This page contains affiliate links. As Amazon Associates we earn from qualifying purchases.
Language:
Genre:
Published:
  • 1876
Edition:
Collection:
Tag:
Buy it on Amazon FREE Audible 30 days

have met with only one exception to the rule of crossed plants being hardier than the self-fertilised: three long rows of Eschscholtzia plants, consisting of crossed seedlings from a fresh stock, of intercrossed seedlings of the same stock, and of self-fertilised ones, were left unprotected during a severe winter, and all perished except two of the self-fertilised. But this case is not so anomalous as it at first appears, for it should be remembered that the self-fertilised plants of Eschscholtzia always grow taller and are heavier than the crossed; the whole benefit of a cross with this species being confined to increased fertility.

Independently of any external cause which could be detected, the self-fertilised plants were more liable to premature death than were the crossed; and this seems to me a curious fact. Whilst the seedlings were very young, if one died its antagonist was pulled up and thrown away, and I believe that many more of the self-fertilised died at this early age than of the crossed; but I neglected to keep any record. With Beta vulgaris, however, it is certain that a large number of the self-fertilised seeds perished after germinating beneath the ground, whereas the crossed seeds sown at the same time did not thus suffer. When a plant died at a somewhat more advanced age the fact was recorded; and I find in my notes that out of several hundred plants, only seven of the crossed died, whilst of the self-fertilised at least twenty-nine were thus lost, that is more than four times as many. Mr. Galton, after examining some of my tables, remarks: “It is very evident that the columns with the self-fertilised plants include the larger number of exceptionally small plants;” and the frequent presence of such puny plants no doubt stands in close relation with their liability to premature death. The self-fertilised plants of Petunia completed their growth and began to wither sooner than did the intercrossed plants; and these latter considerably before the offspring from a cross with a fresh stock.

PERIOD OF FLOWERING.

In some cases, as with Digitalis, Dianthus, and Reseda, a larger number of the crossed than of the self-fertilised plants threw up flower-stems; but this probably was merely the result of their greater power of growth; for in the first generation of Lobelia fulgens, in which the self-fertilised plants greatly exceeded in height the crossed plants, some of the latter failed to throw up flower-stems. With a large number of species, the crossed plants exhibited a well-marked tendency to flower before the self-fertilised ones growing in the same pots. It should however be remarked that no record was kept of the flowering of many of the species; and when a record was kept, the flowering of the first plant in each pot was alone observed, although two or more pairs grew in the same pot. I will now give three lists,–one of the species in which the first plant that flowered was a crossed one,–a second in which the first that flowered was a self-fertilised plant,–and a third of those which flowered at the same time.

[SPECIES, OF WHICH THE FIRST PLANTS THAT FLOWERED WERE OF CROSSED PARENTAGE.

Ipomoea purpurea.

I record in my notes that in all ten generations many of the crossed plants flowered before the self-fertilised; but no details were kept.

Mimulus luteus (First Generation).

Ten flowers on the crossed plants were fully expanded before one on the self-fertilised.

Mimulus luteus (Second and Third Generation).

In both these generations a crossed plant flowered before one of the self-fertilised in all three pots.

Mimulus luteus (Fifth Generation).

In all three pots a crossed plant flowered first; yet the self-fertilised plants, which belonged to the new tall variety, were in height to the crossed as 126 to 100.

Mimulus luteus.

Plants derived from a cross with a fresh stock as well as the intercrossed plants of the old stock, flowered before the self-fertilised plants in nine out of the ten pots.

Salvia coccinea.

A crossed plant flowered before any one of the self-fertilised in all three pots.

Origanum vulgare.

During two successive seasons several crossed plants flowered before the self-fertilised.

Brassica oleracea (First Generation).

All the crossed plants growing in pots and in the open ground flowered first.

Brassica oleracea (Second Generation).

A crossed plant in three out of the four pots flowered before any one of the self-fertilised.

Iberis umbellata.

In both pots a crossed plant flowered first.

Eschscholtzia californica.

Plants derived from the Brazilian stock crossed by the English stock flowered in five out of the nine pots first; in four of them a self-fertilised plant flowered first; and not in one pot did an intercrossed plant of the old stock flower first.

Viola tricolor.

A crossed plant in five out of the six pots flowered before any one of the self-fertilised.

Dianthus caryophyllus (First Generation).

In two large beds of plants, four of the crossed plants flowered before any one of the self-fertilised.

Dianthus caryophyllus (Second Generation).

In both pots a crossed plant flowered first.

Dianthus caryophyllus (Third Generation).

In three out of the four pots a crossed plant flowered first; yet the crossed were to the self-fertilised in height only as 100 to 99, but in weight as 100 to 49.

Dianthus caryophyllus.

Plants derived from a cross with a fresh stock, and the intercrossed plants of the old stock, both flowered before the self-fertilised in nine out of the ten pots.

Hibiscus africanus.

In three out of the four pots a crossed plant flowered before any one of the self-fertilised; yet the latter were to the crossed in height as 109 to 100.

Tropaeolum minus.

A crossed plant flowered before any one of the self-fertilised in three out of the four pots, and simultaneously in the fourth pot.

Limnanthes douglasii.

A crossed plant flowered before any one of the self-fertilised in four out of the five pots.

Phaseolus multiflorus.

In both pots a crossed plant flowered first.

Specularia speculum.

In all four pots a crossed plant flowered first.

Lobelia ramosa (First Generation).

In all four pots a crossed plant flowered before any one of the self-fertilised.

Lobelia ramosa (Second Generation).

In all four pots a crossed plant flowered some days before any one of the self-fertilised.

Nemophila insignis.

In four out of the five pots a crossed plant flowered first.

Borago officinalis.

In both pots a crossed plant flowered first.

Petunia violacea (Second Generation).

In all three pots a crossed plant flowered first.

Nicotiana tabacum.

A plant derived from a cross with a fresh stock flowered before any one of the self-fertilised plants of the fourth generation, in fifteen out of the sixteen pots.

Cyclamen persicum.

During two successive seasons a crossed plant flowered some weeks before any one of the self-fertilised in all four pots.

Primula veris (equal-styled var.)

In all three pots a crossed plant flowered first.

Primula sinensis.

In all four pots plants derived from an illegitimate cross between distinct plants flowered before any one of the self-fertilised plants.

Primula sinensis.

A legitimately crossed plant flowered before any one of the self-fertilised plants in seven out of the eight pots.

Fagopyrum esculentum.

A legitimately crossed plant flowered from one to two days before any one of the self-fertilised plants in all three pots.

Zea mays.

In all four pots a crossed plant flowered first.

Phalaris canariensis.

The crossed plants flowered before the self-fertilised in the open ground, but simultaneously in the pots.

SPECIES OF WHICH THE FIRST PLANTS THAT FLOWERED WERE OF SELF-FERTILISED PARENTAGE.

Eschscholtzia californica (First Generation).

The crossed plants were at first taller than the self-fertilised, but on their second growth during the following year the self-fertilised exceeded the crossed in height, and now they flowered first in three out of the four pots.

Lupinus luteus.

Although the crossed plants were to the self-fertilised in height as 100 to 82; yet in all three pots the self-fertilised plants flowered first.

Clarkia elegans.

Although the crossed plants were, as in the last case, to the self-fertilised in height as 100 to 82, yet in the two pots the self-fertilised flowered first.

Lobelia fulgens (First Generation).

The crossed plants were to the self-fertilised in height only as 100 to 127, and the latter flowered much before the crossed.

Petunia violacea (Third Generation).

The crossed plants were to the self-fertilised in height as 100 to 131, and in three out of the four pots a self-fertilised plant flowered first; in the fourth pot simultaneously.

Petunia violacea (Fourth generation).

Although the crossed plants were to the self-fertilised in height as 100 to 69, yet in three out of the five pots a self-fertilised plant flowered first; in the fourth pot simultaneously, and only in the fifth did a crossed plant flower first.

