Lathyrus odoratus (Second Generation).
Although the crossed plants were to the self-fertilised in height as 100 to 88, yet there was no marked difference in their period of flowering.
Lobelia fulgens (Second Generation).
Although the crossed plants were to the self-fertilised in height as 100 to 91, yet they flowered simultaneously.
Nicotiana tabac.u.m (Third Generation).
Although the crossed plants were to the self-fertilised in height as 100 to 83, yet in half the pots a self-fertilised plant flowered first, and in the other half a crossed plant.]
These three lists include fifty-eight cases, in which the period of flowering of the crossed and self-fertilised plants was recorded. In forty-four of them a crossed plant flowered first either in a majority of the pots or in all; in nine instances a self-fertilised plant flowered first, and in five the two lots flowered simultaneously. One of the most striking cases is that of Cyclamen, in which the crossed plants flowered some weeks before the self-fertilised in all four pots during two seasons. In the second generation of Lobelia ramosa, a crossed plant flowered in all four pots some days before any one of the self-fertilised. Plants derived from a cross with a fresh stock generally showed a very strongly marked tendency to flower before the self-fertilised and the intercrossed plants of the old stock; all three lots growing in the same pots. Thus with Mimulus and Dianthus, in only one pot out of ten, and in Nicotiana in only one pot out of sixteen, did a self-fertilised plant flower before the plants of the two crossed kinds,--these latter flowering almost simultaneously.
A consideration of the two first lists, especially of the second one, shows that a tendency to flower first is generally connected with greater power of growth, that is, with greater height. But there are some remarkable exceptions to this rule, proving that some other cause comes into play. Thus the crossed plants both of Lupinus luteus and Clarkia elegans were to the self-fertilised plants in height as 100 to 82, and yet the latter flowered first. In the third generation of Nicotiana, and in all three generations of Canna, the crossed and self-fertilised plants were of nearly equal height, yet the self-fertilised tended to flower first. On the other hand, with Primula sinensis, plants raised from a cross between two distinct individuals, whether these were legitimately or illegitimately crossed, flowered before the illegitimately self-fertilised plants, although all the plants were of nearly equal height in both cases. So it was with respect to height and flowering with Phaseolus, Specularia, and Borago. The crossed plants of Hibiscus were inferior in height to the self-fertilised, in the ratio of 100 to 109, and yet they flowered before the self-fertilised in three out of the four pots. On the whole, there can be no doubt that the crossed plants exhibit a tendency to flower before the self-fertilised, almost though not quite so strongly marked as to grow to a greater height, to weigh more, and to be more fertile.
A few other cases not included in the above three lists deserve notice.
In all three pots of Viola tricolor, naturally crossed plants the offspring of crossed plants flowered before naturally crossed plants the offspring of self-fertilised plants. Flowers on two plants, both of self-fertilised parentage, of the sixth generation of Mimulus luteus were intercrossed, and other flowers on the same plants were fertilised with their own pollen; intercrossed seedlings and seedlings of the seventh self-fertilised generation were thus raised, and the latter flowered before the intercrossed in three out of the five pots. Flowers on a plant both of Mimulus luteus and of Ipomoea purpurea were crossed with pollen from other flowers on the same plant, and other flowers were fertilised with their own pollen; intercrossed seedlings of this peculiar kind, and others strictly self-fertilised being thus raised. In the case of the Mimulus the self-fertilised plants flowered first in seven out of the eight pots, and in the case of the Ipomoea in eight out of the ten pots; so that an intercross between the flowers on the same plant was very far from giving to the offspring thus raised, any advantage over the strictly self-fertilised plants in their period of flowering.
EFFECTS OF CROSSING FLOWERS ON THE SAME PLANT.
