[This book, as pointed out in the 'Autobiography,' is a complement to the 'Fertilisation of Orchids,' because it shows how important are the results of cross-fertilisation which are ensured by the mechanisms described in that book.
By proving that the offspring of cross-fertilisation are more vigorous than the offspring of self-fertilisation, he showed that one circumstance which influences the fate of young plants in the struggle for life is the degree to which their parents are fitted for cross-fertilisation. He thus convinced himself that the intensity of the struggle (which he had elsewhere shown to exist among young plants) is a measure of the strength of a selective agency perpetually sifting out every modification in the structure of flowers which can effect its capabilities for cross-fertilisation.
The book is also valuable in another respect, because it throws light on the difficult problems of the origin of sexuality. The increased vigour resulting from cross-fertilisation is allied in the closest manner to the advantage gained by change of conditions. So strongly is this the case, that in some instances cross-fertilisation gives no advantage to the offspring, unless the parents have lived under slightly different conditions. So that the really important thing is not that two individuals of different BLOOD shall unite, but two individuals which have been subjected to different conditions. We are thus led to believe that sexuality is a means for infusing vigour into the offspring by the coalescence of differentiated elements, an advantage which could not follow if reproductions were entirely asexual.
It is remarkable that this book, the result of eleven years of experimental work, owed its origin to a chance observation. My father had raised two beds of Linaria vulgaris--one set being the offspring of cross- and the other of self-fertilisation. These plants were grown for the sake of some observations on inheritance, and not with any view to cross-breeding, and he was astonished to observe that the offspring of self-fertilisation were clearly less vigorous than the others. It seemed incredible to him that this result could be due to a single act of self-fertilisation, and it was only in the following year when precisely the same result occurred in the case of a similar experiment on inheritance in Carnations, that his attention was "thoroughly aroused" and that he determined to make a series of experiments specially directed to the question. The following letters give some account of the work in question.]
CHARLES DARWIN TO ASA GRAY.
September 10, [1866?].
...I have just begun a large course of experiments on the germination of the seed, and on the growth of the young plants when raised from a pistil fertilised by pollen from the same flower, and from pollen from a distinct plant of the same, or of some other variety. I have not made sufficient experiments to judge certainly, but in some cases the difference in the growth of the young plants is highly remarkable. I have taken every kind of precaution in getting seed from the same plant, in germinating the seed on my own chimney-piece, in planting the seedlings in the same flower-pot, and under this similar treatment I have seen the young seedlings from the crossed seed exactly twice as tall as the seedlings from the self-fertilised seed; both seeds having germinated on the same day. If I can establish this fact (but perhaps it will all go to the dogs), in some fifty cases, with plants of different orders, I think it will be very important, for then we shall positively know why the structure of every flower permits, or favours, or necessitates an occasional cross with a distinct individual. But all this is rather cooking my hare before I have caught it. But somehow it is a great pleasure to me to tell you what I am about.
Believe me, my dear Gray, Ever yours most truly, and with cordial thanks, CH. DARWIN.
CHARLES DARWIN TO G. BENTHAM.
April 22, 1868.