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

3. Bra.s.sica oleracea.

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

4. Iberis umbellata.

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

5. Eschscholtzia californica.

This plant offers an almost unique case, inasmuch as the good effects of a cross are confined to the reproductive system. Intercrossed and self-fertilised plants of the English stock did not differ in height (nor in weight, as far as was ascertained) in any constant manner; the self-fertilised plants usually having the advantage. So it was with the offspring of plants of the Brazilian stock, tried in the same manner.

The parent-plants, however, of the English stock produced many more seeds when fertilised with pollen from another plant than when self-fertilised; and in Brazil the parent-plants were absolutely sterile unless they were fertilised with pollen from another plant. Intercrossed seedlings, raised in England from the Brazilian stock, compared with self-fertilised seedlings of the corresponding second generation, yielded seeds in number as 100 to 89; both lots of plants being left freely exposed to the visits of insects. If we now turn to the effects of crossing plants of the Brazilian stock with pollen from the English stock,--so that plants which had been long exposed to very different conditions were intercrossed,--we find that the offspring were, as before, inferior in height and weight to the plants of the Brazilian stock after two generations of self-fertilisation, but were superior to them in the most marked manner in the number of seeds produced, namely, as 100 to 40; both lots of plants being left freely exposed to the visits of insects.

In the case of Ipomoea, we have seen that the plants derived from a cross with a fresh stock were superior in height as 100 to 78, and in fertility as 100 to 51, to the plants of the old stock, although these had been intercrossed during the last ten generations. With Eschscholtzia we have a nearly parallel case, but only as far as fertility is concerned, for the plants derived from a cross with a fresh stock were superior in fertility in the ratio of 100 to 45 to the Brazilian plants, which had been artificially intercrossed in England for the two last generations, and which must have been naturally intercrossed by insects during all previous generations in Brazil, where otherwise they are quite sterile.

6. Dianthus caryophyllus.

Plants self-fertilised for three generations were crossed with pollen from a fresh stock, and their offspring were grown in compet.i.tion with plants of the fourth self-fertilised generation. The crossed plants thus obtained were to the self-fertilised in height as 100 to 81, and in fertility (both lots being left to be naturally fertilised by insects) as 100 to 33.

These same crossed plants were also to the offspring from the plants of the third generation crossed by the intercrossed plants of the corresponding generation, in height as 100 to 85, and in fertility as 100 to 45.

We thus see what a great advantage the offspring from a cross with a fresh stock had, not only over the self-fertilised plants of the fourth generation, but over the offspring from the self-fertilised plants of the third generation, when crossed by the intercrossed plants of the old stock.

7. Pisum sativum.

It has been shown under the head of this species, that the several varieties in this country almost invariably fertilise themselves, owing to insects rarely visiting the flowers; and as the plants have been long cultivated under nearly similar conditions, we can understand why a cross between two individuals of the same variety does not do the least good to the offspring either in height or fertility. This case is almost exactly parallel with that of Mimulus, or that of the Ipomoea named Hero; for in these two instances, crossing plants which had been self-fertilised for seven generations did not at all benefit the offspring. On the other hand, a cross between two varieties of the pea causes a marked superiority in the growth and vigour of the offspring, over the self-fertilised plants of the same varieties, as shown by two excellent observers. From my own observations (not made with great care) the offspring from crossed varieties were to self-fertilised plants in height, in one case as 100 to about 75, and in a second case as 100 to 60.

8. Lathyrus odoratus.

The sweet-pea is in the same state in regard to self-fertilisation as the common pea; and we have seen that seedlings from a cross between two varieties, which differed in no respect except in the colour of their flowers, were to the self-fertilised seedlings from the same mother-plant in height as 100 to 80; and in the second generation as 100 to 88. Unfortunately I did not ascertain whether crossing two plants of the same variety failed to produce any beneficial effect, but I venture to predict such would be the result.

