THE EFFECTS OF A CROSS WITH A FRESH STOCK.
I now tried a different experiment. Eight flowers on the self-fertilised plants of the last experiment (i.e., grandchildren of the plants which grew in Brazil) were again fertilised with pollen from the same plant, and produced five capsules, containing on an average 27.4 seeds, with a maximum in one of forty-two seeds. The seedlings raised from these seeds formed the second SELF-FERTILISED generation of the Brazilian stock.
Eight flowers on one of the crossed plants of the last experiment were crossed with pollen from another grandchild, and produced five capsules. These contained on an average 31.6 seeds, with a maximum in one of forty-nine seeds. The seedlings raised from these seeds may be called the INTERCROSSED.
Lastly, eight other flowers on the crossed plants of the last experiment were fertilised with pollen from a plant of the English stock, growing in my garden, and which must have been exposed during many previous generations to very different conditions from those to which the Brazilian progenitors of the mother-plant had been subjected. These eight flowers produced only four capsules, containing on an average 63.2 seeds, with a maximum in one of ninety. The plants raised from these seeds may be called the ENGLISH-CROSSED. As far as the above averages can be trusted from so few capsules, the English-crossed capsules contained twice as many seeds as the intercrossed, and rather more than twice as many as the self-fertilised capsules. The plants which yielded these capsules were grown in pots in the greenhouse, so that their absolute productiveness must not be compared with that of plants growing out of doors.
The above three lots of seeds, namely, the self-fertilised, intercrossed, and English-crossed, were planted in an equal state of germination (having been as usual sown on bare sand) in nine large pots, each divided into three parts by superficial partitions. Many of the self-fertilised seeds germinated before those of the two crossed lots, and these were of course rejected. The seedlings thus raised are the great-grandchildren of the plants which grew in Brazil. When they were from 2 to 4 inches in height, the three lots were equal. They were measured when four-fifths grown, and again when fully grown, and as their relative heights were almost exactly the same at these two ages, I will give only the last measurements. The average height of the nineteen English-crossed plants was 45.92 inches; that of the eighteen intercrossed plants (for one died), 43.38; and that of the nineteen self-fertilised plants, 50.3 inches. So that we have the following ratios in height:—
The English-crossed to the self-fertilised plants, as 100 to 109.
The English-crossed to the intercrossed plants, as 100 to 94.
The intercrossed to the self-fertilised plants, as 100 to 116.
After the seed-capsules had been gathered, all these plants were cut down close to the ground and weighed. The nineteen English crossed plants weighed 18.25 ounces; the intercrossed plants (with their weight calculated as if there had been nineteen) weighed 18.2 ounces; and the nineteen self-fertilised plants, 21.5 ounces. We have therefore for the weights of the three lots of plants the following ratios:—
The English-crossed to the self-fertilised plants, as 100 to 118.
The English-crossed to the intercrossed plants, as 100 to 100.
The intercrossed to the self-fertilised plants, as 100 to 118.
We thus see that in weight, as in height, the self-fertilised plants had a decided advantage over the English-crossed and intercrossed plants.
The remaining seeds of the three kinds, whether or not in a state of germination, were sown in three long parallel rows in the open ground; and here again the self-fertilised seedlings exceeded in height by between 2 and 3 inches the seedlings in the two other rows, which were of nearly equal heights. The three rows were left unprotected throughout the winter, and all the plants were killed, with the exception of two of the self-fertilised; so that as far as this little bit of evidence goes, some of the self-fertilised plants were more hardy than any of the crossed plants of either lot.
We thus see that the self-fertilised plants which were grown in the nine pots were superior in height (as 116 to 100), and in weight (as 118 to 100), and apparently in hardiness, to the intercrossed plants derived from a cross between the grandchildren of the Brazilian stock. The superiority is here much more strongly marked than in the second trial with the plants of the English stock, in which the self-fertilised were to the crossed in height as 101 to 100. It is a far more remarkable fact—if we bear in mind the effects of crossing plants with pollen from a fresh stock in the cases of Ipomoea, Mimulus, Brassica, and Iberis—that the self-fertilised plants exceeded in height (as 109 to 100), and in weight (as 118 to 100), the offspring of the Brazilian stock crossed by the English stock; the two stocks having been long subjected to widely different conditions.