Nicotiana tabacum (First Generation).

The crossed plants were to the self-fertilised in height only as 100 to 178, and a self-fertilised plant flowered first in all four pots.

Nicotiana tabacum (Third Generation).

The crossed plants were to the self-fertilised in height as 100 to 101, and in four out of the five pots a self-fertilised plant flowered first.

Canna warscewiczi.

In the three generations taken together the crossed were to the self-fertilised in height as 100 to 101; in the first generation the self-fertilised plants showed some tendency to flower first, and in the third generation they flowered first in nine out of the twelve pots.

SPECIES IN WHICH THE CROSSED AND SELF-FERTILISED PLANTS FLOWERED ALMOST SIMULTANEOUSLY.

Mimulus luteus (Sixth Generation).

The crossed plants were inferior in height and vigour to the self-fertilised plants, which all belonged to the new white-flowered tall variety, yet in only half the pots did the self-fertilised plants flower first, and in the other half the crossed plants.

Viscaria oculata.

The crossed plants were only a little taller than the self-fertilised (namely, as 100 to 97), but considerably more fertile, yet both lots flowered almost simultaneously.

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 tabacum (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 constitution, 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 constitution. 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. Analogous 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 assuming new external characters, as well as new constitutional 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 sexual organs of monoecious and polygamous plants,–in the structure of the circumferential 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 competition 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 competition 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 competition 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 competition 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 constitutional 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 constitutional 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 constitutions, 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 constitutional 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 constitutional 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 competition 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 constitutional 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 constitutional 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 constitution; and as the advantage from a cross depends on the plants which are crossed differing somewhat in constitution, 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 constitutions. (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 circumstances. 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 constitution 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. Self-sterile plants.
Causes of self-sterility.
The appearance of highly self-fertile varieties. Self-fertilisation apparently in some respects beneficial, independently of the assured production of seeds.
Relative weights and rates of germination of seeds from crossed and self-fertilised flowers.

The present chapter is devoted to the Fertility of plants, as influenced by cross-fertilisation and self-fertilisation. The subject consists of two distinct branches; firstly, the relative productiveness or fertility of flowers crossed with pollen from a distinct plant and with their own pollen, as shown by the proportional number of capsules which they produce, together with the number of the contained seeds. Secondly, the degree of innate fertility or sterility of the seedlings raised from crossed and self-fertilised seeds; such seedlings being of the same age, grown under the same conditions, and fertilised in the same manner. These two branches of the subject correspond with the two which have to be considered by any one treating of hybrid plants; namely, in the first place the comparative productiveness of a species when fertilised with pollen from a distinct species and with its own pollen; and in the second place, the fertility of its hybrid offspring. These two classes of cases do not always run parallel; thus some plants, as Gartner has shown, can be crossed with great ease, but yield excessively sterile hybrids; while others are crossed with extreme difficulty, but yield fairly fertile hybrids.

The natural order to follow in this chapter would have been first to consider the effects on the fertility of the parent-plants of crossing them, and of fertilising them with their own pollen; but as we have discussed in the two last chapters the relative height, weight, and constitutional vigour of crossed and self-fertilised plants–that is, of plants raised from crossed and self-fertilised seeds–it will be convenient here first to consider their relative fertility. The cases observed by me are given in Table 9/D, in which plants of crossed and self-fertilised parentage were left to fertilise themselves, being either crossed by insects or spontaneously self-fertilised. It should be observed that the results cannot be considered as fully trustworthy, for the fertility of a plant is a most variable element, depending on its age, health, nature of the soil, amount of water given, and temperature to which it is exposed. The number of the capsules produced and the number of the contained seeds, ought to have been ascertained on a large number of crossed and self-fertilised plants of the same age and treated in every respect alike. In these two latter respects my observations may be trusted, but a sufficient number of capsules were counted only in a few instances. The fertility, or as it may perhaps better be called the productiveness, of a plant depends on the number of capsules produced, and on the number of seeds which these contain. But from various causes, chiefly from the want of time, I was often compelled to rely on the number of the capsules alone. Nevertheless, in the more interesting cases, the seeds were also counted or weighed. The average number of seeds per capsule is a more valuable criterion of fertility than the number of capsules produced. This latter circumstance depends partly on the size of the plant; and we know that crossed plants are generally taller and heavier than the self-fertilised; but the difference in this respect is rarely sufficient to account for the difference in the number of the capsules produced. It need hardly be added that in Table 9/D the same number of crossed and self-fertilised plants are always compared. Subject to the foregoing sources of doubt I will now give the table, in which the parentage of the plants experimented on, and the manner of determining their fertility are explained. Fuller details may be found in the previous part of this work, under the head of each species.

TABLE 9/D.–RELATIVE FERTILITY OF PLANTS OF CROSSED AND SELF-FERTILISED PARENTAGE, BOTH SETS BEING FERTILISED IN THE SAME MANNER. FERTILITY JUDGED OF BY VARIOUS STANDARDS. THAT OF THE CROSSED PLANTS TAKEN AS 100.

Column 1: Name of plant and feature observed.

Column 2: x, in the expression, as 100 to x.

Ipomoea purpurea–first generation: seeds per capsule on crossed and self-fertilised plants, not growing much crowded, spontaneously self-fertilised under a net, in number: 99.

Ipomoea purpurea–seeds per capsule on crossed and self-fertilised plants from the same parents as in the last case, but growing much crowded, spontaneously self-fertilised under a net, in number: 93.

Ipomoea purpurea–productiveness of the same plants, as judged by the number of capsules produced, and average number of seeds per capsule: 45.

Ipomoea purpurea–third generation: seeds per capsule on crossed and self-fertilised plants, spontaneously self-fertilised under a net, in number: 94.

Ipomoea purpurea–productiveness of the same plants, as judged by the number of capsules produced, and the average number of seeds per capsule: 35.

Ipomoea purpurea–fifth generation: seeds per capsule on crossed and self-fertilised plants, left uncovered in the hothouse, and spontaneously fertilised: 89.

Ipomoea purpurea–ninth generation: number of capsules on crossed plants to those on self-fertilised plants, spontaneously self-fertilised under a net: 26.

Mimulus luteus–an equal number of capsules on plants descended from self-fertilised plants of the 8th generation crossed by a fresh stock, and on plants of the 9th self-fertilised generation, both sets having been left uncovered and spontaneously fertilised, contained seeds, by weight: 30.

Mimulus luteus–productiveness of the same plants, as judged by the number of capsules produced, and the average weight of seeds per capsule: 3.

Vandellia nummularifolia–seeds per capsule from cleistogene flowers on the crossed and self-fertilised plants, in number: 106.

Salvia coccinea–crossed plants, compared with self-fertilised plants, produced flowers, in number: 57.

Iberis umbellata–plants left uncovered in greenhouse; intercrossed plants of the 3rd generation, compared with self-fertilised plants of the 3rd generation, yielded seeds, in number: 75.

Iberis umbellata–plants from a cross between two varieties, compared with self-fertilised plants of the 3rd generation, yielded seeds, by weight : 75.

Papaver vagum–crossed and self-fertilised plants, left uncovered, produced capsules, in number: 99.

Eschscholtzia californica–Brazilian stock; plants left uncovered and cross-fertilised by bees; capsules on intercrossed plants of the 2nd generation, compared with capsules on self-fertilised plants of 2nd generation, contained seeds, in number: 78.

Eschscholtzia californica–productiveness of the same plants, as judged by the number of capsules produced, and the average number of seeds per capsule: 89.

Eschscholtzia californica–plants left uncovered and cross-fertilised by bees; capsules on plants derived from intercrossed plants of the 2nd generation of the Brazilian stock crossed by English stock, compared with capsules on self-fertilised plants of 2nd generation, contained seeds, in number: 63.

Eschscholtzia californica–productiveness of the same plants, as judged by the number of capsules produced, and the average number of seeds per capsule: 40.