9. Petunia violacea.

The intercrossed plants of the same stock in four out of the five successive generations plainly exceeded in height the self-fertilised plants. The latter in the fourth generation were crossed by a fresh stock, and the seedlings thus obtained were put into compet.i.tion with the self-fertilised plants of the fifth generation. The crossed plants exceeded the self-fertilised in height in the ratio of 100 to 66, and in weight as 100 to 23; but this difference, though so great, is not much greater than that between the intercrossed plants of the same stock in comparison with the self-fertilised plants of the corresponding generation. This case, therefore, seems at first sight opposed to the rule that a cross with a fresh stock is much more beneficial than a cross between individuals of the same stock. But as with Eschscholtzia, the reproductive system was here chiefly benefited; for the plants raised from the cross with the fresh stock were to the self-fertilised plants in fertility, both lots being naturally fertilised, as 100 to 46, whereas the intercrossed plants of the same stock were to the self-fertilised plants of the corresponding fifth generation in fertility only as 100 to 86.

Although at the time of measurement the plants raised from the cross with the fresh stock did not exceed in height or weight the intercrossed plants of the old stock (owing to the growth of the former not having been completed, as explained under the head of this species), yet they exceeded the intercrossed plants in fertility in the ratio of 100 to 54.

This fact is interesting, as it shows that plants self-fertilised for four generations and then crossed by a fresh stock, yielded seedlings which were nearly twice as fertile as those from plants of the same stock which had been intercrossed for the five previous generations. We here see, as with Eschscholtzia and Dianthus, that the mere act of crossing, independently of the state of the crossed plants, has little efficacy in giving increased fertility to the offspring. The same conclusion holds good, as we have already seen, in the a.n.a.logous cases of Ipomoea, Mimulus, and Dianthus, with respect to height.

10. Nicotiana tabac.u.m.

My plants were remarkably self-fertile, and the capsules from the self-fertilised flowers apparently yielded more seeds than those which were cross-fertilised. No insects were seen to visit the flowers in the hothouse, and I suspect that the stock on which I experimented had been raised under gla.s.s, and had been self-fertilised during several previous generations; if so, we can understand why, in the course of three generations, the crossed seedlings of the same stock did not uniformly exceed in height the self-fertilised seedlings. But the case is complicated by individual plants having different const.i.tutions, so that some of the crossed and self-fertilised seedlings raised at the same time from the same parents behaved differently. However this may be, plants raised from self-fertilised plants of the third generation crossed by a slightly different sub-variety, exceeded greatly in height and weight the self-fertilised plants of the fourth generation; and the trial was made on a large scale. They exceeded them in height when grown in pots, and not much crowded, in the ratio of 100 to 66; and when much crowded, as 100 to 54. These crossed plants, when thus subjected to severe compet.i.tion, also exceeded the self-fertilised in weight in the ratio of 100 to 37. So it was, but in a less degree (as may be seen in Table 7/C), when the two lots were grown out of doors and not subjected to any mutual compet.i.tion. Nevertheless, strange as is the fact, the flowers on the mother-plants of the third self-fertilised generation did not yield more seed when they were crossed with pollen from plants of the fresh stock than when they were self-fertilised.

11. Anagallis collina.

Plants raised from a red variety crossed by another plant of the same variety were in height to the self-fertilised plants from the red variety as 100 to 73. When the flowers on the red variety were fertilised with pollen from a closely similar blue-flowered variety, they yielded double the number of seeds to what they did when crossed by pollen from another individual of the same red variety, and the seeds were much finer. The plants raised from this cross between the two varieties were to the self-fertilised seedlings from the red variety, in height as 100 to 66, and in fertility as 100 to 6.

12. Primula veris.

Some flowers on long-styled plants of the third illegitimate generation were legitimately crossed with pollen from a fresh stock, and others were fertilised with their own pollen. From the seeds thus produced crossed plants, and self-fertilised plants of the fourth illegitimate generation, were raised. The former were to the latter in height as 100 to 46, and in fertility during one year as 100 to 5, and as 100 to 3.5 during the next year. In this case, however, we have no means of distinguishing between the evil effects of illegitimate fertilisation continued during four generations (that is, by pollen of the same form, but taken from a distinct plant) and strict self-fertilisation. But it is probable that these two processes do not differ so essentially as at first appears to be the case. In the following experiment any doubt arising from illegitimate fertilisation was completely eliminated.

