The Different Forms of Flowers on Plants of the Same Species
by Charles Darwin
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When, however, any one form of Oxalis is fertilised illegitimately with pollen from the same form, the seedlings appear to belong invariably to this form. Thus Hildebrand states that long-styled plants of O. rosea growing by themselves have been propagated in Germany year after year by seed, and have always produced long-styled plants. (5/6. 'Ueber den Trimorphismus in der Gattung Oxalis: Monatsberichte der Akad. der Wissen. zu Berlin' 21 June 1866 page 373 and 'Botanische Zeitung' 1871 page 435.) Again, 17 seedlings were raised from mid- styled plants of O. hedysaroides growing by themselves, and these were all mid- styled. So that the forms of Oxalis, when illegitimately fertilised with their own pollen, behave like the long-styled form of Lythrum salicaria, which when thus fertilised always produced with me long-styled offspring.]


Primula Sinensis.

I raised during February 1862, from some long-styled plants illegitimately fertilised with pollen from the same form, twenty-seven seedlings. These were all long-styled. They proved fully fertile or even fertile in excess; for ten flowers, fertilised with pollen from other plants of the same lot, yielded nine capsules, containing on an average 39.75 seeds, with a maximum in one capsule of 66 seeds. Four other flowers legitimately crossed with pollen from a legitimate plant, and four flowers on the latter crossed with pollen from the illegitimate seedlings, yielded seven capsules with an average of 53 seeds, with a maximum of 72. I must here state that I have found some difficulty in estimating the normal standard of fertility for the several unions of this species, as the results differ much during successive years, and the seeds vary so greatly in size that it is hard to decide which ought to be considered good. In order to avoid over- estimating the infertility of the several illegitimate unions, I have taken the normal standard as low as possible.

From the foregoing twenty-seven illegitimate plants, fertilised with their own- form pollen, twenty-five seedling grandchildren were raised; and these were all long-styled; so that from the two illegitimate generations fifty-two plants were raised, and all without exception proved long-styled. These grandchildren grew vigorously, and soon exceeded in height two other lots of illegitimate seedlings of different parentage and one lot of equal-styled seedlings presently to be described. Hence I expected that they would have turned out highly ornamental plants; but when they flowered, they seemed, as my gardener remarked, to have gone back to the wild state; for the petals were pale-coloured, narrow, sometimes not touching each other, flat, generally deeply notched in the middle, but not flexuous on the margin, and with the yellow eye or centre conspicuous. Altogether these flowers were strikingly different from those of their progenitors; and this I think, can only be accounted for on the principle of reversion. Most of the anthers on one plant were contabescent. Seventeen flowers on the grandchildren were illegitimately fertilised with pollen taken from other seedlings of the same lot, and produced fourteen capsules, containing on an average 29.2 seeds; but they ought to have contained about 35 seeds. Fifteen flowers legitimately fertilised with pollen from an illegitimate short-styled plant (belonging to the lot next to be described) produced fourteen capsules, containing an average of 46 seeds; they ought to have contained at least 50 seeds. Hence these grandchildren of illegitimate descent appear to have lost, though only in a very slight degree, their full fertility.

We will now turn to the short-styled form: from a plant of this kind, fertilised with its own-form pollen, I raised, during February 1862, eight seedlings, seven of which were short-styled and one long-styled. They grew slowly, and never attained to the full stature of ordinary plants; some of them flowered precociously, and others late in the season. Four flowers on these short-styled seedlings and four on the one long-styled seedling were illegitimately fertilised with their own-form pollen and produced only three capsules, containing on an average 23.6 seeds, with a maximum of 29; but we cannot judge of their fertility from so few capsules; and I have greater doubts about the normal standard for this union than about any other; but I believe that rather above 25 seeds would be a fair estimate. Eight flowers on these same short- styled plants, and the one long-styled illegitimate plant were reciprocally and legitimately crossed; they produced five capsules, which contained an average of 28.6 seeds, with a maximum of 36. A reciprocal cross between legitimate plants of the two forms would have yielded an average of at least 57 seeds, with a possible maximum of 74 seeds; so that these illegitimate plants were sterile when legitimately crossed.