In the discussion on the results of a cross with a fresh stock, given under Table 7/C in the last chapter, it was shown that the mere act of crossing by itself does no good; but that the advantages thus derived depend on the plants which are crossed, either consisting of distinct varieties which will almost certainly differ somewhat in const.i.tution, or on the progenitors of the plants which are crossed, though identical in every external character, having been subjected to somewhat different conditions and having thus acquired some slight difference in const.i.tution. All the flowers produced by the same plant have been developed from the same seed; those which expand at the same time have been exposed to exactly the same climatic influences; and the stems have all been nourished by the same roots. Therefore in accordance with the conclusion just referred to, no good ought to result from crossing flowers on the same plant. (8/1. It is, however, possible that the stamens which differ in length or construction in the same flower may produce pollen differing in nature, and in this manner a cross might be made effective between the several flowers on the same plant. Mr. Macnab states in a communication to M. Verlot "La Production des Varietes" 1865 page 42, that seedlings raised from the shorter and longer stamens of rhododendron differ in character; but the shorter stamens apparently are becoming rudimentary, and the seedlings are dwarfs, so that the result may be simply due to a want of fertilising power in the pollen, as in the case of the dwarfed plants of Mirabilis raised by Naudin by the use of too few pollen-grains. a.n.a.logous statements have been made with respect to the stamens of Pelargonium. With some of the Melastomaceae, seedlings raised by me from flowers fertilised by pollen from the shorter stamens, certainly differed in appearance from those raised from the longer stamens, with differently coloured anthers; but here, again, there is some reason for believing that the shorter stamens are tending towards abortion. In the very different case of trimorphic heterostyled plants, the two sets of stamens in the same flower have widely different fertilising powers.) In opposition to this conclusion is the fact that a bud is in one sense a distinct individual, and is capable of occasionally or even not rarely a.s.suming new external characters, as well as new const.i.tutional peculiarities. Plants raised from buds which have thus varied may be propagated for a great length of time by grafts, cuttings, etc., and sometimes even by seminal generation. (8/2. I have given numerous cases of such bud-variations in my "Variation of Animals and Plants under Domestication" chapter 11 2nd edition volume 1 page 448.) There exist also numerous species in which the flowers on the same plant differ from one another,--as in the s.e.xual organs of monoecious and polygamous plants,--in the structure of the circ.u.mferential flowers in many Compositae, Umbelliferae, etc.,--in the structure of the central flower in some plants,--in the two kinds of flowers produced by cleistogene species,--and in several other such cases. These instances clearly prove that the flowers on the same plant have often varied independently of one another in many important respects, such variations having been fixed, like those on distinct plants during the development of species.
It was therefore necessary to ascertain by experiment what would be the effect of intercrossing flowers on the same plant, in comparison with fertilising them with their own pollen or crossing them with pollen from a distinct plant. Trials were carefully made on five genera belonging to four families; and in only one case, namely, Digitalis, did the offspring from a cross between the flowers on the same plant receive any benefit, and the benefit here was small compared with that derived from a cross between distinct plants. In the chapter on Fertility, when we consider the effects of cross-fertilisation and self-fertilisation on the productiveness of the parent-plants we shall arrive at nearly the same result, namely, that a cross between the flowers on the same plant does not at all increase the number of the seeds, or only occasionally and to a slight degree. I will now give an abstract of the results of the five trials which were made.
1. Digitalis purpurea.
Seedlings raised from intercrossed flowers on the same plant, and others from flowers fertilised with their own pollen, were grown in the usual manner in compet.i.tion with one another on the opposite sides of ten pots. In this and the four following cases, the details may be found under the head of each species. In eight pots, in which the plants did not grow much crowded, the flower-stems on sixteen intercrossed plants were in height to those on sixteen self-fertilised plants, as 100 to 94.
In the two other pots on which the plants grew much crowded, the flower-stems on nine intercrossed plants were in height to those on nine self-fertilised plants, as 100 to 90. That the intercrossed plants in these two latter pots had a real advantage over their self-fertilised opponents, was well shown by their relative weights when cut down, which was as 100 to 78. The mean height of the flower-stems on the twenty-five intercrossed plants in the ten pots taken together, was to that of the flower-stems on the twenty-five self-fertilised plants, as 100 to 92.
Thus the intercrossed plants were certainly superior to the self-fertilised in some degree; but their superiority was small compared with that of the offspring from a cross between distinct plants over the self-fertilised, this being in the ratio of 100 to 70 in height. Nor does this latter ratio show at all fairly the great superiority of the plants derived from a cross between distinct individuals over the self-fertilised, as the former produced more than twice as many flower-stems as the latter, and were much less liable to premature death.