13. Primula veris. (Equal-styled, red-flowered variety.)

Flowers on plants of the second self-fertilised generation were crossed with pollen from a distinct variety or fresh stock, and others were again self-fertilised. Crossed plants and plants of the third self-fertilised generation, all of legitimate origin, were thus raised; and the former was to the latter in height as 100 to 85, and in fertility (as judged by the number of capsules produced, together with the average number of seeds) as 100 to 11.

SUMMARY OF THE MEASUREMENTS IN TABLE 7/C.

This table includes the heights and often the weights of 292 plants derived from a cross with a fresh stock, and of 305 plants, either of self-fertilised origin, or derived from an intercross between plants of the same stock. These 597 plants belong to thirteen species and twelve genera. The various precautions which were taken to ensure a fair comparison have already been stated. If we now look down the right hand column, in which the mean height, weight, and fertility of the plants derived from a cross with a fresh stock are represented by 100, we shall see by the other figures how wonderfully superior they are both to the self-fertilised and to the intercrossed plants of the same stock. With respect to height and weight, there are only two exceptions to the rule, namely, with Eschscholtzia and Petunia, and the latter is probably no real exception. Nor do these two species offer an exception in regard to fertility, for the plants derived from the cross with a fresh stock were much more fertile than the self-fertilised plants. The difference between the two sets of plants in the table is generally much greater in fertility than in height or weight. On the other hand, with some of the species, as with Nicotiana, there was no difference in fertility between the two sets, although a great difference in height and weight.

Considering all the cases in this table, there can be no doubt that plants profit immensely, though in different ways, by a cross with a fresh stock or with a distinct sub-variety. It cannot be maintained that the benefit thus derived is due merely to the plants of the fresh stock being perfectly healthy, whilst those which had been long intercrossed or self-fertilised had become unhealthy; for in most cases there was no appearance of such unhealthiness, and we shall see under Table 7/A that the intercrossed plants of the same stock are generally superior to a certain extent to the self-fertilised,--both lots having been subjected to exactly the same conditions and being equally healthy or unhealthy.

We further learn from Table 7/C, that a cross between plants that have been self-fertilised during several successive generations and kept all the time under nearly uniform conditions, does not benefit the offspring in the least or only in a very slight degree. Mimulus and the descendants of Ipomoea named Hero offer instances of this rule. Again, plants self-fertilised during several generations profit only to a small extent by a cross with intercrossed plants of the same stock (as in the case of Dianthus), in comparison with the effects of a cross by a fresh stock. Plants of the same stock intercrossed during several generations (as with Petunia) were inferior in a marked manner in fertility to those derived from the corresponding self-fertilised plants crossed by a fresh stock. Lastly, certain plants which are regularly intercrossed by insects in a state of nature, and which were artificially crossed in each succeeding generation in the course of my experiments, so that they can never or most rarely have suffered any evil from self-fertilisation (as with Eschscholtzia and Ipomoea), nevertheless profited greatly by a cross with a fresh stock. These several cases taken together show us in the clearest manner that it is not the mere crossing of any two individuals which is beneficial to the offspring. The benefit thus derived depends on the plants which are united differing in some manner, and there can hardly be a doubt that it is in the const.i.tution or nature of the s.e.xual elements. Anyhow, it is certain that the differences are not of an external nature, for two plants which resemble each other as closely as the individuals of the same species ever do, profit in the plainest manner when intercrossed, if their progenitors have been exposed during several generations to different conditions. But to this latter subject I shall have to recur in a future chapter.

TABLE 7/A.

We will now turn to our first table, which relates to crossed and self-fertilised plants of the same stock. These consist of fifty-four species belonging to thirty natural orders. The total number of crossed plants of which measurements are given is 796, and of self-fertilised 809; that is altogether 1,605 plants. Some of the species were experimented on during several successive generations; and it should be borne in mind that in such cases the crossed plants in each generation were crossed with pollen from another crossed plant, and the flowers on the self-fertilised plants were almost always fertilised with their own pollen, though sometimes with pollen from other flowers on the same plant. The crossed plants thus became more or less closely inter-related in the later generations; and both lots were subjected in each generation to almost absolutely the same conditions, and to nearly the same conditions in the successive generations. It would have been a better plan in some respects if I had always crossed some flowers either on the self-fertilised or intercrossed plants of each generation with pollen from a non-related plant, grown under different conditions, as was done with the plants in Table 7/C; for by this procedure I should have learnt how much the offspring became deteriorated through continued self-fertilisation in the successive generations. As the case stands, the self-fertilised plants of the successive generations in Table 7/A were put into compet.i.tion with and compared with intercrossed plants, which were probably deteriorated in some degree by being more or less inter-related and grown under similar conditions. Nevertheless, had I always followed the plan in Table 7/C, I should not have discovered the important fact that, although a cross between plants which are rather closely related and which had been subjected to closely similar conditions, gives during several generations some advantage to the offspring, yet that after a time they may be intercrossed with no advantage whatever to the offspring. Nor should I have learnt that the self-fertilised plants of the later generations might be crossed with intercrossed plants of the same stock with little or no advantage, although they profited to an extraordinary degree by a cross with a fresh stock.

