeded by them, and when, having at length struggled through them, they reach the slippery precipice above, they are completely baffled.” Mr. Belt says that he watched many flowers during a whole season in North Wales, and “only once saw a small bee reach the nectary, though many were seen trying in vain to do so.” (3/5. ‘The Naturalist in Nicaragua’ 1874 page 132. But it appears from H. Muller ‘Die Befruchtung der Blumen’ 1873 page 285, that small insects sometimes succeed in entering the flowers.)

I covered a plant growing in its native soil in North Wales with a net, and fertilised six flowers each with its own pollen, and six others with pollen from a distinct plant growing within the distance of a few feet. The covered plant was occasionally shaken with violence, so as to imitate the effects of a gale of wind, and thus to facilitate as far as possible self-fertilisation. It bore ninety-two flowers (besides the dozen artificially fertilised), and of these only twenty-four produced capsules; whereas almost all the flowers on the surrounding uncovered plants were fruitful. Of the twenty-four spontaneously self-fertilised capsules, only two contained their full complement of seed; six contained a moderate supply; and the remaining sixteen extremely few seeds. A little pollen adhering to the anthers after they had dehisced, and accidentally falling on the stigma when mature, must have been the means by which the above twenty-four flowers were partially self-fertilised; for the margins of the corolla in withering do not curl inwards, nor do the flowers in dropping off turn round on their axes, so as to bring the pollen-covered hairs, with which the lower surface is clothed, into contact with the stigma—by either of which means self-fertilisation might be effected.

Seeds from the above crossed and self-fertilised capsules, after germinating on bare sand, were planted in pairs on the opposite sides of five moderately-sized pots, which were kept in the greenhouse. The plants after a time appeared starved, and were therefore, without being disturbed, turned out of their pots, and planted in the open ground in two close parallel rows. They were thus subjected to tolerably severe competition with one another; but not nearly so severe as if they had been left in the pots. At the time when they were turned out, their leaves were between 5 and 8 inches in length, and the longest leaf on the finest plant on each side of each pot was measured, with the result that the leaves of the crossed plants exceeded, on an average, those of the self-fertilised plants by .4 of an inch.

In the following summer the tallest flower-stem on each plant, when fully grown, was measured. There were seventeen crossed plants; but one did not produce a flower-stem. There were also, originally, seventeen self-fertilised plants, but these had such poor constitutions that no less than nine died in the course of the winter and spring, leaving only eight to be measured, as in Table 3/23.

TABLE 3/23. Digitalis purpurea.

The tallest Flower-stem on each Plant measured in inches: 0 means that the Plant died before a Flower-stem was produced.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 53 6/8 : 27 4/8. Pot 1 : 57 4/8 : 55 6/8. Pot 1 : 57 6/8 : 0. Pot 1 : 65 : 0.

Pot 2 : 34 4/8 : 39. Pot 2 : 52 4/8 : 32. Pot 2 : 63 6/8 : 21.

Pot 3 : 57 4/8 : 53 4/8. Pot 3 : 53 4/8 : 0. Pot 3 : 50 6/8 : 0. Pot 3 : 37 2/8 : 0.

Pot 4 : 64 4/8 : 34 4/8. Pot 4 : 37 4/8 : 23 6/8. Pot 4 : — : 0.

Pot 5 : 53 : 0. Pot 5 : 47 6/8 : 0. Pot 5 : 34 6/8 : 0.

Total : 821.25 : 287.00.

The average height of the flower-stems of the sixteen crossed plants is here 51.33 inches; and that of the eight self-fertilised plants, 35.87; or as 100 to 70. But this difference in height does not give at all a fair idea of the vast superiority of the crossed plants. These latter produced altogether sixty-four flower-stems, each plant producing, on an average, exactly four flower-stems, whereas the eight self-fertilised plants produced only fifteen flower-stems, each producing an average only of 1.87 stems, and these had a less luxuriant appearance. We may put the result in another way: the number of flower-stems on the crossed plants was to those on an equal number of self-fertilised plants as 100 to 48.