Reseda odorata–crossed and self-fertilised plants, left uncovered and cross-fertilised by bees; produced capsules in number (about): 100.

Viola tricolor–crossed and self-fertilised plants, left uncovered and cross-fertilised by bees, produced capsules in number: 10.

Delphinium consolida–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number: 56.

Viscaria oculata–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number: 77.

Dianthus caryophyllus–plants spontaneously self-fertilised under a net; capsules on intercrossed and self-fertilised plants of the 3rd generation contained seeds in number: 125.

Dianthus caryophyllus–plants left uncovered and cross-fertilised by insects: offspring from plants self-fertilised for three generations and then crossed by an intercrossed plant of the same stock, compared with plants of the 4th self-fertilised generation, produced seeds by weight: 73.

Dianthus caryophyllus–plants left uncovered and cross-fertilised by insects: offspring from plants self-fertilised for three generations and then crossed by a fresh stock, compared with plants of the 4th self-fertilised generation, produced seeds by weight: 33.

Tropaeolum minus–crossed and self-fertilised plants, left uncovered in the greenhouse, produced seeds in number: 64.

Limnanthes douglasii–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number (about): 100.

Lupinus luteus–crossed and self-fertilised plants of the 2nd generation, left uncovered in the greenhouse, produced seeds in number (judged from only a few pods): 88.

Phaseolus multiflorus–crossed and self-fertilised plants, left uncovered in the greenhouse, produced seeds in number (about): 100.

Lathyrus odoratus–crossed and self-fertilised plants of the 2nd generation, left uncovered in the greenhouse, but certainly self-fertilised, produced pods in number: 91.

Clarkia elegans–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number: 60.

Nemophila insignis–crossed and self-fertilised plants, covered by a net and spontaneously self-fertilised in the greenhouse, produced capsules in number: 29.

Petunia violacea–left uncovered and cross-fertilised by insects: plants of the 5th intercrossed and self-fertilised generations produced seeds, as judged by the weight of an equal number of capsules: 86.

Petunia violacea–left uncovered as above: offspring of plants self-fertilised for four generations and then crossed by a fresh stock, compared with plants of the 5th self-fertilised generation, produced seeds, as judged by the weight of an equal number of capsules: 46.

Cyclamen persicum–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number: 12.

Anagallis collina–crossed and self-fertilised plants, left uncovered in the greenhouse, produced capsules in number: 8.

Primula veris–left uncovered in open ground and cross-fertilised by insects: offspring from plants of the 3rd illegitimate generation crossed by a fresh stock, compared with plants of the 4th illegitimate and self-fertilised generation, produced capsules in number: 5.

Same plants in the following year: 3.5.

Primula veris–(equal-styled variety): left uncovered in open ground and cross-fertilised by insects: offspring from plants self-fertilised for two generations and then crossed by another variety, compared with plants of the 3rd self-fertilised generation, produced capsules in number: 15.

Primula veris–(equal-styled variety) same plants; average number of seeds per capsule: 71.

Primula veris–(equal-styled variety) productiveness of the same plants, as judged by the number of capsules produced and the average number of seeds per capsule: 11.

This table includes thirty-three cases relating to twenty-three species, and shows the degree of innate fertility of plants of crossed parentage in comparison with those of self-fertilised parentage; both lots being fertilised in the same manner. With several of the species, as with Eschscholtzia, Reseda, Viola, Dianthus, Petunia, and Primula, both lots were certainly cross-fertilised by insects, and so it probably was with several of the others; but in some of the species, as with Nemophila, and in some of the trials with Ipomoea and Dianthus, the plants were covered up, and both lots were spontaneously self-fertilised. This also was necessarily the case with the capsules produced by the cleistogene flowers of Vandellia.

The fertility of the crossed plants is represented in Table 9/D by 100, and that of the self-fertilised by the other figures. There are five cases in which the fertility of the self-fertilised plants is approximately equal to that of the crossed; nevertheless, in four of these cases the crossed plants were plainly taller, and in the fifth somewhat taller than the self-fertilised. But I should state that in some of these five cases the fertility of the two lots was not strictly ascertained, as the capsules were not actually counted, from appearing equal in number and from all apparently containing a full complement of seeds. In only two instances in the table, namely, with Vandellia and in the third generation of Dianthus, the capsules on the self-fertilised plants contained more seed than those on the crossed plants. With Dianthus the ratio between the number of seeds contained in the self-fertilised and crossed capsules was as 125 to 100; both sets of plants were left to fertilise themselves under a net; and it is almost certain that the greater fertility of the self-fertilised plants was here due merely to their having varied and become less strictly dichogamous, so as to mature their anthers and stigmas more nearly at the same time than is proper to the species. Excluding the seven cases now referred to, there remain twenty-six in which the crossed plants were manifestly much more fertile, sometimes to an extraordinary degree, than the self-fertilised with which they grew in competition. The most striking instances are those in which plants derived from a cross with a fresh stock are compared with plants of one of the later self-fertilised generations; yet there are some striking cases, as that of Viola, between the intercrossed plants of the same stock and the self-fertilised, even in the first generation. The results most to be trusted are those in which the productiveness of the plants was ascertained by the number of capsules produced by an equal number of plants, together with the actual or average number of seeds in each capsule. Of such cases there are twelve in the table, and the mean of their mean fertility is as 100 for the crossed plants, to 59 for the self-fertilised plants. The Primulaceae seem eminently liable to suffer in fertility from self-fertilisation.

The following short table, Table 9/E, includes four cases which have already been partly given in the last table.

TABLE 9/E.–INNATE FERTILITY OF PLANTS FROM A CROSS WITH A FRESH STOCK, COMPARED WITH THAT OF INTERCROSSED PLANTS OF THE SAME STOCK, AND WITH THAT OF SELF-FERTILISED PLANTS, ALL OF THE CORRESPONDING GENERATION. FERTILITY JUDGED OF BY THE NUMBER OR WEIGHT OF SEEDS PRODUCED BY AN EQUAL NUMBER OF PLANTS.

Column 1: Name of plant and feature observed.

Column 2: Plants from a cross with a fresh stock.

Column 3: Intercrossed plants of the same stock.

Column 4: Self-fertilised plants.

Mimulus luteus–the intercrossed plants are derived from a cross between two plants of the 8th self-fertilised generation. The self-fertilised plants belong to the 9th generation: 100 : 4 : 3.

Eschscholtzia californica–the intercrossed and self-fertilised plants belong to the 2nd generation: 100 : 45 : 40.

Dianthus caryophyllus–the intercrossed plants are derived from self-fertilised of the 3rd generation, crossed by intercrossed plants of the 3rd generation. The self-fertilised plants belong to the 4th generation: 100 : 45 : 33.

Petunia violacea–the intercrossed and self-fertilised plants belong to the 5th generation: 100 : 54 : 46.

NB.–In the above cases, excepting in that of Eschscholtzia, the plants derived from a cross with a fresh stock belong on the mother-side to the same stock with the intercrossed and self-fertilised plants, and to the corresponding generation.

These cases show us how greatly superior in innate fertility the seedlings from plants self-fertilised or intercrossed for several generations and then crossed by a fresh stock are, in comparison with the seedlings from plants of the old stock, either intercrossed or self-fertilised for the same number of generations. The three lots of plants in each case were left freely exposed to the visits of insects, and their flowers without doubt were cross-fertilised by them.

Table 9/E further shows us that in all four cases the intercrossed plants of the same stock still have a decided though small advantage in fertility over the self-fertilised plants.

With respect to the state of the reproductive organs in the self-fertilised plants of Tables 9/D and 9/E, only a few observations were made. In the seventh and eighth generation of Ipomoea, the anthers in the flowers of the self-fertilised plants were plainly smaller than those in the flowers of the intercrossed plants. The tendency to sterility in these same plants was also shown by the first-formed flowers, after they had been carefully fertilised, often dropping off, in the same manner as frequently occurs with hybrids. The flowers likewise tended to be monstrous. In the fourth generation of Petunia, the pollen produced by the self-fertilised and intercrossed plants was compared, and they were far more empty and shrivelled grains in the former.