I succeeded in raising from the above seven short-styled illegitimate plants, fertilised with their own-form pollen, only six plants—grandchildren of the first union. These, like their parents, were of low stature, and had so poor a constitution that four died before flowering. With ordinary plants it has been a rare event with me to have more than a single plant die out of a large lot. The two grandchildren which lived and flowered were short-styled; and twelve of their flowers were fertilised with their own-form pollen and produced twelve capsules containing an average of 28.2 seeds; so that these two plants, though belonging to so weakly a set, were rather more fertile than their parents, and perhaps not in any degree sterile. Four flowers on the same two grandchildren were legitimately fertilised by a long-styled illegitimate plant, and produced four capsules, containing only 32.2 seeds instead of about 64 seeds, which is the normal average for legitimate short-styled plants legitimately crossed.

By looking back, it will be seen that I raised at first from a short-styled plant fertilised with its own-form pollen one long-styled and seven short-styled illegitimate seedlings. These seedlings were legitimately intercrossed, and from their seed fifteen plants were raised, grandchildren of the first illegitimate union, and to my surprise all proved short-styled. Twelve short-styled flowers borne by these grandchildren were illegitimately fertilised with pollen taken from other plants of the same lot, and produced eight capsules which contained an average of 21.8 seeds, with a maximum of 35. These figures are rather below the normal standard for such a union. Six flowers were also legitimately fertilised with pollen from an illegitimate long-styled plant and produced only three capsules, containing on an average 23.6 seeds, with a maximum of 35. Such a union in the case of a legitimate plant ought to have yielded an average of 64 seeds, with a possible maximum of 73 seeds.


In regard to the long-styled plants, their illegitimate offspring, of which fifty-two were raised in the course of two generations, were all long-styled. (5/7. Dr. Hildebrand, who first called attention to this subject 'Botanische Zeitung' 1864 page 5, raised from a similar illegitimate union seventeen plants, of which fourteen were long-styled and three short-styled. From a short-styled plant illegitimately fertilised with its own pollen he raised fourteen plants, of which eleven were short-styled and three long-styled.) These plants grew vigorously; but the flowers in one instance were small, appearing as if they had reverted to the wild state. In the first illegitimate generation they were perfectly fertile, and in the second their fertility was only very slightly impaired. With respect to the short-styled plants, twenty-four out of twenty- five of their illegitimate offspring were short-styled. They were dwarfed in stature, and one lot of grandchildren had so poor a constitution that four out of six plants perished before flowering. The two survivors, when illegitimately fertilised with their own-form pollen, were rather less fertile than they ought to have been; but their loss of fertility was clearly shown in a special and unexpected manner, namely, when legitimately fertilised by other illegitimate plants: thus altogether eighteen flowers were fertilised in this manner, and yielded twelve capsules, which included on an average only 28.5 seeds, with a maximum of 45. Now a legitimate short-styled plant would have yielded, when legitimately fertilised, an average of 64 seeds, with a possible maximum of 74. This particular kind of infertility will perhaps be best appreciated by a simile: we may assume that with mankind six children would be born on an average from an ordinary marriage; but that only three would be born from an incestuous marriage. According to the analogy of Primula Sinensis, the children of such incestuous marriages, if they continued to marry incestuously, would have their sterility only slightly increased; but their fertility would not be restored by a proper marriage; for if two children, both of incestuous origin, but in no degree related to each other, were to marry, the marriage would of course be strictly legitimate, nevertheless they would not give birth to more than half the full and proper number of children.


As any variation in the structure of the reproductive organs, combined with changed function, is a rare event, the following cases are worth giving in detail. My attention was first called to the subject by observing, in 1862, a long-styled plant, descended from a self-fertilised long-styled parent, which had some of its flowers in an anomalous state, namely, with the stamens placed low down in the corolla as in the ordinary long-styled form, but with the pistils so short that the stigmas stood on a level with the anthers. These stigmas were nearly as globular and as smooth as in the short-styled form, instead of being elongated and rough as in the long-styled form. Here, then, we have combined in the same flower, the short stamens of the long-styled form with a pistil closely resembling that of the short-styled form. But the structure varied much even on the same umbel: for in two flowers the pistil was intermediate in length between that of the long and that of the short-styled form, with the stigma elongated as in the former, and smooth as in the latter; and in three other flowers the structure was in all respects like that of the long-styled form. These modifications appeared to me so remarkable that I fertilised eight of the flowers with their own pollen, and obtained five capsules, which contained on an average 43 seeds; and this number shows that the flowers had become abnormally fertile in comparison with those of ordinary long- styled plants when self-fertilised. I was thus led to examine the plants in several small collections, and the result showed that the equal-styled variety was not rare.