Three crossed seeds in a state of germination were also planted in three separate pots; and three self-fertilised seeds in the same state in three other pots. These plants were therefore at first exposed to no competition with one another, and when turned out of their pots into the open ground they were planted at a moderate distance apart, so that they were exposed to much less severe competition than in the last case. The longest leaves on the three crossed plants, when turned out, exceeded those on the self-fertilised plants by a mere trifle, namely, on an average by .17 of an inch. When fully grown the three crossed plants produced twenty-six flower-stems; the two tallest of which on each plant were on an average 54.04 inches in height. The three self-fertilised plants produced twenty-three flower-stems, the two tallest of which on each plant had an average height of 46.18 inches. So that the difference between these two lots, which hardly competed together, is much less than in the last case when there was moderately severe competition, namely, as 100 to 85, instead of as 100 to 70.

THE EFFECTS ON THE OFFSPRING OF INTERCROSSING DIFFERENT FLOWERS ON THE SAME PLANT, INSTEAD OF CROSSING DISTINCT INDIVIDUALS.

A fine plant growing in my garden (one of the foregoing seedlings) was covered with a net, and six flowers were crossed with pollen from another flower on the same plant, and six others were fertilised with their own pollen. All produced good capsules. The seeds from each were placed in separate watch-glasses, and no difference could be perceived by the eye between the two lots of seeds; and when they were weighed there was no difference of any significance, as the seeds from the self-fertilised capsules weighed 7.65 grains, whilst those from the crossed capsules weighed 7.7 grains. Therefore the sterility of the present species, when insects are excluded, is not due to the impotence of pollen on the stigma of the same flower. Both lots of seeds and seedlings were treated in exactly the same manner as in Table 3/23, excepting that after the pairs of germinating seeds had been planted on the opposite sides of eight pots, all the remaining seeds were thickly sown on the opposite sides of Pots 9 and 10 in Table 3/24. The young plants during the following spring were turned out of their pots, without being disturbed, and planted in the open ground in two rows, not very close together, so that they were subjected to only moderately severe competition with one another. Very differently to what occurred in the first experiment, when the plants were subjected to somewhat severe mutual competition, an equal number on each side either died or did not produce flower-stems. The tallest flower-stems on the surviving plants were measured, as shown in Table 3/24.

TABLE 3/24. Digitalis purpurea.

The tallest Flower-stem on each Plant measured in inches: 0 signifies that the Plant died, or did not produce a Flower-stem.

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 49 4/8 : 45 5/8. Pot 1 : 46 7/8 : 52. Pot 1 : 43 6/8 : 0.

Pot 2 : 38 4/8 : 54 4/8. Pot 2 : 47 4/8 : 47 4/8. Pot 2 : 0 : 32 5/8.

Pot 3 : 54 7/8 : 46 5/8.

Pot 4 : 32 1/8 : 41 3/8. Pot 4 : 0 : 29 7/8. Pot 4 : 43 7/8 : 37 1/8.

Pot 5 : 46 6/8 : 42 1/8. Pot 5 : 40 4/8 : 42 1/8. Pot 5 : 43 : 0.

Pot 6 : 48 2/8 : 47 7/8. Pot 6 : 46 2/8 : 48 3/8.

Pot 7 : 48 5/8 : 25. Pot 7 : 42 : 40 5/8.

Pot 8 : 46 7/8 : 39 1/8.

Pot 9 : 49 : 30 3/8. Pot 9 : 50 3/8 : 15. Pot 9 : 46 3/8 : 36 7/8. Pot 9 : 47 6/8 : 44 1/8. Pot 9 : 0 : 31 6/8. Crowded Plants.

Pot 10 : 46 4/8 : 47 7/8. Pot 10 : 35 2/8 : 0. Pot 10 : 24 5/8 : 34 7/8. Pot 10 : 41 4/8 : 40 7/8. Pot 10 : 17 3/8 : 41 1/8. Crowded Plants.