If we now turn to the fertility of the three lots of plants we find a very different result. I may premise that in five out of the nine pots the first plant which flowered was one of the English-crossed; in four of the pots it was a self-fertilised plant; and in not one did an intercrossed plant flower first; so that these latter plants were beaten in this respect, as in so many other ways. The three closely adjoining rows of plants growing in the open ground flowered profusely, and the flowers were incessantly visited by bees, and certainly thus intercrossed. The manner in which several plants in the previous experiments continued to be almost sterile as long as they were covered by a net, but set a multitude of capsules immediately that they were uncovered, proves how effectually the bees carry pollen from plant to plant. My gardener gathered, at three successive times, an equal number of ripe capsules from the plants of the three lots, until he had collected forty-five from each lot. It is not possible to judge from external appearance whether or not a capsule contains any good seeds; so that I opened all the capsules. Of the forty-five from the English-crossed plants, four were empty; of those from the intercrossed, five were empty; and of those from the self-fertilised, nine were empty. The seeds were counted in twenty-one capsules taken by chance out of each lot, and the average number of seeds in the capsules from the English-crossed plants was 67; from the intercrossed, 56; and from the self-fertilised, 48.52. It therefore follows that:—
The forty-five capsules (the four empty ones included) from the English-crossed plants contained 2747 seeds.
The forty-five capsules (the five empty ones included) from the intercrossed plants contained 2240 seeds.
The forty-five capsules (the nine empty ones included) from the self-fertilised plants contained 1746.7 seeds.
The reader should remember that these capsules are the product of cross-fertilisation, effected by the bees; and that the difference in the number of the contained seeds must depend on the constitution of the plants;—that is, on whether they were derived from a cross with a distinct stock, or from a cross between plants of the same stock, or from self-fertilisation. From the above facts we obtain the following ratios:—
Number of seeds contained in an equal number of naturally fertilised capsules produced:—
By the English-crossed and self-fertilised plants, as 100 to 63.
By the English-crossed and intercrossed plants, as 100 to 81.
By the intercrossed and self-fertilised plants, as 100 to 78.
But to have ascertained the productiveness of the three lots of plants, it would have been necessary to know how many capsules were produced by the same number of plants. The three long rows, however, were not of quite equal lengths, and the plants were much crowded, so that it would have been extremely difficult to have ascertained how many capsules were produced by them, even if I had been willing to undertake so laborious a task as to collect and count all the capsules. But this was feasible with the plants grown in pots in the greenhouse; and although these were much less fertile than those growing out of doors, their relative fertility appeared, after carefully observing them, to be the same. The nineteen plants of the English-crossed stock in the pots produced altogether 240 capsules; the intercrossed plants (calculated as nineteen) produced 137.22 capsules; and the nineteen self-fertilised plants, 152 capsules. Now, knowing the number of seeds contained in forty-five capsules of each lot, it is easy to calculate the relative numbers of seeds produced by an equal number of the plants of the three lots.
Number of seeds produced by an equal number of naturally-fertilised plants:—
Plants of English-crossed and self-fertilised parentage, as 100 to 40 seeds.
Plants of English-crossed and intercrossed parentage, as 100 to 45 seeds.
Plants of intercrossed and self-fertilised parentage, as 100 to 89 seeds.
The superiority in productiveness of the intercrossed plants (that is, the product of a cross between the grandchildren of the plants which grew in Brazil) over the self-fertilised, small as it is, is wholly due to the larger average number of seeds contained in the capsules; for the intercrossed plants produced fewer capsules in the greenhouse than did the self-fertilised plants. The great superiority in productiveness of the English-crossed over the self-fertilised plants is shown by the larger number of capsules produced, the larger average number of contained seeds, and the smaller number of empty capsules. As the English-crossed and intercrossed plants were the offspring of crosses in every previous generation (as must have been the case from the flowers being sterile with their own pollen), we may conclude that the great superiority in productiveness of the English-crossed over the intercrossed plants is due to the two parents of the former having been long subjected to different conditions.
The English-crossed plants, though so superior in productiveness, were, as we have seen, decidedly inferior in height and weight to the self-fertilised, and only equal to, or hardly superior to, the intercrossed plants. Therefore, the whole advantage of a cross with a distinct stock is here confined to productiveness, and I have met with no similar case.
8. RESEDACEAE.—Reseda lutea.
Seeds collected from wild plants growing in this neighbourhood were sown in the kitchen-garden; and several of the seedlings thus raised were covered with a net. Of these, some were found (as will hereafter be more fully described) to be absolutely sterile when left to fertilise themselves spontaneously, although plenty of pollen fell on their stigmas; and they were equally sterile when artificially and repeatedly fertilised with their own pollen; whilst other plants produced a few spontaneously self-fertilised capsules. The remaining plants were left uncovered, and as pollen was carried from plant to plant by the hive and humble-bees which incessantly visit the flowers, they produced an abundance of capsules. Of the necessity of pollen being carried from one plant to another, I had ample evidence in the case of this species and of R. odorata; for those plants, which set no seeds or very few as long as they were protected from insects, became loaded with capsules immediately that they were uncovered.
Seeds from the flowers spontaneously self-fertilised under the net, and from flowers naturally crossed by the bees, were sown on opposite sides of five large pots. The seedlings were thinned as soon as they appeared above ground, so that an equal number were left on the two sides. After a time the pots were plunged into the open ground. The same number of plants of crossed and self-fertilised parentage were measured up to the summits of their flower-stems, with the result given in Table 4/35. Those which did not produce flower-stems were not measured.
