NINTH GENERATION OF CROSSED AND SELF-FERTILISED PLANTS.
Fourteen crossed plants, spontaneously self-fertilised, and fourteen self-fertilised plants spontaneously self-fertilised, yielded capsules (the average number of seeds per capsule not having been ascertained) in the proportion of : 100 to 26.
PLANTS DERIVED FROM A CROSSED WITH A FRESH STOCK COMPARED WITH INTERCROSSED PLANTS.
The offspring of intercrossed plants of the ninth generation, crossed by a fresh stock, compared with plants of the same stock intercrossed during ten generations, both sets of plants left uncovered and naturally fertilised, produced capsules by weight as : 100 to 51.
We see in this table that the crossed plants are always in some degree more productive than the self-fertilised plants, by whatever standard they are compared. The degree differs greatly; but this depends chiefly on whether an average was taken of the seeds alone, or of the capsules alone, or of both combined. The relative superiority of the crossed plants is chiefly due to their producing a much greater number of capsules, and not to each capsule containing a larger average number of seeds. For instance, in the third generation the crossed and self-fertilised plants produced capsules in the ratio of 100 to 38, whilst the seeds in the capsules on the crossed plants were to those on the self-fertilised plants only as 100 to 94. In the eighth generation the capsules on two self-fertilised plants (not included in table 2/18), grown in separate pots and thus not subjected to any competition, yielded the large average of 5.1 seeds. The smaller number of capsules produced by the self-fertilised plants may be in part, but not altogether, attributed to their lessened size or height; this being chiefly due to their lessened constitutional vigour, so that they were not able to compete with the crossed plants growing in the same pots. The seeds produced by the crossed flowers on the crossed plants were not always heavier than the self-fertilised seeds on the self-fertilised plants. The lighter seeds, whether produced from crossed or self-fertilised flowers, generally germinated before the heavier seeds. I may add that the crossed plants, with very few exceptions, flowered before their self-fertilised opponents, as might have been expected from their greater height and vigour.
The impaired fertility of the self-fertilised plants was shown in another way, namely, by their anthers being smaller than those in the flowers on the crossed plants. This was first observed in the seventh generation, but may have occurred earlier. Several anthers from flowers on the crossed and self-fertilised plants of the eighth generation were compared under the microscope; and those from the former were generally longer and plainly broader than the anthers of the self-fertilised plants. The quantity of pollen contained in one of the latter was, as far as could be judged by the eye, about half of that contained in one from a crossed plant. The impaired fertility of the self-fertilised plants of the eighth generation was also shown in another manner, which may often be observed in hybrids—namely, by the first-formed flowers being sterile. For instance, the fifteen first flowers on a self-fertilised plant of one of the later generations were carefully fertilised with their own pollen, and eight of them dropped off; at the same time fifteen flowers on a crossed plant growing in the same pot were self-fertilised, and only one dropped off. On two other crossed plants of the same generation, several of the earliest flowers were observed to fertilise themselves and to produce capsules. In the plants of the ninth, and I believe of some previous generations, very many of the flowers, as already stated, were slightly monstrous; and this probably was connected with their lessened fertility.
All the self-fertilised plants of the seventh generation, and I believe of one or two previous generations, produced flowers of exactly the same tint, namely, of a rich dark purple. So did all the plants, without any exception, in the three succeeding generations of self-fertilised plants; and very many were raised on account of other experiments in progress not here recorded. My attention was first called to this fact by my gardener remarking that there was no occasion to label the self-fertilised plants, as they could always be known by their colour. The flowers were as uniform in tint as those of a wild species growing in a state of nature; whether the same tint occurred, as is probable, in the earlier generations, neither my gardener nor self could recollect. The flowers on the plants which were first raised from purchased seed, as well as during the first few generations, varied much in the depth of the purple tint; many were more or less pink, and occasionally a white variety appeared. The crossed plants continued to the tenth generation to vary in the same manner as before, but to a much less degree, owing, probably, to their having become more or less closely inter-related. We must therefore attribute the extraordinary uniformity of colour in the flowers on the plants of the seventh and succeeding self-fertilised generations, to inheritance not having been interfered with by crosses during several preceding generations, in combination with the conditions of life having been very uniform.
A plant appeared in the sixth self-fertilised generation, named the Hero, which exceeded by a little in height its crossed antagonist, and which transmitted its powers of growth and increased self-fertility to its children and grandchildren. A cross between the children of Hero did not give to the grandchildren any advantage over the self-fertilised grandchildren raised from the self-fertilised children. And as far as my observations can be trusted, which were made on very unhealthy plants, the great-grandchildren raised from intercrossing the grandchildren had no advantage over the seedlings from the grandchildren the product of continued self-fertilisation; and what is far more remarkable, the great-grandchildren raised by crossing the grandchildren with a fresh stock, had no advantage over either the intercrossed or self-fertilised great-grandchildren. It thus appears that Hero and its descendants differed in constitution in an extraordinary manner from ordinary plants of the present species.
Although the plants raised during ten successive generations from crosses between distinct yet inter-related plants almost invariably exceeded in height, constitutional vigour, and fertility their self-fertilised opponents, it has been proved that seedlings raised by intercrossing flowers on the same plant are by no means superior, on the contrary are somewhat inferior in height and weight, to seedlings raised from flowers fertilised with their own pollen. This is a remarkable fact, which seems to indicate that self-fertilisation is in some manner more advantageous than crossing, unless the cross brings with it, as is generally the case, some decided and preponderant advantage; but to this subject I shall recur in a future chapter.
The benefits which so generally follow from a cross between two plants apparently depend on the two differing somewhat in constitution or character. This is shown by the seedlings from the intercrossed plants of the ninth generation, when crossed with pollen from a fresh stock, being as superior in height and almost as superior in fertility to the again intercrossed plants, as these latter were to seedlings from self-fertilised plants of the corresponding generation. We thus learn the important fact that the mere act of crossing two distinct plants, which are in some degree inter-related and which have been long subjected to nearly the same conditions, does little good as compared with that from a cross between plants belonging to different stocks or families, and which have been subjected to somewhat different conditions. We may attribute the good derived from the crossing of the intercrossed plants during the ten successive generations to their still differing somewhat in constitution or character, as was indeed proved by their flowers still differing somewhat in colour. But the several conclusions which may be deduced from the experiments on Ipomoea will be more fully considered in the final chapters, after all my other observations have been given.