2. Ipomoea purpurea.
Thirty-one intercrossed plants raised from a cross between flowers on the same plants were grown in ten pots in compet.i.tion with the same number of self-fertilised plants, and the former were to the latter in height as 100 to 105. So that the self-fertilised plants were a little taller than the intercrossed; and in eight out of the ten pots a self-fertilised plant flowered before any one of the crossed plants in the same pots. The plants which were not greatly crowded in nine of the pots (and these offer the fairest standard of comparison) were cut down and weighed; and the weight of the twenty-seven intercrossed plants was to that of the twenty-seven self-fertilised as 100 to 124; so that by this test the superiority of the self-fertilised was strongly marked. To this subject of the superiority of the self-fertilised plants in certain cases, I shall have to recur in a future chapter. If we now turn to the offspring from a cross between distinct plants when put into compet.i.tion with self-fertilised plants, we find that the mean height of seventy-three such crossed plants, in the course of ten generations, was to that of the same number of self-fertilised plants as 100 to 77; and in the case of the plants of the tenth generation in weight as 100 to 44. Thus the contrast between the effects of crossing flowers on the same plant, and of crossing flowers on distinct plants, is wonderfully great.
3. Mimulus luteus.
Twenty-two plants raised by crossing flowers on the same plant were grown in compet.i.tion with the same number of self-fertilised plants; and the former were to the latter in height as 100 to 105, and in weight as 100 to 103. Moreover, in seven out of the eight pots a self-fertilised plant flowered before any of the intercrossed plants. So that here again the self-fertilised exhibit a slight superiority over the intercrossed plants. For the sake of comparison, I may add that seedlings raised during three generations from a cross between distinct plants were to the self-fertilised plants in height as 100 to 65.
4. Pelargonium zonale.
Two plants growing in separate pots, which had been propagated by cuttings from the same plant, and therefore formed in fact parts of the same individual, were intercrossed, and other flowers on one of these plants were self-fertilised; but the seedlings obtained by the two processes did not differ in height. When, on the other hand, flowers on one of the above plants were crossed with pollen taken from a distinct seedling, and other flowers were self-fertilised, the crossed offspring thus obtained were to the self-fertilised in height as 100 to 74.
5. Origanum vulgare.
A plant which had been long cultivated in my kitchen garden, had spread by stolons so as to form a large bed or clump. Seedlings raised by intercrossing flowers on these plants, which strictly consisted of the same plant, and other seedlings raised from self-fertilised flowers, were carefully compared from their earliest youth to maturity; and they did not differ at all in height or in const.i.tutional vigour. Some flowers on these seedlings were then crossed with pollen taken from a distinct seedling, and other flowers were self-fertilised; two fresh lots of seedlings being thus raised, which were the grandchildren of the plant that had spread by stolons and formed a large clump in my garden.
These differed much in height, the crossed plants being to the self-fertilised as 100 to 86. They differed, also, to a wonderful degree in const.i.tutional vigour. The crossed plants flowered first, and produced exactly twice as many flower-stems; and they afterwards increased by stolons to such an extent as almost to overwhelm the self-fertilised plants.
Reviewing these five cases, we see that in four of them, the effect of a cross between flowers on the same plant (even on offsets of the same plant growing on separate roots, as with the Pelargonium and Origanum) does not differ from that of the strictest self-fertilisation. Indeed, in two of the cases the self-fertilised plants were superior to such intercrossed plants. With Digitalis a cross between the flowers on the same plant certainly did do some good, yet very slight compared with that from a cross between distinct plants. On the whole the results here arrived at, if we bear in mind that the flower-buds are to a certain extent distinct individuals and occasionally vary independently of one another, agree well with our general conclusion, that the advantages of a cross depend on the progenitors of the crossed plants possessing somewhat different const.i.tutions, either from having been exposed to different conditions, or to their having varied from unknown causes in a manner which we in our ignorance are forced to speak of as spontaneous.
Hereafter I shall have to recur to this subject of the inefficiency of a cross between the flowers on the same plant, when we consider the part which insects play in the cross-fertilisation of flowers.
ON THE TRANSMISSION OF THE GOOD EFFECTS FROM A CROSS AND OF THE EVIL EFFECTS FROM SELF-FERTILISATION.