TABLE 4/35. Reseda lutea, in pots.
Heights of plants to the summits of the flower-stems measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 21 : 12 7/8. Pot 1 : 14 2/8 : 16. Pot 1 : 19 1/8 : 11 7/8. Pot 1 : 7 : 15 2/8. Pot 1 : 15 1/8 : 19 1/8.
Pot 2 : 20 4/8 : 12 4/8. Pot 2 : 17 3/8 : 16 2/8. Pot 2 : 23 7/8 : 16 2/8. Pot 2 : 17 1/8 : 13 3/8. Pot 2 : 20 6/8 : 13 5/8.
Pot 3 : 16 1/8 : 14 4/8. Pot 3 : 17 6/8 : 19 4/8. Pot 3 : 16 2/8 : 20 7/8. Pot 3 : 10 : 7 7/8. Pot 3 : 10 : 17 6/8.
Pot 4 : 22 1/8 : 9. Pot 4 : 19 : 11 4/8. Pot 4 : 18 7/8 : 11. Pot 4 : 16 4/8 : 16. Pot 4 : 19 2/8 : 16 3/8.
Pot 5 : 25 2/8 : 14 6/8. Pot 5 : 22 : 16. Pot 5 : 8 6/8 : 14 3/8. Pot 5 : 14 2/8 : 14 2/8.
Total : 412.25 : 350.86.
The average height of the twenty-four crossed plants is here 17.17 inches, and that of the same number of self-fertilised plants 14.61; or as 100 to 85. Of the crossed plants all but five flowered, whilst several of the self-fertilised did not do so. The above pairs, whilst still in flower, but with some capsules already formed, were afterwards cut down and weighed. The crossed weighed 90.5 ounces; and an equal number of the self-fertilised only 19 ounces, or as 100 to 21; and this is an astonishing difference.
Seeds of the same two lots were also sown in two adjoining rows in the open ground. There were twenty crossed plants in the one row and thirty-two self-fertilised plants in the other row, so that the experiment was not quite fair; but not so unfair as it at first appears, for the plants in the same row were not crowded so much as seriously to interfere with each other’s growth, and the ground was bare on the outside of both rows. These plants were better nourished than those in the pots and grew to a greater height. The eight tallest plants in each row were measured in the same manner as before, with the following result:—
TABLE 4/36. Reseda lutea, growing in the open ground.
Heights of plants to the summits of the flower-stems measured in inches.
Column 1: Crossed Plants.
Column 2: Self-fertilised Plants.
28 : 33 2/8.
27 3/8 : 23.
27 5/8 : 21 5/8.
28 6/8 : 20 4/8.
29 7/8 : 21 5/8.
26 6/8 : 22.
26 2/8 : 21 2/8.
30 1/8 : 21 7/8.
Total : 224.75 : 185.13
The average height of the crossed plants, whilst in full flower, was here 28.09, and that of the self-fertilised 23.14 inches; or as 100 to 82. It is a singular fact that the tallest plant in the two rows, was one of the self-fertilised. The self-fertilised plants had smaller and paler green leaves than the crossed. All the plants in the two rows were afterwards cut down and weighed. The twenty crossed plants weighed 65 ounces, and twenty self-fertilised (by calculation from the actual weight of the thirty-two self-fertilised plants) weighed 26.25 ounces; or as 100 to 40. Therefore the crossed plants did not exceed in weight the self-fertilised plants in nearly so great a degree as those growing in the pots, owing probably to the latter having been subjected to more severe mutual competition. On the other hand, they exceeded the self-fertilised in height in a slightly greater degree.
Reseda odorata.
Plants of the common mignonette were raised from purchased seed, and several of them were placed under separate nets. Of these some became loaded with spontaneously self-fertilised capsules; others produced a few, and others not a single one. It must not be supposed that these latter plants produced no seed because their stigmas did not receive any pollen, for they were repeatedly fertilised with pollen from the same plant with no effect; but they were perfectly fertile with pollen from any other plant. Spontaneously self-fertilised seeds were saved from one of the highly self-fertile plants, and other seeds were collected from the plants growing outside the nets, which had been crossed by the bees. These seeds after germinating on sand were planted in pairs on the opposite sides of five pots. The plants were trained up sticks, and measured to the summits of their leafy stems—the flower-stems not being included. We here have the result:—
TABLE 4/37. Reseda odorata (seedlings from a highly self-fertile plant).
Heights of plants to the summits of the leafy stems, flower-stems not included, measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 20 7/8 : 22 4/8. Pot 1 : 34 7/8 : 28 5/8. Pot 1 : 26 6/8 : 23 2/8. Pot 1 : 32 6/8 : 30 4/8.