RELATIVE FERTILITY OF FLOWERS CROSSED WITH POLLEN FROM A DISTINCT PLANT AND WITH THEIR OWN POLLEN. THIS HEADING INCLUDES FLOWERS ON THE PARENT-PLANTS, AND ON THE CROSSED AND SELF-FERTILISED SEEDLINGS OF THE FIRST OR A SUCCEEDING GENERATION.

I will first treat of the parent-plants, which were raised from seeds purchased from nursery-gardens, or taken from plants growing in my garden, or growing wild, and surrounded in every case by many individuals of the same species. Plants thus circumstanced will commonly have been intercrossed by insects; so that the seedlings which were first experimented on will generally have been the product of a cross. Consequently any difference in the fertility of their flowers, when crossed and self-fertilised, will have been caused by the nature of the pollen employed; that is, whether it was taken from a distinct plant or from the same flower. The degrees of fertility shown in Table 9/F, were determined in each case by the average number of seeds per capsule, ascertained either by counting or weighing.

Another element ought properly to have been taken into account, namely, the proportion of flowers which yielded capsules when they were crossed and self-fertilised; and as crossed flowers generally produce a larger proportion of capsules, their superiority in fertility, if this element had been taken into account, would have been much more strongly marked than appears in Table 9/F. But had I thus acted, there would have been greater liability to error, as pollen applied to the stigma at the wrong time fails to produce any effect, independently of its greater or less potency. A good illustration of the great difference in the results which sometimes follows, if the number of capsules produced relatively to the number of flowers fertilised be included in the calculation, was afforded by Nolana prostrata. Thirty flowers on some plants of this species were crossed and produced twenty-seven capsules, each containing five seeds; thirty-two flowers on the same plants were self-fertilised and produced only six capsules, each containing five seeds. As the number of seeds per capsule is here the same, the fertility of the crossed and self-fertilised flowers is given in Table 9/F as equal, or as 100 to 100. But if the flowers which failed to produce capsules be included, the crossed flowers yielded on an average 4.50 seeds, whilst the self-fertilised flowers yielded only 0.94 seeds, so that their relative fertility would have been as 100 to 21. I should here state that it has been found convenient to reserve for separate discussion the cases of flowers which are usually quite sterile with their own pollen.

TABLE 9/f.–relative fertility of the flowers on the parent-plants used in my experiments, when fertilised with pollen from a distinct plant and with their own pollen. Fertility judged of by the average number of seeds per capsule. Fertility of crossed flowers taken as 100.

Column 1: Name of plant and feature observed.

Column 2: x, in the expression 100 to x.

Ipomoea purpurea–crossed and self-fertilised flowers yielded seeds as (about): 100.

Mimulus luteus–crossed and self-fertilised flowers yielded seeds as (by weight): 79.

Linaria vulgaris–crossed and self-fertilised flowers yielded seeds as: 14.

Vandellia nummularifolia–crossed and self-fertilised flowers yielded seeds as: 67?

Gesneria pendulina–crossed and self-fertilised flowers yielded seeds as (by weight): 100.

Salvia coccinea–crossed and self-fertilised flowers yielded seeds as (about): 100.

Brassica oleracea–crossed and self-fertilised flowers yielded seeds as: 25.

Eschscholtzia californica–(English stock) crossed and self-fertilised flowers yielded seeds as (by weight): 71.

Eschscholtzia californica–(Brazilian stock grown in England) crossed and self-fertilised flowers yielded seeds (by weight) as (about): 15.

Delphinium consolida–crossed and self-fertilised flowers (self-fertilised capsules spontaneously produced, but result supported by other evidence) yielded seeds as: 59.

Viscaria oculata–crossed and self-fertilised flowers yielded seeds as (by weight): 38.

Viscaria oculata–crossed and self-fertilised flowers (crossed capsules compared on following year with spontaneously self-fertilised capsules) yielded seeds as : 58.

Dianthus caryophyllus–crossed and self-fertilised flowers yielded seeds as: 92.

Tropaeolum minus–crossed and self-fertilised flowers yielded seeds as: 92.

Tropaeolum tricolorum–crossed and self-fertilised flowers yielded seeds as: 115. (9/1. Tropaeolum tricolorum and Cuphea purpurea have been introduced into this table, although seedlings were not raised from them; but of the Cuphea only six crossed and six self-fertilised capsules, and of the Tropaeolum only six crossed and eleven self-fertilised capsules, were compared. A larger proportion of the self-fertilised than of the crossed flowers of the Tropaeolum produced fruit.)

Limnanthes douglasii–crossed and self-fertilised flowers yielded seeds as (about): 100.

Sarothamnus scoparius–crossed and self-fertilised flowers yielded seeds as: 41.

Ononis minutissima–crossed and self-fertilised flowers yielded seeds as: 65.

Cuphea purpurea–crossed and self-fertilised flowers yielded seeds as: 113.

Passiflora gracilis–crossed and self-fertilised flowers yielded seeds as: 85.

Specularia speculum–crossed and self-fertilised flowers yielded seeds as: 72.

Lobelia fulgens–crossed and self-fertilised flowers yielded seeds as (about): 100.

Nemophila insignis–crossed and self-fertilised flowers yielded seeds as (by weight): 69.

Borago officinalis–crossed and self-fertilised flowers yielded seeds as: 60.

Nolana prostrata–crossed and self-fertilised flowers yielded seeds as: 100.

Petunia violacea–crossed and self-fertilised flowers yielded seeds as (by weight): 67.

Nicotiana tabacum–crossed and self-fertilised flowers yielded seeds as (by weight): 150.

Cyclamen persicum–crossed and self-fertilised flowers yielded seeds as: 38.

Anagallis collina–crossed and self-fertilised flowers yielded seeds as: 96.

Canna warscewiczi–crossed and self-fertilised flowers (on three generations of crossed and self-fertilised plants taken all together) yielded seeds as: 85.

Table 9/G gives the relative fertility of flowers on crossed plants again cross-fertilised, and of flowers on self-fertilised plants again self-fertilised, either in the first or in a later generation. Here two causes combine to diminish the fertility of the self-fertilised flowers; namely, the lesser efficacy of pollen from the same flower, and the innate lessened fertility of plants derived from self-fertilised seeds, which as we have seen in the previous Table 9/D is strongly marked. The fertility was determined in the same manner as in Table 9/F, that is, by the average number of seeds per capsule; and the same remarks as before, with respect to the different proportion of flowers which set capsules when they are cross-fertilised and self-fertilised, are here likewise applicable.

TABLE 9/G.–RELATIVE FERTILITY OF FLOWERS ON CROSSED AND SELF-FERTILISED PLANTS OF THE FIRST OR SOME SUCCEEDING GENERATION; THE FORMER BEING AGAIN FERTILISED WITH POLLEN FROM A DISTINCT PLANT, AND THE LATTER AGAIN WITH THEIR OWN POLLEN. FERTILITY JUDGED OF BY THE AVERAGE NUMBER OF SEEDS PER CAPSULE. FERTILITY OF CROSSED FLOWERS TAKEN AS 100.

Column 1: Name of plant and feature observed.

Column 2: x, in the expression, 100 to x.

Ipomoea purpurea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the first generation yielded seeds as: 93.

Ipomoea purpurea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 3rd generation yielded seeds as: 94.

Ipomoea purpurea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 4th generation yielded seeds as: 94.

Ipomoea purpurea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 5th generation yielded seeds as: 107.

Mimulus luteus–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 3rd generation yielded seeds as (by weight): 65.

Mimulus luteus–same plants of the 3rd generation treated in the same manner on the following year yielded seeds as (by weight): 34.

Mimulus luteus–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 4th generation yielded seeds as (by weight): 40.