We have seen that seedlings from a cross between distinct plants almost always exceed their self-fertilised opponents in height, weight, and const.i.tutional vigour, and, as will hereafter be shown, often in fertility. To ascertain whether this superiority would be transmitted beyond the first generation, seedlings were raised on three occasions from crossed and self-fertilised plants, both sets being fertilised in the same manner, and therefore not as in the many cases given in Tables 7/A, 7/B, 7/C, in which the crossed plants were again crossed and the self-fertilised again self-fertilised.
Firstly, seedlings were raised from self-fertilised seeds produced under a net by crossed and self-fertilised plants of Nemophila insignis; and the latter were to the former in height as 133 to 100. But these seedlings became very unhealthy early in life, and grew so unequally that some of them in both lots were five times as tall as the others.
Therefore this experiment was quite worthless; but I have felt bound to give it, as opposed to my general conclusion. I should state that in this and the two following trials, both sets of plants were grown on the opposite sides of the same pots, and treated in all respects alike. The details of the experiments may be found under the head of each species.
Secondly, a crossed and a self-fertilised plant of Heartsease (Viola tricolor) grew near together in the open ground and near to other plants of heartsease; and as both produced an abundance of very fine capsules, the flowers on both were certainly cross-fertilised by insects. Seeds were collected from both plants, and seedlings raised from them. Those from the crossed plants flowered in all three pots before those from the self-fertilised plants; and when fully grown the former were to the latter in height as 100 to 82. As both sets of plants were the product of cross-fertilisation, the difference in their growth and period of flowering was clearly due to their parents having been of crossed and self-fertilised parentage; and it is equally clear that they transmitted different const.i.tutional powers to their offspring, the grandchildren of the plants which were originally crossed and self-fertilised.
Thirdly, the Sweet Pea (Lathyrus odoratus) habitually fertilises itself in this country. As I possessed plants, the parents and grandparents of which had been artificially crossed and other plants descended from the same parents which had been self-fertilised for many previous generations, these two lots of plants were allowed to fertilise themselves under a net, and their self-fertilised seeds saved. The seedlings thus raised were grown in compet.i.tion with each other in the usual manner, and differed in their powers of growth. Those from the self-fertilised plants which had been crossed during the two previous generations were to those from the plants self-fertilised during many previous generations in height as 100 to 90. These two lots of seeds were likewise tried by being sown under very unfavourable conditions in poor exhausted soil, and the plants whose grandparents and great-grandparents had been crossed showed in an unmistakable manner their superior const.i.tutional vigour. In this case, as in that of the heartsease, there could be no doubt that the advantage derived from a cross between two plants was not confined to the offspring of the first generation. That const.i.tutional vigour due to cross-parentage is transmitted for many generations may also be inferred as highly probable, from some of Andrew Knight"s varieties of the common pea, which were raised by crossing distinct varieties, after which time they no doubt fertilised themselves in each succeeding generation. These varieties lasted for upwards of sixty years, "but their glory is now departed." (8/3. See the evidence on this head in my "Variation under Domestication" chapter 9 volume 1 2nd edition page 397.) On the other hand, most of the varieties of the common pea, which there is no reason to suppose owe their origin to a cross, have had a much shorter existence. Some also of Mr. Laxton"s varieties produced by artificial crosses have retained their astonishing vigour and luxuriance for a considerable number of generations; but as Mr. Laxton informs me, his experience does not extend beyond twelve generations, within which period he has never perceived any diminution of vigour in his plants.