Pot 2 : 34 3/8 : 28 5/8. Pot 2 : 34 5/8 : 30 5/8. Pot 2 : 11 6/8 : 23. Pot 2 : 33 3/8 : 30 1/8.
Pot 3 : 17 7/8 : 4 4/8. Pot 3 : 27 : 25. Pot 3 : 30 1/8 : 26 3/8. Pot 3 : 30 2/8 : 25 1/8.
Pot 4 : 21 5/8 : 22 6/8. Pot 4 : 28 : 25 4/8. Pot 4 : 32 5/8 : 15 1/8. Pot 4 : 32 3/8 : 24 6/8.
Pot 5 : 21 : 11 6/8. Pot 5 : 25 2/8 : 19 7/8. Pot 5 : 26 6/8 : 10 4/8.
Total : 522.25 : 428.50.
The average height of the nineteen crossed plants is here 27.48, and that of the nineteen self-fertilised 22.55 inches; or as 100 to 82. All these plants were cut down in the early autumn and weighed: the crossed weighed 11.5 ounces, and the self-fertilised 7.75 ounces, or as 100 to 67. These two lots having been left freely exposed to the visits of insects, did not present any difference to the eye in the number of seed-capsules which they produced.
The remainder of the same two lots of seeds were sown in two adjoining rows in the open ground; so that the plants were exposed to only moderate competition. The eight tallest on each side were measured, as shown in Table 4/38.
TABLE 4/38. Reseda odorata, growing in the open ground.
Heights of plants measured in inches.
Column 1: Crossed Plants.
Column 2: Self-fertilised Plants.
24 4/8 : 26 5/8.
27 2/8 : 25 7/8.
24 : 25.
26 6/8 : 28 3/8.
25 : 29 7/8.
26 2/8 : 25 7/8.
27 2/8 : 26 7/8.
25 1/8 : 28 2/8.
Total : 206.13 : 216.75
The average height of the eight crossed plants is 25.76, and that of the eight self-fertilised 27.09; or as 100 to 105.
We here have the anomalous result of the self-fertilised plants being a little taller than the crossed; of which fact I can offer no explanation. It is of course possible, but not probable, that the labels may have been interchanged by accident.
Another experiment was now tried: all the self-fertilised capsules, though very few in number, were gathered from one of the semi-self-sterile plants under a net; and as several flowers on this same plant had been fertilised with pollen from a distinct individual, crossed seeds were thus obtained. I expected that the seedlings from this semi-self-sterile plant would have profited in a higher degree from a cross, than did the seedlings from the fully self-fertile plants. But my anticipation was quite wrong, for they profited in a less degree. An analogous result followed in the case of Eschscholtzia, in which the offspring of the plants of Brazilian parentage (which were partially self-sterile) did not profit more from a cross, than did the plants of the far more self-fertile English stock. The above two lots of crossed and self-fertilised seeds from the same plant of Reseda odorata, after germinating on sand, were planted on opposite sides of five pots, and measured as in the last case, with the result in Table 4/39.
TABLE 4/39. Reseda odorata (seedlings from a semi-self-sterile plant).
Heights of plants to the summits of the leafy stems, flower-stems not included, measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 33 4/8 : 31. Pot 1 : 30 6/8 : 28. Pot 1 : 29 6/8 : 13 2/8. Pot 1 : 20 : 32.
Pot 2 : 22 : 21 6/8. Pot 2 : 33 4/8 : 26 6/8. Pot 2 : 31 2/8 : 25 2/8. Pot 2 : 32 4/8 : 30 4/8.
Pot 3 : 30 1/8 : 17 2/8. Pot 3 : 32 1/8 : 29 6/8. Pot 3 : 31 4/8 : 24 6/8. Pot 3 : 32 2/8 : 34 2/8.
Pot 4 : 19 1/8 : 20 6/8. Pot 4 : 30 1/8 : 32 6/8. Pot 4 : 24 3/8 : 31 4/8. Pot 4 : 30 6/8 : 36 6/8.
Pot 5 : 34 6/8 : 24 5/8. Pot 5 : 37 1/8 : 34. Pot 5 : 31 2/8 : 22 2/8. Pot 5 : 33 : 37 1/8.
Total : 599.75 : 554.25.
The average height of the twenty crossed plants is here 29.98, and that of the twenty self-fertilised 27.71 inches; or as 100 to 92. These plants were then cut down and weighed; and the crossed in this case exceeded the self-fertilised in weight by a mere trifle, namely, in the ratio of 100 to 99. The two lots, left freely exposed to insects, seemed to be equally fertile.
The remainder of the seed was sown in two adjoining rows in the open ground; and the eight tallest plants in each row were measured, with the result in Table 4/40.
TABLE 4/40. Reseda odorata, (seedlings from a semi-self-sterile plant, planted in the open ground).
Heights of plants measured in inches.
