The Effects of Cross & Self-Fertilisation in the Vegetable Kingdom

by Charles Darwin

This species offers another interesting case. In the five first generations in which intercrossed and self-fertilised plants were put into competition with one another, every single intercrossed plant beat its self-fertilised antagonist, except in one instance, in which they were equal in height. But in the sixth generation a plant appeared, named by me the Hero, remarkable for its tallness and increased self-fertility, and which transmitted its characters to the next three generations. The children of Hero were again self-fertilised, forming the eighth self-fertilised generation, and were likewise intercrossed one with another; but this cross between plants which had been subjected to the same conditions and had been self-fertilised during the seven previous generations, did not effect the least good; for the intercrossed grandchildren were actually shorter than the self-fertilised grandchildren, in the ratio of 100 to 107. We here see that the mere act of crossing two distinct plants does not by itself benefit the offspring. This case is almost the converse of that in the last paragraph, on which the offspring profited so greatly by a cross with a fresh stock. A similar trial was made with the descendants of Hero in the following generation, and with the same result. But the trial cannot be fully trusted, owing to the extremely unhealthy condition of the plants. Subject to this same serious cause of doubt, even a cross with a fresh stock did not benefit the great-grandchildren of Hero; and if this were really the case, it is the greatest anomaly observed by me in all my experiments.

2. Mimulus luteus.

During the three first generations the intercrossed plants taken together exceeded in height the self-fertilised taken together, in the ratio of 100 to 65, and in fertility in a still higher degree. In the fourth generation a new variety, which grew taller and had whiter and larger flowers than the old varieties, began to prevail, especially amongst the self-fertilised plants. This variety transmitted its characters with remarkable fidelity, so that all the plants in the later self-fertilised generations belonged to it. These consequently exceeded the intercrossed plants considerably in height. Thus in the seventh generation the intercrossed plants were to the self-fertilised in height as 100 to 137. It is a more remarkable fact that the self-fertilised plants of the sixth generation had become much more fertile than the intercrossed plants, judging by the number of capsules spontaneously produced, in the ratio of 147 to 100. This variety, which as we have seen appeared amongst the plants of the fourth self-fertilised generation, resembles in almost all its constitutional peculiarities the variety called Hero which appeared in the sixth self-fertilised generation of Ipomoea. No other such case, with the partial exception of that of Nicotiana, occurred in my experiments, carried on during eleven years.

Two plants of this variety of Mimulus, belonging to the sixth self-fertilised generation, and growing in separate pots, were intercrossed; and some flowers on the same plants were again self-fertilised. From the seeds thus obtained, plants derived from a cross between the self-fertilised plants, and others of the seventh self-fertilised generation, were raised. But this cross did not do the least good, the intercrossed plants being inferior in height to the self-fertilised, in the ratio of 100 to 110. This case is exactly parallel with that given under Ipomoea, of the grandchildren of Hero, and apparently of its great-grandchildren; for the seedlings raised by intercrossing these plants were not in any way superior to those of the corresponding generation raised from the self-fertilised flowers. Therefore in these several cases the crossing of plants, which had been self-fertilised for several generations and which had been cultivated all the time under as nearly as possible the same conditions, was not in the least beneficial.

Another experiment was now tried. Firstly, plants of the eighth self-fertilised generation were again self-fertilised, producing plants of the ninth self-fertilised generation. Secondly, two of the plants of the eighth self-fertilised generation were intercrossed one with another, as in the experiment above referred to; but this was now effected on plants which had been subjected to two additional generations of self-fertilisation. Thirdly, the same plants of the eighth self-fertilised generation were crossed with pollen from plants of a fresh stock brought from a distant garden. Numerous plants were raised from these three sets of seeds, and grown in competition with one another. The plants derived from a cross between the self-fertilised plants exceeded in height by a little the self-fertilised, namely, as 100 to 92; and in fertility in a greater degree, namely, as 100 to 73. I do not know whether this difference in the result, compared with that in the previous case, can be accounted for by the increased deterioration of the self-fertilised plants from two additional generations of self-fertilisation, and the consequent advantage of any cross whatever, along merely between the self-fertilised plants. But however this may be, the effects of crossing the self-fertilised plants of the eighth generation with a fresh stock were extremely striking; for the seedlings thus raised were to the self-fertilised of the ninth generation as 100 to 52 in height, and as 100 to 3 in fertility! They were also to the intercrossed plants (derived from crossing two of the self-fertilised plants of the eighth generation) in height as 100 to 56, and in fertility as 100 to 4. Better evidence could hardly be desired of the potent influence of a cross with a fresh stock on plants which had been self-fertilised for eight generations, and had been cultivated all the time under nearly uniform conditions, in comparison with plants self-fertilised for nine generations continuously, or then once intercrossed, namely in the last generation.

3. Brassica oleracea.

Some flowers on cabbage plants of the second self-fertilised generation were crossed with pollen from a plant of the same variety brought from a distant garden, and other flowers were again self-fertilised. Plants derived from a cross with a fresh stock and plants of the third self-fertilised generation were thus raised. The former were to the self-fertilised in weight as 100 to 22; and this enormous difference must be attributed in part to the beneficial effects of a cross with a fresh stock, and in part to the deteriorating effects of self-fertilisation continued during three generations.

4. Iberis umbellata.

Seedlings from a crimson English variety crossed by a pale-coloured variety which had been grown for some generations in Algiers, were to the self-fertilised seedlings from the crimson variety in height as 100 to 89, and as 100 to 75 in fertility. I am surprised that this cross with another variety did not produce a still more strongly marked beneficial effect; for some intercrossed plants of the crimson English variety, put into competition with plants of the same variety self-fertilised during three generations, were in height as 100 to 86, and in fertility as 100 to 75. The slightly greater difference in height in this latter case, may possibly be attributed to the deteriorating effects of self-fertilisation carried on for two additional generations.

5. Eschscholtzia californica.