04 - Natural Selection

04-08 - On the Intercrossing of Individuals

In the first place, I have collected so large a body of facts, and made so many experiments, showing, in accordance with the almost universal belief of breeders, that with animals and plants a cross between different varieties, or between individuals of the same variety but of another strain, gives vigour and fertility to the offspring; and on the other hand, that close interbreeding diminishes vigour and fertility; that these facts alone incline me to believe that it is a general law of
nature that no organic being fertilises itself for a perpetuity of generations; but that a cross with another individual is occasionally- perhaps at long intervals of time- indispensable.

04 - Natural Selection

04-08 - On the Intercrossing of Individuals

When the stamens of a flower suddenly spring towards the pistil, or slowly move one after the other towards it, the contrivance seems adapted solely to ensure self-fertilisation; and no doubt it is useful for this end: but the agency of insects is often required to cause the stamens to spring forward, as Kolreuter has shown to be the case with the barberry; and in this very genus, which seems to have a special contrivance for self-fertilisation, it is well known that, if closely allied forms or varieties are planted near each other, it is hardly possible to raise pure seedlings, so largely do they naturally cross.

barberry

In numerous other cases, far from self-fertilisation being favoured, there are special contrivances which effectually prevent the stigma receiving pollen from its own flower, as I could show from the works of Sprengel and others, as well as from my own observations: for instance, in Lobelia fulgens, there is a really beautiful and elaborate contrivance by which all the infinitely numerous pollen-granules are swept out of the conjoined anthers of each flower, before the stigma of that individual flower is ready to receive them; and as this flower is never visited, at least in my garden, by insects, it never sets a seed, though by placing pollen from one flower on the stigma of another, I raise plenty of seedlings.

Lobelia Fulgens

Another species of Lobelia which is visited by bees, seeds freely in my garden.

In very many other cases, though there is no special mechanical contrivance to prevent the stigma receiving pollen from the same flower, yet, as Sprengel, and more recently Hildebrand, and others, have shown, and as I can confirm, either the anthers burst before the stigma is ready for fertilisation, or the stigma is ready before the pollen of that flower is ready, so that these so-named dichogamous plants have in fact separated sexes, and must habitually be crossed.

So it is with the reciprocally dimorphic and trimorphic plants previously alluded to. How strange are these facts!

How strange that the pollen and stigmatic surface of the same flower, though placed so close together, as if for the very purpose of self-fertilisation, should be in so many cases mutually useless to each other! How simply are these facts explained on the view of an occasional cross with a distinct individual being advantageous or indispensable!

04 - Natural Selection

04-08 - On the Intercrossing of Individuals

In the case of a large tree covered with innumerable flowers, it may be objected that pollen could seldom be carried from tree to tree, and at most only from flower to flower on the same tree; and flowers on the same tree can be considered as distinct individuals only in a limited sense.

tree

I believe this objection to be valid, but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes.

When the sexes are separated, although the male and female flowers may be produced on the same tree, pollen must be regularly carried from flower to flower; and this will give a better chance of pollen being occasionally carried from tree to tree.

That trees belonging to all Orders have their sexes more often separated than other plants, I find to be the case in this country; and at my request Dr. Hooker tabulated the trees of New Zealand, and Dr. Asa Gray those of the United States, and the result was as I anticipated.

England

New Zealand

United States

On the other hand, Dr. Hooker informs me that the rule does not hold good in Australia but if most of the Australian trees are dichogamous, the same result would follow as if they bore flowers with separated sexes.

Australia

I have made these few remarks on trees simply to call attention to the subject.

04 - Natural Selection

04-08 - On the Intercrossing of Individuals

It must have struck most naturalists as a strange anomaly that, both with animals and plants, some species of the same family and even of the same genus, though agreeing closely with each other in their whole organisation, are hermaphrodites, and some unisexual.

But if, in fact, all hermaphrodites do occasionally intercross, the difference between them and unisexual species is, as far as function is concerned, very small.

From these several considerations and from the many special facts which I have collected, but which I am unable here to give, it appears that with animals and plants an occasional intercross between distinct individuals is a very general, if not universal, law of nature.