An allied point may be here noticed. As the force of inheritance is strong with plants (of which abundant evidence could be given), it is almost certain that seedlings from the same capsule or from the same plant would tend to inherit nearly the same const.i.tution; and as the advantage from a cross depends on the plants which are crossed differing somewhat in const.i.tution, it may be inferred as probable that under similar conditions a cross between the nearest relations would not benefit the offspring so much as one between non-related plants. In support of this conclusion we have some evidence, as Fritz Muller has shown by his valuable experiments on hybrid Abutilons, that the union of brothers and sisters, parents and children, and of other near relations is highly injurious to the fertility of the offspring. In one case, moreover, seedlings from such near relations possessed very weak const.i.tutions. (8/4. "Jenaische Zeitschrift fur Naturw." B. 7 pages 22 and 45 1872 and 1873 pages 441-450.) This same observer also found three plants of a Bignonia growing near together. (8/5. "Botanische Zeitung"
1868 page 626.) He fertilised twenty-nine flowers on one of them with their own pollen, and they did not set a single capsule. Thirty flowers were then fertilised with pollen from a distinct plant, one of the three growing together, and they yielded only two capsules. Lastly, five flowers were fertilised with pollen from a fourth plant growing at a distance, and all five produced capsules. It seems therefore probable, as Fritz Muller suggests, that the three plants growing near together were seedlings from the same parent, and that from being closely related they had little power of fertilising one another. (8/6. Some remarkable cases are given in my "Variation under Domestication" chapter 17 2nd edition volume 2 page 121, of hybrids of Gladiolus and Cistus, any one of which could be fertilised by pollen from any other, but not by its own pollen.)
Lastly, the fact of the intercrossed plants in Table 7/A not exceeding in height the self-fertilised plants in a greater and greater degree in the later generations, is probably the result of their having become more and more closely inter-related.
UNIFORM COLOUR OF THE FLOWERS ON PLANTS, SELF-FERTILISED AND GROWN UNDER SIMILAR CONDITIONS FOR SEVERAL GENERATIONS.
At the commencement of my experiments, the parent-plants of Mimulus luteus, Ipomoea purpurea, Dianthus caryophyllus, and Petunia violacea, raised from purchased seeds, varied greatly in the colour of their flowers. This occurs with many plants which have been long cultivated as an ornament for the flower-garden, and which have been propagated by seeds. The colour of the flowers was a point to which I did not at first in the least attend, and no selection whatever was practised.
Nevertheless, the flowers produced by the self-fertilised plants of the above four species became absolutely uniform in tint, or very nearly so, after they had been grown for some generations under closely similar conditions. The intercrossed plants, which were more or less closely inter-related in the later generations, and which had been likewise cultivated all the time under similar conditions, became more uniform in the colour of their flowers than were the original parent-plants, but much less so than the self-fertilised plants. When self-fertilised plants of one of the later generations were crossed with a fresh stock, and seedlings thus raised, these presented a wonderful contrast in the diversified tints of their flowers compared with those of the self-fertilised seedlings. As such cases of flowers becoming uniformly coloured without any aid from selection seem to me curious, I will give a full abstract of my observations.
Mimulus luteus.
A tall variety, bearing large, almost white flowers blotched with crimson, appeared amongst the intercrossed and self-fertilised plants of the third and fourth generations. This variety increased so rapidly, that in the sixth generation of self-fertilised plants every single one consisted of it. So it was with all the many plants which were raised, up to the last or ninth self-fertilised generation. Although this variety first appeared amongst the intercrossed plants, yet from their offspring being intercrossed in each succeeding generation, it never prevailed amongst them; and the flowers on the several intercrossed plants of the ninth generation differed considerably in colour. On the other hand, the uniformity in colour of the flowers on the plants of all the later self-fertilised generations was quite surprising; on a casual inspection they might have been said to be quite alike, but the crimson blotches were not of exactly the same shape, or in exactly the same position. Both my gardener and myself believe that this variety did not appear amongst the parent-plants, raised from purchased seeds, but from its appearance amongst both the crossed and self-fertilised plants of the third and fourth generations; and from what I have seen of the variation of this species on other occasions, it is probable that it would occasionally appear under any circ.u.mstances. We learn, however, from the present case that under the peculiar conditions to which my plants were subjected, this particular variety, remarkable for its colouring, largeness of the corolla, and increased height of the whole plant, prevailed in the sixth and all the succeeding self-fertilised generations to the complete exclusion of every other variety.
Ipomoea purpurea.