Viola tricolor–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as: 69.

Dianthus caryophyllus–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as: 65.

Dianthus caryophyllus–flowers on self-fertilised plants of the 3rd generation crossed by intercrossed plants, and other flowers again self-fertilised yielded seeds as: 97.

Dianthus caryophyllus–flowers on self-fertilised plants of the 3rd generation crossed by a fresh stock, and other flowers again self-fertilised yielded seeds as: 127.

Lathytus odoratus–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as: 65.

Lobelia ramosa–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as (by weight): 60.

Petunia violacea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as (by weight): 68.

Petunia violacea–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 4th generation yielded seeds as (by weight): 72.

Petunia violacea–flowers on self-fertilised plants of the 4th generation crossed by a fresh stock, and other flowers again self-fertilised yielded seeds as (by weight): 48.

Nicotiana tabacum–crossed and self-fertilised flowers on the crossed and self-fertilised plants of the 1st generation yielded seeds as (by weight): 97.

Nicotiana tabacum–flowers on self-fertilised plants of the 2nd generation crossed by intercrossed plants, and other flowers again self-fertilised yielded seeds as (by estimation): 110.

Nicotiana tabacum–flowers on self-fertilised plants of the 3rd generation crossed by a fresh stock, and other flowers again self-fertilised yielded seeds as (by estimation): 110.

Anagallis collina–flowers on red variety crossed by a blue variety, and other flowers on the red variety self-fertilised yielded seeds as: 48.

Canna warscewiczi–crossed and self-fertilised flowers on the crossed and self-fertilised plants of three generations taken together yielded seeds as: 85.

As both these tables relate to the fertility of flowers fertilised by pollen from another plant and by their own pollen, they may be considered together. The difference between them consists in the self-fertilised flowers in Table 9/G, being produced by self-fertilised parents, and the crossed flowers by crossed parents, which in the later generations had become somewhat closely inter-related, and had been subjected all the time to nearly the same conditions. These two tables include fifty cases relating to thirty-two species. The flowers on many other species were crossed and self-fertilised, but as only a few were thus treated, the results cannot be trusted, as far as fertility is concerned, and are not here given. Some other cases have been rejected, as the plants were in an unhealthy condition. If we look to the figures in the two tables expressing the ratios between the mean relative fertility of the crossed and self-fertilised flowers, we see that in a majority of cases (i.e., in thirty-five out of fifty) flowers fertilised by pollen from a distinct plant yield more, sometimes many more, seeds than flowers fertilised with their own pollen; and they commonly set a larger proportion of capsules. The degree of infertility of the self-fertilised flowers differs extremely in the different species, and even, as we shall see in the section on self-sterile plants, in the individuals of the same species, as well as under slightly changed conditions of life. Their fertility ranges from zero to fertility equalling that of the crossed flowers; and of this fact no explanation can be offered. There are fifteen cases in the two tables in which the number of seeds per capsule produced by the self-fertilised flowers equals or even exceeds that yielded by the crossed flowers. Some few of these cases are, I believe, accidental; that is, would not recur on a second trial. This was apparently the case with the plants of the fifth generation of Ipomoea, and in one of the experiments with Dianthus. Nicotiana offers the most anomalous case of any, as the self-fertilised flowers on the parent-plants, and on their descendants of the second and third generations, produced more seeds than did the crossed flowers; but we shall recur to this case when we treat of highly self-fertile varieties.

It might have been expected that the difference in fertility between the crossed and self-fertilised flowers would have been more strongly marked in Table 9/G, in which the plants of one set were derived from self-fertilised parents, than in Table 9/F, in which flowers on the parent-plants were self-fertilised for the first time. But this is not the case, as far as my scanty materials allow of any judgment. There is therefore no evidence at present, that the fertility of plants goes on diminishing in successive self-fertilised generations, although there is some rather weak evidence that this does occur with respect to their height or growth. But we should bear in mind that in the later generations the crossed plants had become more or less closely inter-related, and had been subjected all the time to nearly uniform conditions.

It is remarkable that there is no close correspondence, either in the parent-plants or in the successive generations, between the relative number of seeds produced by the crossed and self-fertilised flowers, and the relative powers of growth of the seedlings raised from such seeds. Thus, the crossed and self-fertilised flowers on the parent-plants of Ipomoea, Gesneria, Salvia, Limnanthes, Lobelia fulgens, and Nolana produced a nearly equal number of seeds, yet the plants raised from the crossed seeds exceeded considerably in height those raised from the self-fertilised seeds. The crossed flowers of Linaria and Viscaria yielded far more seeds than the self-fertilised flowers; and although the plants raised from the former were taller than those from the latter, they were not so in any corresponding degree. With Nicotiana the flowers fertilised with their own pollen were more productive than those crossed with pollen from a slightly different variety; yet the plants raised from the latter seeds were much taller, heavier, and more hardy than those raised from the self-fertilised seeds. On the other hand, the crossed seedlings of Eschscholtzia were neither taller nor heavier than the self-fertilised, although the crossed flowers were far more productive than the self-fertilised. But the best evidence of a want of correspondence between the number of seeds produced by crossed and self-fertilised flowers, and the vigour of the offspring raised from them, is afforded by the plants of the Brazilian and European stocks of Eschscholtzia, and likewise by certain individual plants of Reseda odorata; for it might have been expected that the seedlings from plants, the flowers of which were excessively self-sterile, would have profited in a greater degree by a cross, than the seedlings from plants which were moderately or fully self-fertile, and therefore apparently had no need to be crossed. But no such result followed in either case: for instance, the crossed and self-fertilised offspring from a highly self-fertile plant of Reseda odorata were in average height to each other as 100 to 82; whereas the similar offspring from an excessively self-sterile plant were as 100 to 92 in average height.

With respect to the innate fertility of the plants of crossed and self-fertilised parentage, given in the previous Table 9/D–that is, the number of seeds produced by both lots when their flowers were fertilised in the same manner,–nearly the same remarks are applicable, in reference to the absence of any close correspondence between their fertility and powers of growth, as in the case of the plants in the Tables 9/F and 9/G, just considered. Thus the crossed and self-fertilised plants of Ipomoea, Papaver, Reseda odorata, and Limnanthes were almost equally fertile, yet the former exceeded considerably in height the self-fertilised plants. On the other hand, the crossed and self-fertilised plants of Mimulus and Primula differed to an extreme degree in innate fertility, but by no means to a corresponding degree in height or vigour.

In all the cases of self-fertilised flowers included in Tables 9/E, 9/F, and 9/G, these were fertilised with their own pollen; but there is another form of self-fertilisation, namely, by pollen from other flowers on the same plant; but this latter method made no difference in comparison with the former in the number of seeds produced, or only a slight difference. Neither with Digitalis nor Dianthus were more seeds produced by the one method than by the other, to any trustworthy degree. With Ipomoea rather more seeds, in the proportion of 100 to 91, were produced from a crossed between flowers on the same plant than from strictly self-fertilised flowers; but I have reason to suspect that the result was accidental. With Origanum vulgare, however, a cross between flowers on plants propagated by stolons from the same stock certainly increased slightly their fertility. This likewise occurred, as we shall see in the next section, with Eschscholtzia, perhaps with Corydalis cava and Oncidium; but not so with Bignonia, Abutilon, Tabernaemontana, Senecio, and apparently Reseda odorata.

SELF-STERILE PLANTS.

The cases here to be described might have been introduced in Table 9/F, which gives the relative fertility of flowers fertilised with their own pollen, and with that from a distinct plant, but it has been found more convenient to keep them for separate discussion. The present cases must not be confounded with those to be given in the next chapter relatively to flowers which are sterile when insects are excluded; for such sterility depends not merely on the flowers being incapable of fertilisation with their own pollen, but on mechanical causes, by which their pollen is prevented from reaching the stigma, or on the pollen and stigma of the same flower being matured at different periods.