TABLE 5.31. Primula Sinensis. Preponderance of long-styled over the short-styled form.

Column 1: Name of owner or place. Column 2: Long-styled form. Column 3: Short-styled form. Column 4: equal-styled variety.

Mr. Horwood : 0 : 0 : 17. Mr. Duck : 20 : 0 : 9. Baston : 30 : 18 : 15. Chichester : 12 : 9 : 2. Holwood : 42 : 12 : 0. High Elms : 16 : 0 : 0. Westerham : 1 : 5 : 0. My own plants from purchased seeds : 13 : 7 : 0. Total : 134 : 51 : 43.

In a state of nature the long and short-styled forms would no doubt occur in nearly equal numbers, as I infer from the analogy of the other heterostyled species of Primula, and from having raised the two forms of the present species in exactly the same number from flowers which had been LEGITIMATELY crossed. The preponderance in Table 5.31 of the long-styled form over the short-styled (in the proportion of 134 to 51) results from gardeners generally collecting seed from self-fertilised flowers; and the long-styled flowers produce spontaneously much more seed (as shown in the first chapter) than the short-styled, owing to the anthers of the long-styled form being placed low down in the corolla, so that, when the flowers fall off, the anthers are dragged over the stigma; and we now also know that long-styled plants, when self-fertilised, very generally reproduce long-styled offspring. From the consideration of this table, it occurred to me in the year 1862, that almost all the plants of the Chinese primrose cultivated in England would sooner or later become long-styled or equal-styled; and now, at the close of 1876, I have had five small collections of plants examined, and almost all consisted of long-styled, with some more or less well-characterised equal-styled plants, but with not one short-styled.

With respect to the equal-styled plants in the table, Mr. Horwood raised from purchased seeds four plants, which he remembered were certainly not long-styled, but either short or equal-styled, probably the latter. These four plants were kept separate and allowed to fertilise themselves; from their seed the seventeen plants in the table were raised, all of which proved equal-styled. The stamens stood low down in the corolla as in the long-styled form; and the stigmas, which were globular and smooth, were either completely surrounded by the anthers, or stood close above them. My son William made drawings for me, by the aid of the camera, of the pollen of one of the above equal-styled plants; and, in accordance with the position of the stamens, the grains resembled in their small size those of the long-styled form. He also examined pollen from two equal- styled plants at Southampton; and in both of them the grains differed extremely in size in the same anthers, a large number being small and shrivelled, whilst many were fully as large as those of the short-styled form and rather more globular. It is probable that the large size of these grains was due, not to their having assumed the character of the short-styled form, but to monstrosity; for Max Wichura has observed pollen-grains of monstrous size in certain hybrids. The vast number of the small shrivelled grains in the above two cases explains the fact that, though equal-styled plants are generally fertile in a high degree, yet some of them yield few seeds. I may add that my son compared, in 1875, the grains from two white-flowered plants, in both of which the pistil projected above the anthers, but neither were properly long-styled or equal- styled; and in the one in which the stigma projected most, the grains were in diameter to those in the other plant, in which the stigma projected less, as 100 to 88; whereas the difference between the grains from perfectly characterised long-styled and short-styled plants is as 100 to 57. So that these two plants were in an intermediate condition. To return to the 17 plants in the first line of Table 5.31: from the relative position of their stigmas and anthers, they could hardly fail to fertilise themselves; and accordingly four of them spontaneously yielded no less than 180 capsules; of these Mr. Horwood selected eight fine capsules for sowing; and they included on an average 54.8 seeds, with a maximum of 72. He gave me thirty other capsules, taken by hazard, of which twenty-seven contained good seeds, averaging 35.5, with a maximum of 70; but if six poor capsules, each with less than 13 seeds, be excluded, the average rises to 42.5. These are higher numbers than could be expected from either well- characterised form if self-fertilised; and this high degree of fertility accords with the view that the male organs belonged to one form, and the female organs partially to the other form; so that a self-union in the case of the equal- styled variety is in fact a legitimate union.

