Mr. Galton sent me at the same time graphical representations which he had made of the measurements, and they evidently form fairly regular curves. He appends the words “very good” to those of Zea and Limnanthes. He also calculated the average height of the crossed and self-fertilised plants in the seven tables by a more correct method than that followed by me, namely, by including the heights, as estimated in accordance with statistical rules, of a few plants which died before they were measured; whereas I merely added up the heights of the survivors, and divided the sum by their number. The difference in our results is in one way highly satisfactory, for the average heights of the self-fertilised plants, as deduced by Mr. Galton, is less than mine in all the cases excepting one, in which our averages are the same; and this shows that I have by no means exaggerated the superiority of the crossed over the self-fertilised plants.
After the heights of the crossed and self-fertilised plants had been taken, they were sometimes cut down close to the ground, and an equal number of both weighed. This method of comparison gives very striking results, and I wish that it had been oftener followed. Finally a record was often kept of any marked difference in the rate of germination of the crossed and self-fertilised seeds,—of the relative periods of flowering of the plants raised from them,—and of their productiveness, that is, of the number of seed-capsules which they produced and of the average number of seeds which each capsule contained.
When I began my experiments I did not intend to raise crossed and self-fertilised plants for more than a single generation; but as soon as the plants of the first generation were in flower I thought that I would raise one more generation, and acted in the following manner. Several flowers on one or more of the self-fertilised plants were again self-fertilised; and several flowers on one or more of the crossed plants were fertilised with pollen from another crossed plant of the same lot. Having thus once begun, the same method was followed for as many as ten successive generations with some of the species. The seeds and seedlings were always treated in exactly the same manner as already described. The self-fertilised plants, whether originally descended from one or two mother-plants, were thus in each generation as closely interbred as was possible; and I could not have improved on my plan. But instead of crossing one of the crossed plants with another crossed plant, I ought to have crossed the self-fertilised plants of each generation with pollen taken from a non-related plant—that is, one belonging to a distinct family or stock of the same species and variety. This was done in several cases as an additional experiment, and gave very striking results. But the plan usually followed was to put into competition and compare intercrossed plants, which were almost always the offspring of more or less closely related plants, with the self-fertilised plants of each succeeding generation;—all having been grown under closely similar conditions. I have, however, learnt more by this method of proceeding, which was begun by an oversight and then necessarily followed, than if I had always crossed the self-fertilised plants of each succeeding generation with pollen from a fresh stock.
I have said that the crossed plants of the successive generations were almost always inter-related. When the flowers on an hermaphrodite plant are crossed with pollen taken from a distinct plant, the seedlings thus raised may be considered as hermaphrodite brothers or sisters; those raised from the same capsule being as close as twins or animals of the same litter. But in one sense the flowers on the same plant are distinct individuals, and as several flowers on the mother-plant were crossed by pollen taken from several flowers on the father-plant, such seedlings would be in one sense half-brothers or sisters, but more closely related than are the half-brothers and sisters of ordinary animals. The flowers on the mother-plant were, however, commonly crossed by pollen taken from two or more distinct plants; and in these cases the seedlings might be called with more truth half-brothers or sisters. When two or three mother-plants were crossed, as often happened, by pollen taken from two or three father-plants (the seeds being all intermingled), some of the seedlings of the first generation would be in no way related, whilst many others would be whole or half-brothers and sisters. In the second generation a large number of the seedlings would be what may be called whole or half first-cousins, mingled with whole and half-brothers and sisters, and with some plants not at all related. So it would be in the succeeding generations, but there would also be many cousins of the second and more remote degrees. The relationship will thus have become more and more inextricably complex in the later generations; with most of the plants in some degree and many of them closely related.
I have only one other point to notice, but this is one of the highest importance; namely, that the crossed and self-fertilised plants were subjected in the same generation to as nearly similar and uniform conditions as was possible. In the successive generations they were exposed to slightly different conditions as the seasons varied, and they were raised at different periods. But in other respects all were treated alike, being grown in pots in the same artificially prepared soil, being watered at the same time, and kept close together in the same greenhouse or hothouse. They were therefore not exposed during successive years to such great vicissitudes of climate as are plants growing out of doors.
ON SOME APPARENT AND REAL CAUSES OF ERROR IN MY EXPERIMENTS.
It has been objected to such experiments as mine, that covering plants with a net, although only for a short time whilst in flower, may affect their health and fertility. I have seen no such effect except in one instance with a Myosotis, and the covering may not then have been the real cause of injury. But even if the net were slightly injurious, and certainly it was not so in any high degree, as I could judge by the appearance of the plants and by comparing their fertility with that of neighbouring uncovered plants, it would not have vitiated my experiments; for in all the more important cases the flowers were crossed as well as self-fertilised under a net, so that they were treated in this respect exactly alike.
As it is impossible to exclude such minute pollen-carrying insects as Thrips, flowers which it was intended to fertilise with their own pollen may sometimes have been afterwards crossed with pollen brought by these insects from another flower on the same plant; but as we shall hereafter see, a cross of this kind does not produce any effect, or at most only a slight one. When two or more plants were placed near one another under the same net, as was often done, there is some real though not great danger of the flowers which were believed to be self-fertilised being afterwards crossed with pollen brought by Thrips from a distinct plant. I have said that the danger is not great because I have often found that plants which are self-sterile, unless aided by insects, remained sterile when several plants of the same species were placed under the same net. If, however, the flowers which had been presumably self-fertilised by me were in any case afterwards crossed by Thrips with pollen brought from a distinct plant, crossed seedlings would have been included amongst the self-fertilised; but it should be especially observed that this occurrence would tend to diminish and not to increase any superiority in average height, fertility, etc., of the crossed over the self-fertilised plants.
As the flowers which were crossed were never castrated, it is probable or even almost certain that I sometimes failed to cross-fertilise them effectually, and that they were afterwards spontaneously self-fertilised. This would have been most likely to occur with dichogamous species, for without much care it is not easy to perceive whether their stigmas are ready to be fertilised when the anthers open. But in all cases, as the flowers were protected from wind, rain, and the access of insects, any pollen placed by me on the stigmatic surface whilst it was immature, would generally have remained there until the stigma was mature; and the flowers would then have been crossed as was intended. Nevertheless, it is highly probable that self-fertilised seedlings have sometimes by this means got included amongst the crossed seedlings. The effect would be, as in the former case, not to exaggerate but to diminish any average superiority of the crossed over the self-fertilised plants.
Errors arising from the two causes just named, and from others,—such as some of the seeds not having been thoroughly ripened, though care was taken to avoid this error—the sickness or unperceived injury of any of the plants,—will have been to a large extent eliminated, in those cases in which many crossed and self-fertilised plants were measured and an average struck. Some of these causes of error will also have been eliminated by the seeds having been allowed to germinate on bare damp sand, and being planted in pairs; for it is not likely that ill-matured and well-matured, or diseased and healthy seeds, would germinate at exactly the same time. The same result will have been gained in the several cases in which only a few of the tallest, finest, and healthiest plants on each side of the pots were measured.