In the seventeenth chapter of my ‘Variation of Animals and Plants under Domestication’ I had occasion to enter fully on the present subject; and I will therefore here give only a brief abstract of the cases there described, but others must be added, as they have an important bearing on the present work. Kolreuter long ago described plants of Verbascum phoeniceum which during two years were sterile with their own pollen, but were easily fertilised by that of four other species; these plants however afterwards became more or less self-fertile in a strangely fluctuating manner. Mr. Scott also found that this species, as well as two of its varieties, were self-sterile, as did Gartner in the case of Verbascum nigrum. So it was, according to this latter author, with two plants of Lobelia fulgens, though the pollen and ovules of both were in an efficient state in relation to other species. Five species of Passiflora and certain individuals of a sixth species have been found sterile with their own pollen; but slight changes in their conditions, such as being grafted on another stock or a change of temperature, rendered them self-fertile. Flowers on a completely self-impotent plant of Passiflora alata fertilised with pollen from its own self-impotent seedlings were quite fertile. Mr. Scott, and afterwards Mr. Munro, found that some species of Oncidium and of Maxillaria cultivated in a hothouse in Edinburgh were quite sterile with their own pollen; and Fritz Muller found this to be the case with a large number of Orchidaceous genera growing in their native home of South Brazil. (9/2. ‘Botanische Zeitung’ 1868 page 114.) He also discovered that the pollen-masses of some orchids acted on their own stigmas like a poison; and it appears that Gartner formerly observed indications of this extraordinary fact in the case of some other plants.

Fritz Muller also states that a species of Bignonia and Tabernaemontana echinata are both sterile with their own pollen in their native country of Brazil. (9/3. Ibid 1868 page 626 and 1870 page 274.) Several Amaryllidaceous and Liliaceous plants are in the same predicament. Hildebrand observed with care Corydalis cava, and found it completely self-sterile (9/4. ‘Report of the International Horticultural Congress’ 1866.); but according to Caspary a few self-fertilised seeds are occasionally produced: Corydalis halleri is only slightly self-sterile, and C. intermedia not at all so. (9/5. ‘Botanische Zeitung’ June 27, 1873.) In another Fumariaceous genus, Hypecoum, Hildebrand observed that H. grandiflorum was highly self-sterile, whilst H. procumbens was fairly self-fertile. (9/6. ‘Jahrb. fur wiss. Botanik’ B. 7 page 464.) Thunbergia alata kept by me in a warm greenhouse was self-sterile early in the season, but at a later period produced many spontaneously self-fertilised fruits. So it was with Papaver vagum: another species, P. alpinum, was found by Professor H. Hoffmann to be quite self-sterile excepting on one occasion (9/7. ‘Zur Speciesfrage’ 1875 page 47.); whilst P. somniferum has been with me always completely self-sterile.

Eschscholtzia californica.

This species deserves a fuller consideration. A plant cultivated by Fritz Muller in South Brazil happened to flower a month before any of the others, and it did not produce a single capsule. This led him to make further observations during the next six generations, and he found that all his plants were completely sterile, unless they were crossed by insects or were artificially fertilised with pollen from a distinct plant, in which case they were completely fertile. (9/8. ‘Botanische Zeitung’ 1868 page 115 and 1869 page 223.) I was much surprised at this fact, as I had found that English plants, when covered by a net, set a considerable number of capsules; and that these contained seeds by weight, compared with those on plants intercrossed by the bees, as 71 to 100. Professor Hildebrand, however, found this species much more self-sterile in Germany than it was with me in England, for the capsules produced by self-fertilised flowers, compared with those from intercrossed flowers, contained seeds in the ratio of only 11 to 100. At my request Fritz Muller sent me from Brazil seeds of his self-sterile plants, from which I raised seedlings. Two of these were covered with a net, and one produced spontaneously only a single capsule containing no good seeds, but yet, when artificially fertilised with its own pollen, produced a few capsules. The other plant produced spontaneously under the net eight capsules, one of which contained no less than thirty seeds, and on an average about ten seeds per capsule. Eight flowers on these two plants were artificially self-fertilised, and produced seven capsules, containing on an average twelve seeds; eight other flowers were fertilised with pollen from a distinct plant of the Brazilian stock, and produced eight capsules, containing on an average about eighty seeds: this gives a ratio of 15 seeds for the self-fertilised capsules to 100 for the crossed capsules. Later in the season twelve other flowers on these two plants were artificially self-fertilised; but they yielded only two capsules, containing three and six seeds. It appears therefore that a lower temperature than that of Brazil favours the self-fertility of this plant, whilst a still lower temperature lessens it. As soon as the two plants which had been covered by the net were uncovered, they were visited by many bees,and it was interesting to observe how quickly they became, even the more sterile plant of the two, covered with young capsules. On the following year eight flowers on plants of the Brazilian stock of self-fertilised parentage (i.e., grandchildren of the plants which grew in Brazil) were again self-fertilised, and produced five capsules, containing on an average 27.4 seeds, with a maximum in one of forty-two seeds; so that their self-fertility had evidently increased greatly by being reared for two generations in England. On the whole we may conclude that plants of the Brazilian stock are much more self-fertile in this country than in Brazil, and less so than plants of the English stock in England; so that the plants of Brazilian parentage retained by inheritance some of their former sexual constitution. Conversely, seeds from English plants sent by me to Fritz Muller and grown in Brazil, were much more self-fertile than his plants which had been cultivated there for several generations; but he informs me that one of the plants of English parentage which did not flower the first year, and was thus exposed for two seasons to the climate of Brazil, proved quite self-sterile, like a Brazilian plant, showing how quickly the climate had acted on its sexual constitution.

Abutilon darwinii.

Seeds of this plant were sent me by Fritz Muller, who found it, as well as some other species of the same genus, quite sterile in its native home of South Brazil, unless fertilised with pollen from a distinct plant, either artificially or naturally by humming-birds. (9/9. ‘Jenaische Zeitschr. fur Naturwiss’ B. 7 1872 page 22 and 1873 page 441.) Several plants were raised from these seeds and kept in the hothouse. They produced flowers very early in the spring, and twenty of them were fertilised, some with pollen from the same flower, and some with pollen from other flowers on the same plants; but not a single capsule was thus produced, yet the stigmas twenty-seven hours after the application of the pollen were penetrated by the pollen-tubes. At the same time nineteen flowers were crossed with pollen from a distinct plant, and these produced thirteen capsules, all abounding with fine seeds. A greater number of capsules would have been produced by the cross, had not some of the nineteen flowers been on a plant which was afterwards proved to be from some unknown cause completely sterile with pollen of any kind. Thus far these plants behaved exactly like those in Brazil; but later in the season, in the latter part of May and in June, they began to produce under a net a few spontaneously self-fertilised capsules. As soon as this occurred, sixteen flowers were fertilised with their own pollen, and these produced five capsules, containing on an average 3.4 seeds. At the same time I selected by chance four capsules from the uncovered plants growing close by, the flowers of which I had seen visited by humble-bees, and these contained on an average 21.5 seeds; so that the seeds in the naturally intercrossed capsules to those in the self-fertilised capsules were as 100 to 16. The interesting point in this case is that these plants, which were unnaturally treated by being grown in pots in a hothouse, under another hemisphere, with a complete reversal of the seasons, were thus rendered slightly self-fertile, whereas they seem always to be completely self-sterile in their native home.

Senecio cruentus (greenhouse varieties, commonly called Cinerarias, probably derived from several fruticose or herbaceous species much intercrossed (9/10. I am much obliged to Mr. Moore and to Mr. Thiselton Dyer for giving me information with respect to the varieties on which I experimented. Mr. Moore believes that Senecio cruentas, tussilaginis, and perhaps heritieri, maderensis and populifolius have all been more or less blended together in our Cinerarias.))