The seed saved from the above seventeen self-fertilised equal-styled plants produced sixteen plants, which all proved equal-styled, and resembled their parents in all the above-specified respects. The stamens, however, in one plant were seated higher up the tube of the corolla than in the true long-styled form; in another plant almost all the anthers were contabescent. These sixteen plants were the grandchildren of the four original plants, which it is believed were equal-styled; so that this abnormal condition was faithfully transmitted, probably through three, and certainly through two generations. The fertility of one of these grandchildren was carefully observed: six flowers were fertilised with pollen from the same flower, and produced six capsules, containing on an average 68 seeds, with a maximum of 82, and a minimum of 40. Thirteen capsules spontaneously self-fertilised yielded an average of 53.2 seeds, with the astonishing maximum in one of 97 seeds. In no legitimate union has so high an average as 68 seeds been observed by me, or nearly so high a maximum as 82 and 97. These plants, therefore, not only have lost their proper heterostyled structure and peculiar functional powers, but have acquired an abnormal grade of fertility—unless, indeed, their high fertility may be accounted for by the stigmas receiving pollen from the circumjacent anthers at exactly the most favourable period.

With respect to Mr. Duck's lot in Table 5.31, seed was saved from a single plant, of which the form was not observed, and this produced nine equal-styled and twenty long-styled plants. The equal-styled resembled in all respects those previously described; and eight of their capsules spontaneously self-fertilised contained on an average 44.4 seeds, with a maximum of 61 and a minimum of 23. In regard to the twenty long-styled plants, the pistil in some of the flowers did not project quite so high as in ordinary long-styled flowers; and the stigmas, though properly elongated, were smooth; so that we have here a slight approach in structure to the pistil of the short-styled form. Some of these long-styled plants also approached the equal-styled in function; for one of them produced no less than fifteen spontaneously self-fertilised capsules, and of these eight contained, on an average, 31.7 seeds, with a maximum of 61. This average would be rather low for a long-styled plant artificially fertilised with its own pollen, but is high for one spontaneously self-fertilised. For instance, thirty- four capsules produced by the illegitimate grandchildren of a long-styled plant, spontaneously self-fertilised, contained on an average only 9.1 seeds, with a maximum of 46. Some seeds indiscriminately saved from the foregoing twenty-nine equal-styled and long-styled plants produced sixteen seedlings, grandchildren of the original plant belonging to Mr. Duck; and these consisted of fourteen equal- styled and two long-styled plants; and I mention this fact as an additional instance of the transmission of the equal-styled variety.

The third lot in Table 5.31, namely the Baston plants, are the last which need be mentioned. The long and short-styled plants, and the fifteen equal-styled plants, were descended from two distinct stocks. The latter were derived from a single plant, which the gardener is positive was not long-styled; hence, probably, it was equal-styled. In all these fifteen plants the anthers, occupying the same position as in the long-styled form, closely surrounded the stigma, which in one instance alone was slightly elongated. Notwithstanding this position of the stigma, the flowers, as the gardener assured me, did not yield many seeds; and this difference from the foregoing cases may perhaps have been caused by the pollen being bad, as in some of the Southampton equal-styled plants.]


That this is a variation, and not a third or distinct form, as in the trimorphic genera Lythrum and Oxalis, is clear; for we have seen its first appearance in one out of a lot of illegitimate long-styled plants; and in the case of Mr. Duck's seedlings, long-styled plants, only slightly deviating from the normal state, as well as equal-styled plants were produced from the same self- fertilised parent. The position of the stamens in their proper place low down in the tube of the corolla, together with the small size of the pollen-grains, show, firstly, that the equal-styled variety is a modification of the long- styled form, and, secondly, that the pistil is the part which has varied most, as indeed was obvious in many of the plants. This variation is of frequent occurrence, and is strongly inherited when it has once appeared. It would, however, have possessed little interest if it had consisted of a mere change of structure; but this is accompanied by modified fertility. Its occurrence apparently stands in close relation with the illegitimate birth of the parent plant; but to this whole subject I shall hereafter recur.

[Primula auricula.