Column 1: Crossed Plants.
Column 2: Self-fertilised Plants.
28 2/8 : 22 3/8.
22 4/8 : 24 3/8.
25 7/8 : 23 4/8.
25 3/8 : 21 4/8.
29 4/8 : 22 5/8.
27 1/8 : 27 3/8.
22 4/8 : 27 3/8.
26 2/8 : 19 2/8.
Total : 207.38 : 188.38.
The average height of the eight crossed plants is here 25.92, and that of the eight self-fertilised plants 23.54 inches; or as 100 to 90.
9. VIOLACEAE.—Viola tricolor.
Whilst the flowers of the common cultivated heartsease are young, the anthers shed their pollen into a little semi-cylindrical passage, formed by the basal portion of the lower petal, and surrounded by papillae. The pollen thus collected lies close beneath the stigma, but can seldom gain access into its cavity, except by the aid of insects, which pass their proboscides down this passage into the nectary. (4/5. The flowers of this plant have been fully described by Sprengel, Hildebrand, Delpino, and H. Muller. The latter author sums up all the previous observations in his ‘Befruchtung der Blumen’ and in ‘Nature’ November 20, 1873 page 44. See also Mr. A.W. Bennett in ‘Nature’ May 15, 1873 page 50 and some remarks by Mr. Kitchener ibid page 143. The facts which follow on the effects of covering up a plant of V. tricolor have been quoted by Sir J. Lubbock in his ‘British Wild Flowers’ etc. page 62.) Consequently when I covered up a large plant of a cultivated variety, it set only eighteen capsules, and most of these contained very few good seeds—several from only one to three; whereas an equally fine uncovered plant of the same variety, growing close by, produced 105 fine capsules. The few flowers which produce capsules when insects are excluded, are perhaps fertilised by the curling inwards of the petals as their wither, for by this means pollen-grains adhering to the papillae might be inserted into the cavity of the stigma. But it is more probable that their fertilisation is effected, as Mr. Bennett suggests, by Thrips and certain minute beetles which haunt the flowers, and which cannot be excluded by any net. Humble-bees are the usual fertilisers; but I have more than once seen flies (Rhingia rostrata) at work, with the under sides of their bodies, heads and legs dusted with pollen; and having marked the flowers which they visited, I found them after a few days fertilised. (4/6. I should add that this fly apparently did not suck the nectar, but was attracted by the papillae which surround the stigma. Hermann Muller also saw a small bee, an Andrena, which could not reach the nectar, repeatedly inserting its proboscis beneath the stigma, where the papillae are situated; so that these papillae must be in some way attractive to insects. A writer asserts ‘Zoologist’ volume 3-4 page 1225, that a moth (Plusia) frequently visits the flowers of the pansy. Hive-bees do not ordinarily visit them, but a case has been recorded ‘Gardeners’ Chronicle’ 1844 page 374, of these bees doing so. Hermann Muller has also seen the hive-bee at work, but only on the wild small-flowered form. He gives a list ‘Nature’ 1873 page 45, of all the insects which he has seen visiting both the large and small-flowered forms. From his account, I suspect that the flowers of plants in a state of nature are visited more frequently by insects than those of the cultivated varieties. He has seen several butterflies sucking the flowers of wild plants, and this I have never observed in gardens, though I have watched the flowers during many years.) It is curious for how long a time the flowers of the heartsease and of some other plants may be watched without an insect being seen to visit them. During the summer of 1841, I observed many times daily for more than a fortnight some large clumps of heartsease growing in my garden, before I saw a single humble-bee at work. During another summer I did the same, but at last saw some dark-coloured humble-bees visiting on three successive days almost every flower in several clumps; and almost all these flowers quickly withered and produced fine capsules. I presume that a certain state of the atmosphere is necessary for the secretion of nectar, and that as soon as this occurs the insects discover the fact by the odour emitted, and immediately frequent the flowers.
As the flowers require the aid of insects for their complete fertilisation, and as they are not visited by insects nearly so often as most other nectar-secreting flowers, we can understand the remarkable fact discovered by H. Muller and described by him in ‘Nature,’ namely, that this species exists under two forms. One of these bears conspicuous flowers, which, as we have seen, require the aid of insects, and are adapted to be cross-fertilised by them; whilst the other form has much smaller and less conspicuously coloured flowers, which are constructed on a slightly different plan, favouring self-fertilisation, and are thus adapted to ensure the propagation of the species. The self-fertile form, however, is occasionally visited, and may be crossed by insects, though this is rather doubtful.