Two purple-flowered varieties were placed under a net in the greenhouse, and four corymbs on each were repeatedly brushed with flowers from the other plant, so that their stigmas were well covered with each other’s pollen. Two of the eight corymbs thus treated produced very few seeds, but the other six produced on an average 41.3 seeds per corymb, and these germinated well. The stigmas on four other corymbs on both plants were well smeared with pollen from the flowers on their own corymbs; these eight corymbs produced altogether ten extremely poor seeds, which proved incapable of germinating. I examined many flowers on both plants, and found the stigmas spontaneously covered with pollen; but they produced not a single seed. These plants were afterwards left uncovered in the same house where many other Cinerarias were in flower; and the flowers were frequently visited by bees. They then produced plenty of seed, but one of the two plants less than the other, as this species shows some tendency to be dioecious.

The trial was repeated on another variety with white petals tipped with red. Many stigmas on two corymbs were covered with pollen from the foregoing purple variety, and these produced eleven and twenty-two seeds, which germinated well. A large number of the stigmas on several of the other corymbs were repeatedly smeared with pollen from their own corymb; but they yielded only five very poor seeds, which were incapable of germination. Therefore the above three plants belonging to two varieties, though growing vigorously and fertile with pollen from either of the other two plants, were utterly sterile with pollen from other flowers on the same plant.

Reseda odorata.

Having observed that certain individuals were self-sterile, I covered during the summer of 1868 seven plants under separate nets, and will call these plants A, B, C, D, E, F, G. They all appeared to be quite sterile with their own pollen, but fertile with that of any other plant.

Fourteen flowers on A were crossed with pollen from B or C, and produced thirteen fine capsules. Sixteen flowers were fertilised with pollen from other flowers on the same plant, but yielded not a single capsule.

Fourteen flowers on B were crossed with pollen from A, C or D, and all produced capsules; some of these were not very fine, yet they contained plenty of seeds. Eighteen flowers were fertilised with pollen from other flowers on the same plant, and produced not one capsule.

Ten flowers on C were crossed with pollen from A, B, D or E, and produced nine fine capsules. Nineteen flowers were fertilised with pollen from other flowers on the same plant, and produced no capsules.

Ten flowers on D were crossed with pollen from A, B, C or E, and produced nine fine capsules. Eighteen flowers were fertilised with pollen from other flowers on the same plant, and produced no capsules.

Seven flowers on E were crossed with pollen from A, C, or D, and all produced fine capsules. Eight flowers were fertilised with pollen from other flowers on the same plant, and produced no capsules.

On the plants F and G no flowers were crossed, but very many (number not recorded) were fertilised with pollen from other flowers on the same plants, and these did not produce a single capsule.

We thus see that fifty-five flowers on five of the above plants were reciprocally crossed in various ways; several flowers on each of these plants being fertilised with pollen from several of the other plants. These fifty-five flowers produced fifty-two capsules, almost all of which were of full size and contained an abundance of seeds. On the other hand, seventy-nine flowers (besides many others not recorded) were fertilised with pollen from other flowers on the same plants, and these did not produce a single capsule. In one case in which I examined the stigmas of the flowers fertilised with their own pollen, these were penetrated by the pollen-tubes, although such penetration produced no effect. Pollen falls generally, and I believe always, from the anthers on the stigmas of the same flower; yet only three out of the above seven protected plants produced spontaneously any capsules, and these it might have been thought must have been self-fertilised. There were altogether seven such capsules; but as they were all seated close to the artificially crossed flowers, I can hardly doubt that a few grains of foreign pollen had accidentally fallen on their stigmas. Besides the above seven plants, four others were kept covered under the SAME large net; and some of these produced here and there in the most capricious manner little groups of capsules; and this makes me believe that a bee, many of which settled on the outside of the net, being attracted by the odour, had on some one occasion found an entrance, and had intercrossed a few of the flowers.

In the spring of 1869 four plants raised from fresh seeds were carefully protected under separate nets; and now the result was widely different to what it was before. Three of these protected plants became actually loaded with capsules, especially during the early part of the summer; and this fact indicates that temperature produces some effect, but the experiment given in the following paragraph shows that the innate constitution of the plant is a far more important element. The fourth plant produced only a few capsules, many of them of small size; yet it was far more self-fertile than any of the seven plants tried during the previous year. The flowers on four small branches of this semi-self-sterile plant were smeared with pollen from one of the other plants, and they all produced fine capsules.

As I was much surprised at the difference in the results of the trials made during the two previous years, six fresh plants were protected by separate nets in the year 1870. Two of these proved almost completely self-sterile, for on carefully searching them I found only three small capsules, each containing either one or two seeds of small size, which, however, germinated. A few flowers on both these plants were reciprocally fertilised with each other’s pollen, and a few with pollen from one of the following self-fertile plants, and all these flowers produced fine capsules. The four other plants whilst still remaining protected beneath the nets presented a wonderful contrast (though one of them in a somewhat less degree than the others), for they became actually covered with spontaneously self-fertilised capsules, as numerous as, or very nearly so, and as fine as those on the unprotected plants growing near.

The above three spontaneously self-fertilised capsules produced by the two almost completely self-sterile plants, contained altogether five seeds; and from these I raised in the following year (1871) five plants, which were kept under separate nets. They grew to an extraordinarily large size, and on August 29th were examined. At first sight they appeared entirely destitute of capsules; but on carefully searching their many branches, two or three capsules were found on three of the plants, half-a-dozen on the fourth, and about eighteen on the fifth plant. But all these capsules were small, some being empty; the greater number contained only a single seed, and very rarely more than one. After this examination the nets were taken off, and the bees immediately carried pollen from one of these almost self-sterile plants to the other, for no other plants grew near. After a few weeks the ends of the branches on all five plants became covered with capsules, presenting a curious contrast with the lower and naked parts of the same long branches. These five plants therefore inherited almost exactly the same sexual constitution as their parents; and without doubt a self-sterile race of Mignonette could have been easily established.

Reseda lutea.

Plants of this species were raised from seeds gathered from a group of wild plants growing at no great distance from my garden. After casually observing that some of these plants were self-sterile, two plants taken by hazard were protected under separate nets. One of these soon became covered with spontaneously self-fertilised capsules, as numerous as those on the surrounding unprotected plants; so that it was evidently quite self-fertile. The other plant was partially self-sterile, producing very few capsules, many of which were of small size. When, however, this plant had grown tall, the uppermost branches became pressed against the net and grew crooked, and in this position the bees were able to suck the flowers through the meshes, and brought pollen to them from the neighbouring plants. These branches then became loaded with capsules; the other and lower branches remaining almost bare. The sexual constitution of this species is therefore similar to that of Reseda odorata.

CONCLUDING REMARKS ON SELF-STERILE PLANTS.

In order to favour as far as possible the self-fertilisation of some of the foregoing plants, all the flowers on Reseda odorata and some of those on the Abutilon were fertilised with pollen from other flowers on the same plant, instead of with their own pollen, and in the case of the Senecio with pollen from other flowers on the same corymb; but this made no difference in the result. Fritz Muller tried both kinds of self-fertilisation in the case of Bignonia, Tabernaemontana and Abutilon, likewise with no difference in the result. With Eschscholtzia, however, he found that pollen from other flowers on the same plant was a little more effective than pollen from the same flower. So did Hildebrand in Germany; as thirteen out of fourteen flowers of Eschscholtzia thus fertilised set capsules, these containing on an average 9.5 seeds; whereas only fourteen flowers out of twenty-one fertilised with their own pollen set capsules, these containing on an average 9.0 seeds. (9/11. ‘Pringsheim’s Jahrbuch fur wiss. Botanik’ 7 page 467.) Hildebrand found a trace of a similar difference with Corydalis cava, as did Fritz Muller with an Oncidium. (9/12. ‘Variation under Domestication’ chapter 17 2nd edition volume 2 pages 113-115.)