Total : 1078.00 : 995.38.

The average height of the flower-stems on the twenty-five crossed plants in all the pots taken together is 43.12 inches, and that of the twenty-five self-fertilised plants 39.82, or as 100 to 92. In order to test this result, the plants planted in pairs in Pots 1 and 8 were considered by themselves, and the average height of the sixteen crossed plants is here 44.9, and that of the sixteen self-fertilised plants 42.03, or as 100 to 94. Again, the plants raised from the thickly sown seed in Pots 9 and 10, which were subjected to very severe mutual competition, were taken by themselves, and the average height of the nine crossed plants is 39.86, and that of the nine self-fertilised plants 35.88, or as 100 to 90. The plants in these two latter pots (9 and 10), after being measured, were cut down close to the ground and weighed: the nine crossed plants weighed 57.66 ounces, and the nine self-fertilised plants 45.25 ounces, or as 100 to 78. On the whole we may conclude, especially from the evidence of weight, that seedlings from a cross between flowers on the same plant have a decided, though not great, advantage over those from flowers fertilised with their own pollen, more especially in the case of the plants subjected to severe mutual competition. But the advantage is much less than that exhibited by the crossed offspring of distinct plants, for these exceeded the self-fertilised plants in height as 100 to 70, and in the number of flower-stems as 100 to 48. Digitalis thus differs from Ipomoea, and almost certainly from Mimulus, as with these two species a cross between flowers on the same plant did no good.

CALCEOLARIA.A BUSHY GREENHOUSE VARIETY, WITH YELLOW FLOWERS BLOTCHED WITH PURPLE.

The flowers in this genus are constructed so as to favour or almost ensure cross-fertilisation (3/6. Hildebrand as quoted by H. Muller ‘Die Befruchtung der Blumen’ 1873 page 277.); and Mr. Anderson remarks that extreme care is necessary to exclude insects in order to preserve any kind true. (3/7. ‘Gardeners’ Chronicle’ 1853 page 534.) He adds the interesting statement, that when the corolla is cut quite away, insects, as far as he has seen, never discover or visit the flowers. This plant is, however, self-fertile if insects are excluded. So few experiments were made by me, that they are hardly worth giving. Crossed and self-fertilised seeds were sown on opposite sides of a pot, and after a time the crossed seedlings slightly exceeded the self-fertilised in height. When a little further grown, the longest leaves on the former were very nearly 3 inches in length, whilst those on the self-fertilised plants were only 2 inches. Owing to an accident, and to the pot being too small, only one plant on each side grew up and flowered; the crossed plant was 19 1/2 inches in height, and the self-fertilised one 15 inches; or as 100 to 77.

Linaria vulgaris.

It has been mentioned in the introductory chapter that two large beds of this plant were raised by me many years ago from crossed and self-fertilised seeds, and that there was a conspicuous difference in height and general appearance between the two lots. The trial was afterwards repeated with more care; but as this was one of the first plants experimented on, my usual method was not followed. Seeds were taken from wild plants growing in this neighbourhood and sown in poor soil in my garden. Five plants were covered with a net, the others being left exposed to the bees, which incessantly visit the flowers of this species, and which, according to H. Muller, are the exclusive fertilisers. This excellent observer remarks that, as the stigma lies between the anthers and is mature at the same time with them, self-fertilisation is possible. (3/8. ‘Die Befruchtung’ etc. page 279.) But so few seeds are produced by protected plants, that the pollen and stigma of the same flower seem to have little power of mutual interaction. The exposed plants bore numerous capsules forming solid spikes. Five of these capsules were examined and appeared to contain an equal number of seeds; and these being counted in one capsule, were found to be 166. The five protected plants produced altogether only twenty-five capsules, of which five were much finer than all the others, and these contained an average of 23.6 seeds, with a maximum in one capsule of fifty-five. So that the number of seeds in the capsules on the exposed plants to the average number in the finest capsules on the protected plants was as 100 to 14.