In my first experiments on Viola tricolor I was unsuccessful in raising seedlings, and obtained only one full-grown crossed and self-fertilised plant. The former was 12 1/2 inches and the latter 8 inches in height. On the following year several flowers on a fresh plant were crossed with pollen from another plant, which was known to be a distinct seedling; and to this point it is important to attend. Several other flowers on the same plant were fertilised with their own pollen. The average number of seeds in the ten crossed capsules was 18.7, and in the twelve self-fertilised capsules 12.83; or as 100 to 69. These seeds, after germinating on bare sand, were planted in pairs on the opposite sides of five pots. They were first measured when about a third of their full size, and the crossed plants then averaged 3.87 inches, and the self-fertilised only 2.00 inches in height; or as 100 to 52. They were kept in the greenhouse, and did not grow vigorously. Whilst in flower they were again measured to the summits of their stems (see Table 4/41), with the following result:—
TABLE 4/41. Viola tricolor.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 8 2/8 : 0 2/8. Pot 1 : 7 4/8 : 2 4/8. Pot 1 : 5 : 1 2/8.
Pot 2 : 5 : 6. Pot 2 : 4 : 4. Pot 2 : 4 4/8 : 3 1/8.
Pot 3 : 9 4/8 : 3 1/8. Pot 3 : 3 3/8 : 1 7/8. Pot 3 : 8 4/8 : 0 5/8.
Pot 4 : 4 7/8 : 2 1/8. Pot 4 : 4 2/8 : 1 6/8. Pot 4 : 4 : 2 1/8.
Pot 5 : 6 : 3. Pot 5 : 3 3/8 : 1 4/8.
Total : 78.13 : 33.25.
The average height of the fourteen crossed plants is here 5.58 inches, and that of the fourteen self-fertilised 2.37; or as 100 to 42. In four out of the five pots, a crossed plant flowered before any one of the self-fertilised; as likewise occurred with the pair raised during the previous year. These plants without being disturbed were now turned out of their pots and planted in the open ground, so as to form five separate clumps. Early in the following summer (1869) they flowered profusely, and being visited by humble-bees set many capsules, which were carefully collected from all the plants on both sides. The crossed plants produced 167 capsules, and the self-fertilised only 17; or as 100 to 10. So that the crossed plants were more than twice the height of the self-fertilised, generally flowered first, and produced ten times as many naturally fertilised capsules.
By the early part of the summer of 1870 the crossed plants in all the five clumps had grown and spread so much more than the self-fertilised, that any comparison between them was superfluous. The crossed plants were covered with a sheet of bloom, whilst only a single self-fertilised plant, which was much finer than any of its brethren, flowered. The crossed and self-fertilised plants had now grown all matted together on the respective sides of the superficial partitions still separating them; and in the clump which included the finest self-fertilised plant, I estimated that the surface covered by the crossed plants was about nine times as large as that covered by the self-fertilised plants. The extraordinary superiority of the crossed over the self-fertilised plants in all five clumps, was no doubt due to the crossed plants at first having had a decided advantage over the self-fertilised, and then robbing them more and more of their food during the succeeding seasons. But we should remember that the same result would follow in a state of nature even to a greater degree; for my plants grew in ground kept clear of weeds, so that the self-fertilised had to compete only with the crossed plants; whereas the whole surface of the ground is naturally covered with various kinds of plants, all of which have to struggle together for existence.
The ensuing winter was very severe, and in the following spring (1871) the plants were again examined. All the self-fertilised were now dead, with the exception of a single branch on one plant, which bore on its summit a minute rosette of leaves about as large as a pea. On the other hand, all the crossed plants without exception were growing vigorously. So that the self-fertilised plants, besides their inferiority in other respects, were more tender.
Another experiment was now tried for the sake of ascertaining how far the superiority of the crossed plants, or to speak more correctly, the inferiority of the self-fertilised plants, would be transmitted to their offspring. The one crossed and one self-fertilised plant, which were first raised, had been turned out of their pot and planted in the open ground. Both produced an abundance of very fine capsules, from which fact we may safely conclude that they had been cross-fertilised by insects. Seeds from both, after germinating on sand, were planted in pairs on the opposite sides of three pots. The naturally crossed seedlings derived from the crossed plants flowered in all three pots before the naturally crossed seedlings derived from the self-fertilised plants. When both lots were in full flower, the two tallest plants on each side of each pot were measured, and the result is shown in Table 4/42.
TABLE 4/42. Viola tricolor: seedlings from crossed and self-fertilised plants, the parents of both sets having been left to be naturally fertilised.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Naturally Crossed Plants from artificially crossed Plants.
Column 3: Naturally Crossed Plants from Self-fertilised Plants.
Pot 1 : 12 1/8 : 9 6/8. Pot 1 : 11 6/8 : 8 3/8.
Pot 2 : 13 2/8 : 9 6/8. Pot 2 : 10 : 11 4/8.
Pot 3 : 14 4/8 : 11 1/8. Pot 3 : 13 6/8 : 11 3/8.
Total : 75.38 : 61.88.