My attention was first drawn to the present subject by observing that the flowers on all the plants of the seventh self-fertilised generation were of a uniform, remarkably rich, dark purple tint. The many plants which were raised during the three succeeding generations, up to the last or tenth, all produced flowers coloured in the same manner. They were absolutely uniform in tint, like those of a constant species living in a state of nature; and the self-fertilised plants might have been distinguished with certainty, as my gardener remarked, without the aid of labels, from the intercrossed plants of the later generations. These, however, had more uniformly coloured flowers than those which were first raised from the purchased seeds. This dark purple variety did not appear, as far as my gardener and myself could recollect, before the fifth or sixth self-fertilised generation. However this may have been, it became, through continued self-fertilisation and the cultivation of the plants under uniform conditions, perfectly constant, to the exclusion of every other variety.
Dianthus caryophyllus.
The self-fertilised plants of the third generation all bore flowers of exactly the same pale rose-colour; and in this respect they differed quite remarkably from the plants growing in a large bed close by and raised from seeds purchased from the same nursery garden. In this case it is not improbable that some of the parent-plants which were first self-fertilised may have borne flowers thus coloured; but as several plants were self-fertilised in the first generation, it is extremely improbable that all bore flowers of exactly the same tint as those of the self-fertilised plants of the third generation. The intercrossed plants of the third generation likewise produced flowers almost, though not quite so uniform in tint as those of the self-fertilised plants.
Petunia violacea.
In this case I happened to record in my notes that the flowers on the parent-plant which was first self-fertilised were of a "dingy purple colour." In the fifth self-fertilised generation, every one of the twenty-one self-fertilised plants growing in pots, and all the many plants in a long row out of doors, produced flowers of absolutely the same tint, namely, of a dull, rather peculiar and ugly flesh colour; therefore, considerably unlike those on the parent-plant. I believe that this change of colour supervened quite gradually; but I kept no record, as the point did not interest me until I was struck with the uniform tint of the flowers on the self-fertilised plants of the fifth generation. The flowers on the intercrossed plants of the corresponding generation were mostly of the same dull flesh colour, but not nearly so uniform as those on the self-fertilised plants, some few being very pale, almost white. The self-fertilised plants which grew in a long row in the open ground were also remarkable for their uniformity in height, as were the intercrossed plants in a less degree, both lots being compared with a large number of plants raised at the same time under similar conditions from the self-fertilised plants of the fourth generation crossed by a fresh stock. I regret that I did not attend to the uniformity in height of the self-fertilised seedlings in the later generations of the other species.
These few cases seem to me to possess much interest. We learn from them that new and slight shades of colour may be quickly and firmly fixed, independently of any selection, if the conditions are kept as nearly uniform as is possible, and no intercrossing be permitted. With Mimulus, not only a grotesque style of colouring, but a larger corolla and increased height of the whole plant were thus fixed; whereas with most plants which have been long cultivated for the flower-garden, no character is more variable than that of colour, excepting perhaps that of height. From the consideration of these cases we may infer that the variability of cultivated plants in the above respects is due, firstly, to their being subjected to somewhat diversified conditions, and, secondly, to their being often intercrossed, as would follow from the free access of insects. I do not see how this inference can be avoided, as when the above plants were cultivated for several generations under closely similar conditions, and were intercrossed in each generation, the colour of their flowers tended in some degree to change and to become uniform. When no intercrossing with other plants of the same stock was allowed,--that is, when the flowers were fertilised with their own pollen in each generation--their colour in the later generations became as uniform as that of plants growing in a state of nature, accompanied at least in one instance by much uniformity in the height of the plants. But in saying that the diversified tints of the flowers on cultivated plants treated in the ordinary manner are due to differences in the soil, climate, etc., to which they are exposed, I do not wish to imply that such variations are caused by these agencies in any more direct manner than that in which the most diversified illnesses, as colds, inflammation of the lungs or pleura, rheumatism, etc., may be said to be caused by exposure to cold. In both cases the const.i.tution of the being which is acted on is of preponderant importance.
CHAPTER IX.
THE EFFECTS OF CROSS-FERTILISATION AND SELF-FERTILISATION ON THE PRODUCTION OF SEEDS.
Fertility of plants of crossed and self-fertilised parentage, both lots being fertilised in the same manner.
Fertility of the parent-plants when first crossed and self-fertilised, and of their crossed and self-fertilised offspring when again crossed and self-fertilised.
Comparison of the fertility of flowers fertilised with their own pollen and with that from other flowers on the same plant.