In considering the several cases above given of complete or almost complete self-sterility, we are first struck with their wide distribution throughout the vegetable kingdom. Their number is not at present large, for they can be discovered only by protecting plants from insects and then fertilising them with pollen from another plant of the same species and with their own pollen; and the latter must be proved to be in an efficient state by other trials. Unless all this be done, it is impossible to know whether their self-sterility may not be due to the male or female reproductive organs, or to both, having been affected by changed conditions of life. As in the course of my experiments I have found three new cases, and as Fritz Muller has observed indications of several others, it is probable that they will hereafter be proved to be far from rare. (9/13. Mr. Wilder, the editor of a horticultural journal in the United States quoted in ‘Gardeners’ Chronicle’ 1868 page 1286, states that Lilium auratum, Impatiens pallida and fulva, and Forsythia viridissima, cannot be fertilised with their own pollen.)

As with plants of the same species and parentage, some individuals are self-sterile and others self-fertile, of which fact Reseda odorata offers the most striking instances, it is not at all surprising that species of the same genus differ in this same manner. Thus Verbascum phoeniceum and nigrum are self-sterile, whilst V. thapsus and lychnitis are quite self-fertile, as I know by trial. There is the same difference between some of the species of Papaver, Corydalis, and of other genera. Nevertheless, the tendency to self-sterility certainly runs to a certain extent in groups, as we see in the genus Passiflora, and with the Vandeae amongst Orchids.

Self-sterility differs much in degree in different plants. In those extraordinary cases in which pollen from the same flower acts on the stigma like a poison, it is almost certain that the plants would never yield a single self-fertilised seed. Other plants, like Corydalis cava, occasionally, though very rarely, produce a few self-fertilised seeds. A large number of species, as may be seen in Table 9/F, are less fertile with their own pollen than with that from another plant; and lastly, some species are perfectly self-fertile. Even with the individuals of the same species, as just remarked, some are utterly self-sterile, others moderately so, and some perfectly self-fertile. The cause, whatever it may be, which renders many plants more or less sterile with their own pollen, that is, when they are self-fertilised, must be different, at least to a certain extent, from that which determines the difference in height, vigour, and fertility of the seedlings raised from self-fertilised and crossed seeds; for we have already seen that the two classes of cases do not by any means run parallel. This want of parallelism would be intelligible, if it could be shown that self-sterility depended solely on the incapacity of the pollen-tubes to penetrate the stigma of the same flower deeply enough to reach the ovules; whilst the greater or less vigorous growth of the seedlings no doubt depends on the nature of the contents of the pollen-grains and ovules. Now it is certain that with some plants the stigmatic secretion does not properly excite the pollen-grains, so that the tubes are not properly developed, if the pollen is taken from the same flower. This is the case according to Fritz Muller with Eschscholtzia, for he found that the pollen-tubes did not penetrate the stigma deeply; and with the Orchidaceous genus Notylia they failed altogether to penetrate it. (9/14. ‘Botanische Zeitung’ 1868 pages 114, 115.)

With dimorphic and trimorphic species, an illegitimate union between plants of the same form presents the closest analogy with self-fertilisation, whilst a legitimate union closely resembles cross-fertilisation; and here again the lessened fertility or complete sterility of an illegitimate union depends, at least in part, on the incapacity for interaction between the pollen-grains and stigma. Thus with Linum grandiflorum, as I have elsewhere shown, not more than two or three out of hundreds of pollen-grains, either of the long-styled or short-styled form, when placed on the stigma of their own form, emit their tubes, and these do not penetrate deeply; nor does the stigma itself change colour, as occurs when it is legitimately fertilised. (9/15. ‘Journal of the Linnean Society Botany’ volume 7 1863 pages 73-75.)

On the other hand the difference in innate fertility, as well as in growth between plants raised from crossed and self-fertilised seeds, and the difference in fertility and growth between the legitimate and illegitimate offspring of dimorphic and trimorphic plants, must depend on some incompatibility between the sexual elements contained within the pollen-grains and ovules, as it is through their union that new organisms are developed.

If we now turn to the more immediate cause of self-sterility, we clearly see that in most cases it is determined by the conditions to which the plants have been subjected. Thus Eschscholtzia is completely self-sterile in the hot climate of Brazil, but is perfectly fertile there with the pollen of any other individual. The offspring of Brazilian plants became in England in a single generation partially self-fertile, and still more so in the second generation. Conversely, the offspring of English plants, after growing for two seasons in Brazil, became in the first generation quite self-sterile. Again, Abutilon darwinii, which is self-sterile in its native home of Brazil, became moderately self-fertile in a single generation in an English hothouse. Some other plants are self-sterile during the early part of the year, and later in the season become self-fertile. Passiflora alata lost its self-sterility when grafted on another species. With Reseda, however, in which some individuals of the same parentage are self-sterile and others are self-fertile, we are forced in our ignorance to speak of the cause as due to spontaneous variability; but we should remember that the progenitors of these plants, either on the male or female side, may have been exposed to somewhat different conditions. The power of the environment thus to affect so readily and in so peculiar a manner the reproductive organs, is a fact which has many important bearings; and I have therefore thought the foregoing details worth giving. For instance, the sterility of many animals and plants under changed conditions of life, such as confinement, evidently comes within the same general principle of the sexual system being easily affected by the environment. It has already been proved, that a cross between plants which have been self-fertilised or intercrossed during several generations, having been kept all the time under closely similar conditions, does not benefit the offspring; and on the other hand, that a cross between plants that have been subjected to different conditions benefits the offspring to an extraordinary degree. We may therefore conclude that some degree of differentiation in the sexual system is necessary for the full fertility of the parent-plants and for the full vigour of their offspring. It seems also probable that with those plants which are capable of complete self-fertilisation, the male and female elements and organs already differ to an extent sufficient to excite their mutual interaction; but that when such plants are taken to another country, and become in consequence self-sterile, their sexual elements and organs are so acted on as to be rendered too uniform for such interaction, like those of a self-fertilised plant long cultivated under the same conditions. Conversely, we may further infer that plants which are self-sterile in their native country, but become self-fertile under changed conditions, have their sexual elements so acted on, that they become sufficiently differentiated for mutual interaction.

We know that self-fertilised seedlings are inferior in many respects to those from a cross; and as with plants in a state of nature pollen from the same flower can hardly fail to be often left by insects or by the wind on the stigma, it seems at first sight highly probable that self-sterility has been gradually acquired through natural selection in order to prevent self-fertilisation. It is no valid objection to this belief that the structure of some flowers, and the dichogamous condition of many others, suffice to prevent the pollen reaching the stigma of the same flower; for we should remember that with most species many flowers expand at the same time, and that pollen from the same plant is equally injurious or nearly so as that from the same flower. Nevertheless, the belief that self-sterility is a quality which has been gradually acquired for the special purpose of preventing self-fertilisation must, I believe, be rejected. In the first place, there is no close correspondence in degree between the sterility of the parent-plants when self-fertilised, and the extent to which their offspring suffer in vigour by this process; and some such correspondence might have been expected if self-sterility had been acquired on account of the injury caused by self-fertilisation. The fact of individuals of the same parentage differing greatly in their degree of self-sterility is likewise opposed to such a belief; unless, indeed, we suppose that certain individuals have been rendered self-sterile to favour intercrossing, whilst other individuals have been rendered self-fertile to ensure the propagation of the species. The fact of self-sterile individuals appearing only occasionally, as in the case of Lobelia, does not countenance this latter view. But the strongest argument against the belief that self-sterility has been acquired to prevent self-fertilisation, is the immediate and powerful effect of changed conditions in either causing or in removing self-sterility. We are not therefore justified in admitting that this peculiar state of the reproductive system has been gradually acquired through natural selection; but we must look at it as an incidental result, dependent on the conditions to which the plants have been subjected, like the ordinary sterility caused in the case of animals by confinement, and in the case of plants by too much manure, heat, etc. I do not, however, wish to maintain that self-sterility may not sometimes be of service to a plant in preventing self-fertilisation; but there are so many other means by which this result might be prevented or rendered difficult,