The average height of the six tallest plants derived from the crossed plants is 12.56 inches; and that of the six tallest plants derived from the self-fertilised plants is 10.31 inches; or as 100 to 82. We here see a considerable difference in height between the two sets, though very far from equalling that in the previous trials between the offspring from crossed and self-fertilised flowers. This difference must be attributed to the latter set of plants having inherited a weak constitution from their parents, the offspring of self-fertilised flowers; notwithstanding that the parents themselves had been freely intercrossed with other plants by the aid of insects.
10. RANUNCULACEAE.—Adonis aestivalis.
The results of my experiments on this plant are hardly worth giving, as I remark in my notes made at the time, “seedlings, from some unknown cause, all miserably unhealthy.” Nor did they ever become healthy; yet I feel bound to give the present case, as it is opposed to the general results at which I have arrived. Fifteen flowers were crossed and all produced fruit, containing on an average 32.5 seeds; nineteen flowers were fertilised with their own pollen, and they likewise all yielded fruit, containing a rather larger average of 34.5 seeds; or as 100 to 106. Seedlings were raised from these seeds. In one of the pots all the self-fertilised plants died whilst quite young; in the two others, the measurements were as follows:
TABLE 4/43. Adonis aestivalis.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 14 : 13 4/8. Pot 1 : 13 4/8 : 13 4/8.
Pot 2 : 16 2/8 : 15 2/8. Pot 2 : 13 2/8 : 15.
Total : 57.00 : 57.25.
The average height of the four crossed plants is 14.25, and that of the four self-fertilised plants 14.31; or as 100 to 100.4; so that they were in fact of equal height. According to Professor H. Hoffman, this plant is proterandrous (4/7. ‘Zur Speciesfrage’ 1875 page 11.); nevertheless it yields plenty of seeds when protected from insects.
Delphinium consolida.
It has been said in the case of this plant, as of so many others, that the flowers are fertilised in the bud, and that distinct plants or varieties can never naturally intercross. (4/8. Decaisne ‘Comptes-Rendus’ July 1863 page 5.) But this is an error, as we may infer, firstly from the flowers being proterandrous,—the mature stamens bending up, one after the other, into the passage which leads to the nectary, and afterwards the mature pistils bending in the same direction; secondly, from the number of humble-bees which visit the flowers (4/9. Their structure is described by H. Muller ‘Befruchtung’ etc., page 122.); and thirdly, from the greater fertility of the flowers when crossed with pollen from a distinct plant than when spontaneously self-fertilised. In the year 1863 I enclosed a large branch in a net, and crossed five flowers with pollen from a distinct plant; these yielded capsules containing on an average 35.2 very fine seeds, with a maximum of forty-two in one capsule. Thirty-two other flowers on the same branch produced twenty-eight spontaneously self-fertilised capsules, containing on an average 17.2 seeds, with a maximum in one of thirty-six seeds. But six of these capsules were very poor, yielding only from one to five seeds; if these are excluded, the remaining twenty-two capsules give an average of 20.9 seeds, though many of these seeds were small. The fairest ratio, therefore, for the number of seeds produced by a cross and by spontaneous self-fertilisation is as 100 to 59. These seeds were not sown, as I had too many other experiments in progress.
In the summer of 1867, which was a very unfavourable one, I again crossed several flowers under a net with pollen from a distinct plant, and fertilised other flowers on the same plant with their own pollen. The former yielded a much larger proportion of capsules than the latter; and many of the seeds in the self-fertilised capsules, though numerous, were so poor that an equal number of seeds from the crossed and self-fertilised capsules were in weight as 100 to 45. The two lots were allowed to germinate on sand, and pairs were planted on the opposite sides of four pots. When nearly two-thirds grown they were measured, as shown in Table 4/44.
TABLE 4/44. Delphinium consolida.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 11 : 11.
Pot 2 : 19 : 16 2/8. Pot 2 : 16 2/8 : 11 4/8.
Pot 3 : 26 : 22.
Pot 4 : 9 4/8 : 8 2/8. Pot 4 : 8 : 6 4/8.
Total : 89.75 : 75.50.
The six crossed plants here average 14.95, and the six self-fertilised 12.50 inches in height; or as 100 to 84. When fully grown they were again measured, but from want of time only a single plant on each side was measured; so that I have thought it best to give the earlier measurements. At the later period the three tallest crossed plants still exceeded considerably in height the three tallest self-fertilised, but not in quite so great a degree as before. The pots were left uncovered in the greenhouse, but whether the flowers were intercrossed by bees or self-fertilised I do not know. The six crossed plants produced 282 mature and immature capsules, whilst the six self-fertilised plants produced only 159; or as 100 to 56. So that the crossed plants were very much more productive than the self-fertilised.