With respect to the greater number of the plants in Table 7/A, nothing special need here be said; full particulars may be found under the head of each species by the aid of the Index. The figures in the right-hand column show the mean height of the self-fertilised plants, that of the crossed plants with which they competed being represented by 100. No notice is here taken of the few cases in which crossed and self-fertilised plants were grown in the open ground, so as not to compete together. The table includes, as we have seen, plants belonging to fifty-four species, but as some of these were measured during several successive generations, there are eighty-three cases in which crossed and self-fertilised plants were compared. As in each generation the number of plants which were measured (given in the table) was never very large and sometimes small, whenever in the right hand column the mean height of the crossed and self-fertilised plants is the same within five per cent, their heights may be considered as practically equal. Of such cases, that is, of self-fertilised plants of which the mean height is expressed by figures between 95 and 105, there are eighteen, either in some one or all the generations. There are eight cases in which the self-fertilised plants exceed the crossed by above five per cent, as shown by the figures in the right hand column being above 105. Lastly, there are fifty-seven cases in which the crossed plants exceed the self-fertilised in a ratio of at least 100 to 95, and generally in a much higher degree.

If the relative heights of the crossed and self-fertilised plants had been due to mere chance, there would have been about as many cases of self-fertilised plants exceeding the crossed in height by above five per cent as of the crossed thus exceeding the self-fertilised; but we see that of the latter there are fifty-seven cases, and of the former only eight cases; so that the cases in which the crossed plants exceed in height the self-fertilised in the above proportion are more than seven times as numerous as those in which the self-fertilised exceed the crossed in the same proportion. For our special purpose of comparing the powers of growth of crossed and self-fertilised plants, it may be said that in fifty-seven cases the crossed plants exceeded the self-fertilised by more than five per cent, and that in twenty-six cases (18 + 8) they did not thus exceed them. But we shall now show that in several of these twenty-six cases the crossed plants had a decided advantage over the self-fertilised in other respects, though not in height; that in other cases the mean heights are not trustworthy, owing to too few plants having been measured, or to their having grown unequally from being unhealthy, or to both causes combined.

Nevertheless, as these cases are opposed to my general conclusion I have felt bound to give them. Lastly, the cause of the crossed plants having no advantage over the self-fertilised can be explained in some other cases. Thus a very small residue is left in which the self-fertilised plants appear, as far as my experiments serve, to be really equal or superior to the crossed plants.

We will now consider in some little detail the eighteen cases in which the self-fertilised plants equalled in average height the crossed plants within five per cent; and the eight cases in which the self-fertilised plants exceeded in average height the crossed plants by above five per cent; making altogether twenty-six cases in which the crossed plants were not taller than the self-fertilised plants in any marked degree.

[1. Dianthus caryophyllus (third generation).

This plant was experimented on during four generations, in three of which the crossed plants exceeded in height the self-fertilised generally by much more than five per cent; and we have seen under Table 7/C that the offspring from the plants of the third self-fertilised generation crossed by a fresh stock profited in height and fertility to an extraordinary degree. But in this third generation the crossed plants of the same stock were in height to the self-fertilised only as 100 to 99, that is, they were practically equal. Nevertheless, when the eight crossed and eight self-fertilised plants were cut down and weighed, the former were to the latter in weight as 100 to 49! There can therefore be not the least doubt that the crossed plants of this species are greatly superior in vigour and luxuriance to the self-fertilised; and what was the cause of the self-fertilised plants of the third generation, though so light and thin, growing up so as almost to equal the crossed in height, I cannot explain.