Some of the spontaneously self-fertilised seeds from under the net, and some seeds from the uncovered plants naturally fertilised and almost certainly intercrossed by the bees, were sown separately in two large pots of the same size; so that the two lots of seedlings were not subjected to any mutual competition. Three of the crossed plants when in full flower were measured, but no care was taken to select the tallest plants; their heights were 7 4/8, 7 2/8, and 6 4/8 inches; averaging 7.08 in height. The three tallest of all the self-fertilised plants were then carefully selected, and their heights were 6 3/8, 5 5/8, and 5 2/8, averaging 5.75 in height. So that the naturally crossed plants were to the spontaneously self-fertilised plants in height, at least as much as 100 to 81.

Verbascum thapsus.

The flowers of this plant are frequented by various insects, chiefly by bees, for the sake of the pollen. Hermann Muller, however, has shown (‘Die Befruchtung’ etc. page 277) that V. nigrum secretes minute drops of nectar. The arrangement of the reproductive organs, though not at all complex, favours cross-fertilisation; and even distinct species are often crossed, for a greater number of naturally produced hybrids have been observed in this genus than in almost any other. (3/9. I have given a striking case of a large number of such hybrids between Verbascum thapsus and lychnitis found growing wild: ‘Journal of Linnean Society Botany’ volume 10 page 451.) Nevertheless the present species is perfectly self-fertile, if insects are excluded; for a plant protected by a net was as thickly loaded with fine capsules as the surrounding uncovered plants. Verbascum lychnitis is rather less self-fertile, for some protected plants did not yield quite so many capsules as the adjoining uncovered plants.

Plants of Verbascum thapsus had been raised for a distinct purpose from self-fertilised seeds; and some flowers on these plants were again self-fertilised, yielding seed of the second self-fertilised generation; and other flowers were crossed with pollen from a distinct plant. The seeds thus produced were sown on the opposite sides of four large pots. They germinated, however, so irregularly (the crossed seedlings generally coming up first) that I was able to save only six pairs of equal age. These when in full flower were measured, as in Table 3/25.

TABLE 3/25. Verbascum thapsus.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants of the Second Generation.

Pot 1 : 76 : 53 4/8.

Pot 2 : 54 : 66.

Pot 3 : 62 : 75. Pot 3 : 60 5/8 : 30 4/8.

Pot 4 : 73 : 62. Pot 4 : 66 4/8 : 52.

Total : 392.13 : 339.00.

We here see that two of the self-fertilised plants exceed in height their crossed opponents. Nevertheless the average height of the six crossed plants is 65.34 inches, and that of the six self-fertilised plants 56.5 inches; or as 100 to 86.

Vandellia nummularifolia.

Seeds were sent to me by Mr. J. Scott from Calcutta of this small Indian weed, which bears perfect and cleistogene flowers. (3/10. The convenient term of CLEISTOGENE was proposed by Kuhn in an article on the present genus in ‘Bot. Zeitung’ 1867 page 65.) The latter are extremely small, imperfectly developed, and never expand, yet yield plenty of seeds. The perfect and open flowers are also small, of a white colour with purple marks; they generally produce seed, although the contrary has been asserted; and they do so even if protected from insects. They have a rather complicated structure, and appear to be adapted for cross-fertilisation, but were not carefully examined by me. They are not easy to fertilise artificially, and it is possible that some of the flowers which I thought that I had succeeded in crossing were afterwards spontaneously self-fertilised under the net. Sixteen capsules from the crossed perfect flowers contained on an average ninety-three seeds (with a maximum in one capsule of 137), and thirteen capsules from the self-fertilised perfect flowers contained sixty-two seeds (with a maximum in one capsule of 135); or as 100 to 67. But I suspect that this considerable excess was accidental, as on one occasion nine crossed capsules were compared with seven self-fertilised capsules (both included in the above number), and they contained almost exactly the same average number of seed. I may add that fifteen capsules from self-fertilised cleistogene flowers contained on an average sixty-four seeds, with a maximum in one of eighty-seven.