11. CARYOPHYLLACEAE.—Viscaria oculata.
Twelve flowers were crossed with pollen from another plant, and yielded ten capsules, containing by weight 5.77 grains of seeds. Eighteen flowers were fertilised with their own pollen and yielded twelve capsules, containing by weight 2.63 grains. Therefore the seeds from an equal number of crossed and self-fertilised flowers would have been in weight as 100 to 38. I had previously selected a medium-sized capsule from each lot, and counted the seeds in both; the crossed one contained 284, and the self-fertilised one 126 seeds; or as 100 to 44. These seeds were sown on opposite sides of three pots, and several seedlings raised; but only the tallest flower-stem of one plant on each side was measured. The three on the crossed side averaged 32.5 inches, and the three on the self-fertilised side 34 inches in height; or as 100 to 104. But this trial was on much too small a scale to be trusted; the plants also grew so unequally that one of the three flower-stems on the crossed plants was very nearly twice as tall as that on one of the others; and one of the three flower-stems on the self-fertilised plants exceeded in an equal degree one of the others.
In the following year the experiment was repeated on a larger scale: ten flowers were crossed on a new set of plants and yielded ten capsules containing by weight 6.54 grains of seed. Eighteen spontaneously self-fertilised capsules were gathered, of which two contained no seed; the other sixteen contained by weight 6.07 grains of seed. Therefore the weight of seed from an equal number of crossed and spontaneously self-fertilised flowers (instead of artificially fertilised as in the previous case) was as 100 to 58.
The seeds after germinating on sand were planted in pairs on the opposite sides of four pots, with all the remaining seeds sown crowded in the opposite sides of a fifth pot; in this latter pot only the tallest plant on each side was measured. Until the seedlings had grown about 5 inches in height no difference could be perceived in the two lots. Both lots flowered at nearly the same time. When they had almost done flowering, the tallest flower-stem on each plant was measured, as shown in Table 4/45.
TABLE 4/45. Viscaria oculata.
Tallest flower-stem on each plant measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Crossed Plants.
Column 3: Self-fertilised Plants.
Pot 1 : 19 : 32 3/8. Pot 1 : 33 : 38. Pot 1 : 41 : 38. Pot 1 : 41 : 28 7/8.
Pot 2 : 37 4/8 : 36. Pot 2 : 36 4/8 : 32 3/8. Pot 2 : 38 : 35 6/8.
Pot 3 : 44 4/8 : 36. Pot 3 : 39 4/8 : 20 7/8. Pot 3 : 39 : 30 5/8.
Pot 4 : 30 2/8 : 36. Pot 4 : 31 : 39. Pot 4 : 33 1/8 : 29. Pot 4 : 24 : 38 4/8.
Pot 5 : 30 2/8 : 32. Crowded.
Total : 517.63 : 503.36.
The fifteen crossed plants here average 34.5, and the fifteen self-fertilised 33.55 inches in height; or as 100 to 97. So that the excess of height of the crossed plants is quite insignificant. In productiveness, however, the difference was much more plainly marked. All the capsules were gathered from both lots of plants (except from the crowded and unproductive ones in Pot 5), and at the close of the season the few remaining flowers were added in. The fourteen crossed plants produced 381, whilst the fourteen self-fertilised plants produced only 293 capsules and flowers; or as 100 to 77.
Dianthus caryophyllus.
The common carnation is strongly proterandrous, and therefore depends to a large extent upon insects for fertilisation. I have seen only humble-bees visiting the flowers, but I dare say other insects likewise do so. It is notorious that if pure seed is desired, the greatest care is necessary to prevent the varieties which grow in the same garden from intercrossing. (4/10. ‘Gardeners’ Chronicle’ 1847 page 268.) The pollen is generally shed and lost before the two stigmas in the same flower diverge and are ready to be fertilised. I was therefore often forced to use for self-fertilisation pollen from the same plant instead of from the same flower. But on two occasions, when I attended to this point, I was not able to detect any marked difference in the number of seeds produced by these two forms of self-fertilisation.
Several single-flowered carnations were planted in good soil, and were all covered with a net. Eight flowers were crossed with pollen from a distinct plant and yielded six capsules, containing on an average 88.6 seeds, with a maximum in one of 112 seeds. Eight other flowers were self-fertilised in the manner above described, and yielded seven capsules containing on an average 82 seeds, with a maximum in one of 112 seeds. So that there was very little difference in the number of seeds produced by cross-fertilisation and self-fertilisation, namely, as 100 to 92. As these plants were covered by a net, they produced spontaneously only a few capsules containing any seeds, and these few may perhaps be attributed to the action of Thrips and other minute insects which haunt the flowers. A large majority of the spontaneously self-fertilised capsules produced by several plants contained no seeds, or only a single one. Excluding these latter capsules, I counted the seeds in eighteen of the finest ones, and these contained on an average 18 seeds. One of the plants was spontaneously self-fertile in a higher degree than any of the others. On another occasion a single covered-up plant produced spontaneously eighteen capsules, but only two of these contained any seed, namely 10 and 15.