2. Lobelia fulgens (first generation).

The crossed plants of this generation were much inferior in height to the self-fertilised, in the proportion of 100 to 127. Although only two pairs were measured, which is obviously much too few to be trusted, yet from other evidence given under the head of this species, it is certain that the self-fertilised plants were very much more vigorous than the crossed. As I used pollen of unequal maturity for crossing and self-fertilising the parent-plants, it is possible that the great difference in the growth of their offspring may have been due to this cause. In the next generation this source of error was avoided, and many more plants were raised, and now the average height of the twenty-three crossed plants was to that of the twenty-three self-fertilised plants as 100 to 91. We can therefore hardly doubt that a cross is beneficial to this species.

3. Petunia violacea (third generation).

Eight crossed plants were to eight self-fertilised of the third generation in average height as 100 to 131; and at an early age the crossed were inferior even in a still higher degree. But it is a remarkable fact that in one pot in which plants of both lots grew extremely crowded, the crossed were thrice as tall as the self-fertilised. As in the two preceding and two succeeding generations, as well as with plants raised by a crossed with a fresh stock, the crossed greatly exceeded the self-fertilised in height, weight, and fertility (when these two latter points were attended to), the present case must be looked at as an anomaly not affecting the general rule. The most probable explanation is that the seeds from which the crossed plants of the third generation were raised were not well ripened; for I have observed an a.n.a.logous case with Iberis. Self-fertilised seedlings of this latter plant, which were known to have been produced from seeds not well matured, grew from the first much more quickly than the crossed plants, which were raised from better matured seeds; so that having thus once got a great start they were enabled ever afterwards to retain their advantage. Some of these same seeds of the Iberis were sown on the opposite sides of pots filled with burnt earth and pure sand, not containing any organic matter; and now the young crossed seedlings grew during their short life to double the height of the self-fertilised, in the same manner as occurred with the above two sets of seedlings of Petunia which were much crowded and thus exposed to very unfavourable conditions. We have seen also in the eighth generation of Ipomoea that the self-fertilised seedlings raised from unhealthy parents grew at first very much more quickly than the crossed seedlings, so that they were for a long time much taller, though ultimately beaten by them.

4, 5, 6. Eschscholtzia californica.

Four sets of measurements are given in Table 7/A. In one of these the crossed plants exceed the self-fertilised in average height, so that this is not one of the exceptions here to be considered. In two other cases the crossed equalled the self-fertilised in height within five per cent; and in the fourth case the self-fertilised exceeded the crossed by above this limit. We have seen in Table 7/C that the whole advantage of a cross by a fresh stock is confined to fertility, and so it was with the intercrossed plants of the same stock compared with the self-fertilised, for the former were in fertility to the latter as 100 to 89. The intercrossed plants thus have at least one important advantage over the self-fertilised. Moreover, the flowers on the parent-plants when fertilised with pollen from another individual of the same stock yield far more seeds than when self-fertilised; the flowers in this latter case being often quite sterile. We may therefore conclude that a cross does some good, though it does not give to the crossed seedlings increased powers of growth.

7. Viscaria oculata.

The average height of the fifteen intercrossed plants to that of the fifteen self-fertilised plants was only as 100 to 97; but the former produced many more capsules than the latter, in the ratio of 100 to 77.

Moreover, the flowers on the parent-plants which were crossed and self-fertilised, yielded seeds on one occasion in the proportion of 100 to 38, and on a second occasion in the proportion of 100 to 58. So that there can be no doubt about the beneficial effects of a cross, although the mean height of the crossed plants was only three per cent above that of the self-fertilised plants.

8. Specularia speculum.

Only the four tallest of the crossed and the four tallest of the self-fertilised plants, growing in four pots, were measured; and the former were to the latter in height as 100 to 98. In all four pots a crossed plant flowered before any one of the self-fertilised plants, and this is usually a safe indication of some real superiority in the crossed plants. The flowers on the parent-plants which were crossed with pollen from another plant yielded seeds compared with the self-fertilised flowers in the ratio of 100 to 72. We may therefore draw the same conclusion as in the last case with respect to a cross being decidedly beneficial.

9. Borago officinalis.