Crossed and self-fertilised seeds from the perfect flowers, and other seeds from the self-fertilised cleistogene flowers, were sown in five pots, each divided superficially into three compartments. The seedlings were thinned at an early age, so that twenty plants were left in each of the three divisions. The crossed plants when in full flower averaged 4.3 inches, and the self-fertilised plants from the perfect flowers 4.27 inches in height; or as 100 to 99. The self-fertilised plants from the cleistogene flowers averaged 4.06 inches in height; so that the crossed were in height to these latter plants as 100 to 94.

I determined to compare again the growth of plants raised from crossed and self-fertilised perfect flowers, and obtained two fresh lots of seeds. These were sown on opposite sides of five pots, but they were not sufficiently thinned, so that they grew rather crowded. When fully grown, all those above 2 inches in height were selected, all below this standard being rejected; the former consisted of forty-seven crossed and forty-one self-fertilised plants; thus a greater number of the crossed than of the self-fertilised plants grew to a height of above 2 inches. Of the crossed plants, the twenty-four tallest were on an average 3.6 inches in height; whilst the twenty-four tallest self-fertilised plants were 3.38 inches in average height; or as 100 to 94. All these plants were then cut down close to the ground, and the forty-seven crossed plants weighed 1090.3 grains, and the forty-one self-fertilised plants weighed 887.4 grains. Therefore an equal number of crossed and self-fertilised would have been to each other in weight as 100 to 97. From these several facts we may conclude that the crossed plants had some real, though very slight, advantage in height and weight over the self-fertilised plants, when grown in competition with one another.

The crossed plants were, however, inferior in fertility to the self-fertilised. Six of the finest plants were selected out of the forty-seven crossed plants, and six out of the forty-one self-fertilised plants; and the former produced 598 capsules, whilst the latter or self-fertilised plants produced 752 capsules. All these capsules were the product of cleistogene flowers, for the plants did not bear during the whole of this season any perfect flowers. The seeds were counted in ten cleistogene capsules produced by crossed plants, and their average number was 46.4 per capsule; whilst the number in ten cleistogene capsules produced by the self-fertilised plants was 49.4; or as 100 to 106.

3. GESNERIACEAE.—Gesneria pendulina.

In Gesneria the several parts of the flower are arranged on nearly the same plan as in Digitalis, and most or all of the species are dichogamous. (3/11. Dr. Ogle ‘Popular Science Review’ January 1870 page 51.) Plants were raised from seed sent me by Fritz Muller from South Brazil. Seven flowers were crossed with pollen from a distinct plant, and produced seven capsules containing by weight 3.01 grains of seeds. Seven flowers on the same plants were fertilised with their own pollen, and their seven capsules contained exactly the same weight of seeds. Germinating seeds were planted on opposite sides of four pots, and when fully grown measured to the tips of their leaves.

TABLE 3/26. Gesneria pendulina.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 42 2/8 : 39. Pot 1 : 24 4/8 : 27 3/8.

Pot 2 : 33 : 30 6/8. Pot 2 : 27 : 19 2/8.

Pot 3 : 33 4/8 : 31 7/8. Pot 3 : 29 4/8 : 28 6/8.

Pot 4 : 30 6/8 : 29 6/8. Pot 4 : 36 : 26 3/8.

Total : 256.50 : 233.13.

The average height of the eight crossed plants is 32.06 inches, and that of the eight self-fertilised plants 29.14; or as 100 to 90.

4. LABIATAE.—Salvia coccinea. (3/12. The admirable mechanical adaptations in this genus for favouring or ensuring cross-fertilisation, have been fully described by Sprengel, Hildebrand, Delpino, H. Muller, Ogle, and others, in their several works.)

This species, unlike most of the others in the same genus, yields a good many seeds when insects are excluded. I gathered ninety-eight capsules produced by flowers spontaneously self-fertilised under a net, and they contained on an average 1.45 seeds, whilst flowers artificially fertilised with their own pollen, in which case the stigma will have received plenty of pollen, yielded on an average 3.3 seeds, or more than twice as many. Twenty flowers were crossed with pollen from a distinct plant, and twenty-six were self-fertilised. There was no great difference in the proportional number of flowers which produced capsules by these two processes, or in the number of the contained seeds, or in the weight of an equal number of seeds.

Seeds of both kinds were sown rather thickly on opposite sides of three pots. When the seedlings were about 3 inches in height, the crossed showed a slight advantage over the self-fertilised. When two-thirds grown, the two tallest plants on each side of each pot were measured; the crossed averaged 16.37 inches, and the self-fertilised 11.75 in height; or as 100 to 71. When the plants were fully grown and had done flowering, the two tallest plants on each side were again measured, with the results shown in Table 3/27.

TABLE 3/27. Salvia coccinea.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 32 6/8 : 25. Pot 1 : 20 : 18 6/8.

Pot 2 : 32 3/8 : 20 6/8. Pot 2 : 24 4/8 : 19 4/8.

Pot 3 : 29 4/8 : 25. Pot 3 : 28 : 18.

Total : 167.13 : 127.00.

It may be here seen that each of the six tallest crossed plants exceeds in height its self-fertilised opponent; the former averaged 27.85 inches, whilst the six tallest self-fertilised plants averaged 21.16 inches; or as 100 to 76. In all three pots the first plant which flowered was a crossed one. All the crossed plants together produced 409 flowers, whilst all the self-fertilised together produced only 232 flowers; or as 100 to 57. So that the crossed plants in this respect were far more productive than the self-fertilised.

Origanum vulgare.

This plant exists, according to H. Muller, under two forms; one hermaphrodite and strongly proterandrous, so that it is almost certain to be fertilised by pollen from another flower; the other form is exclusively female, has a smaller corolla, and must of course be fertilised by pollen from a distinct plant in order to yield any seeds. The plants on which I experimented were hermaphrodites; they had been cultivated for a long period as a pot-herb in my kitchen garden, and were, like so many long-cultivated plants, extremely sterile. As I felt doubtful about the specific name I sent specimens to Kew, and was assured that the species was Origanum vulgare. My plants formed one great clump, and had evidently spread from a single root by stolons. In a strict sense, therefore, they all belonged to the same individual. My object in experimenting on them was, firstly, to ascertain whether crossing flowers borne by plants having distinct roots, but all derived asexually from the same individual, would be in any respect more advantageous than self-fertilisation; and, secondly, to raise for future trial seedlings which would constitute really distinct individuals. Several plants in the above clump were covered by a net, and about two dozen seeds (many of which, however, were small and withered) were obtained from the flowers thus spontaneously self-fertilised. The remainder of the plants were left uncovered and were incessantly visited by bees, so that they were doubtless crossed by them. These exposed plants yielded rather more and finer seed (but still very few) than did the covered plants. The two lots of seeds thus obtained were sown on opposite sides of two pots; the seedlings were carefully observed from their first growth to maturity, but they did not differ at any period in height or in vigour, the importance of which latter observation we shall presently see. When fully grown, the tallest crossed plant in one pot was a very little taller than the tallest self-fertilised plant on the opposite side, and in the other pot exactly the reverse occurred. So that the two lots were in fact equal; and a cross of this kind did no more good than crossing two flowers on the same plant of Ipomoea or Mimulus.

The plants were turned out of the two pots without being disturbed and planted in the open ground, in order that they might grow more vigorously. In the following summer all the self-fertilised and some of the quasi-crossed plants were covered by a net. Many flowers on the latter were crossed by me with pollen from a distinct plant, and others were left to be crossed by the bees. These quasi-crossed plants produced rather more seed than did the original ones in the great clump when left to the action of the bees. Many flowers on the self-fertilised plants were artificially self-fertilised, and others were allowed to fertilise themselves spontaneously under the net, but they yielded altogether very few seeds. These two lots of seeds—the product of a cross between distinct seedlings, instead of as in the last case between plants multiplied by stolons, and the product of self-fertilised flowers—were allowed to germinate on bare sand, and several equal pairs were planted on opposite sides of two LARGE pots. At a very early age the crossed plants showed some superiority over the self-fertilised, which was ever afterwards retained. When the plants were fully grown, the two tallest crossed and the two tallest self-fertilised plants in each pot were measured, as shown in Table 3/28. I regret that from want of time I did not measure all the pairs; but the tallest on each side seemed fairly to represent the average difference between the two lots.

TABLE 3/28. Origanum vulgare.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants (two tallest in each pot).

Column 3: Self-fertilised Plants (two tallest in each pot).

Pot 1 : 26 : 24. Pot 1 : 21 : 21.

Pot 2 : 17 : 12. Pot 2 : 16 : 11 4/8.

Total : 80.0 : 68.5.

The average height of the crossed plants is here 20 inches, and that of the self-fertilised 17.12; or as 100 to 86. But this excess of height by no means gives a fair idea of the vast superiority in vigour of the crossed over the self-fertilised plants. The crossed flowered first and produced thirty flower-stems, whilst the self-fertilised produced only fifteen, or half the number. The pots were then bedded out, and the roots probably came out of the holes at the bottom and thus aided their growth. Early in the following summer the superiority of the crossed plants, owing to their increase by stolons, over the self-fertilised plants was truly wonderful. In Pot 1, and it should be remembered that very large pots had been used, the oval clump of crossed plants was 10 by 4 1/2 inches across, with the tallest stem, as yet young, 5 1/2 inches in height; whilst the clump of self-fertilised plants, on the opposite side of the same pot, was only 3 1/2 by 2 1/2 inches across, with the tallest young stem 4 inches in height. In Pot 2, the clump of crossed plants was 18 by 9 inches across, with the tallest young stem 8 1/2 inches in height; whilst the clump of self-fertilised plants on the opposite side of the same pot was 12 by 4 1/2 inches across, with the tallest young stem 6 inches in height. The crossed plants during this season, as during the last, flowered first. Both the crossed and self-fertilised plants being left freely exposed to the visits of bees, manifestly produced much more seed than their grand-parents,—the plants of the original clump still growing close by in the same garden, and equally left to the action of the bees.

5. ACANTHACEAE.—Thunbergia alata.

It appears from Hildebrand’s description (‘Botanische Zeitung’ 1867 page 285) that the conspicuous flowers of this plant are adapted for cross-fertilisation. Seedlings were twice raised from purchased seed; but during the early summer, when first experimented on, they were extremely sterile, many of the anthers containing hardly any pollen. Nevertheless, during the autumn these same plants spontaneously produced a good many seeds. Twenty-six flowers during the two years were crossed with pollen from a distinct plant, but they yielded only eleven capsules; and these contained very few seeds! Twenty-eight flowers were fertilised with pollen from the same flower, and these yielded only ten capsules, which, however, contained rather more seed than the crossed capsules. Eight pairs of germinating seeds were planted on opposite sides of five pots; and exactly half the crossed and half the self-fertilised plants exceeded their opponents in height. Two of the self-fertilised plants died young, before they were measured, and their crossed opponents were thrown away. The six remaining pairs of these grew very unequally, some, both of the crossed and self-fertilised plants, being more than twice as tall as the others. The average height of the crossed plants was 60 inches, and that of the self-fertilised plants 65 inches, or as 100 to 108. A cross, therefore, between distinct individuals here appears to do no good; but this result deduced from so few plants in a very sterile condition and growing very unequally, obviously cannot be trusted.]

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This book is part of the public domain. Charles Darwin (2003). The Effects of Cross & Self-Fertilisation in the Vegetable Kingdom. Urbana, Illinois: Project Gutenberg. Retrieved October 2022, https://www.gutenberg.org/cache/epub/4346/pg4346-images.html

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