Although I made no experiments on the illegitimate offspring of this species, I refer to it for two reasons:—First, because I have observed two equal-styled plants in which the pistil resembled in all respects that of the long-styled form, whilst the stamens had become elongated as in the short-styled form, so that the stigma was almost surrounded by the anthers. The pollen-grains, however, of the elongated stamens resembled in their small size those of the shorter stamens proper to the long-styled form. Hence these plants have become equal-styled by the increased length of the stamens, instead of, as with P. Sinensis, by the diminished length of the pistil. Mr. J. Scott observed five other plants in the same state, and he shows that one of them, when self- fertilised, yielded more seed than an ordinary long- or short-styled form would have done when similarly fertilised, but that it was far inferior in fertility to either form when legitimately crossed. (5/8. 'Journal of the Proceedings of the Linnean Society' 8 1864 page 91.) Hence it appears that the male and female organs of this equal-styled variety have been modified in some special manner, not only in structure but in functional powers. This, moreover, is shown by the singular fact that both the long-styled and short-styled plants, fertilised with pollen from the equal-styled variety, yield a lower average of seed than when these two forms are fertilised with their own pollen.

The second point which deserves notice is that florists always throw away the long-styled plants, and save seed exclusively from the short-styled form. Nevertheless, as Mr. Scott was informed by a man who raises this species extensively in Scotland, about one-fourth of the seedlings appear long-styled; so that the short-styled form of the Auricula, when fertilised by its own pollen, does not reproduce the same form in so large a proportion as in the case of P. Sinensis. We may further infer that the short-styled form is not rendered quite sterile by a long course of fertilisation with pollen of the same form: but as there would always be some liability to an occasional cross with the other form, we cannot tell how long self-fertilisation has been continued.

Primula farinosa.

Mr. Scott says that it is not at all uncommon to find equal-styled plants of this heterostyled species. (5/9. 'Journal of the Proceedings of the Linnean Society' 8 1864 page 115.) Judging from the size of the pollen-grains, these plants owe their structure, as in the case of P. auricula, to the abnormal elongation of the stamens of the long-styled form. In accordance with this view, they yield less seed when crossed with the long-styled form than with the short- styled. But they differ in an anomalous manner from the equal-styled plants of P. auricula in being extremely sterile with their own pollen.

Primula elatior.

It was shown in the first chapter, on the authority of Herr Breitenbach, that equal-styled flowers are occasionally found on this species whilst growing in a state of nature; and this is the only instance of such an occurrence known to me, with the exception of some wild plants of the Oxlip—a hybrid between P. veris and vulgaris—which were equal-styled. Herr Breitenbach's case is remarkable in another way; for equal-styled flowers were found in two instances on plants which bore both long-styled and short-styled flowers. In every other instance these two forms and the equal-styled variety have been produced by distinct plants.]

Primula vulgaris, BRIT. FL.

VAR. acaulis OF LINN. AND P. acaulis OF JACQ.


Mr. Scott states that this variety, which grew in the Botanic Garden in Edinburgh, was quite sterile when fertilised with pollen from the common primrose, as well as from a white variety of the same species, but that some of the plants, when artificially fertilised with their own pollen, yielded a moderate supply of seed. (5/10. 'Journal of the Proceedings of the Linnean Society' 8 1864 page 98.) He was so kind as to send me some of these self- fertilised seeds, from which I raised the plants immediately to be described. I may premise that the results of my experiments on the seedlings, made on a large scale, do not accord with those by Mr. Scott on the parent-plant.

First, in regard to the transmission of form and colour. The parent-plant was long-styled, and of a rich purple colour. From the self-fertilised seed 23 plants were raised; of these 18 were purple of different shades, with two of them a little streaked and freckled with yellow, thus showing a tendency to reversion; and 5 were yellow, but generally with a brighter orange centre than in the wild flower. All the plants were profuse flowerers. All were long-styled; but the pistil varied a good deal in length even on the same plant, being rather shorter, or considerably longer, than in the normal long-styled form; and the stigmas likewise varied in shape. It is, therefore, probable that an equal- styled variety of the primrose might be found on careful search; and I have received two accounts of plants apparently in this condition. The stamens always occupied their proper position low down in the corolla; and the pollen-grains were of the small size proper to the long-styled form, but were mingled with many minute and shrivelled grains. The yellow-flowered and the purple-flowered plants of this first generation were fertilised under a net with their own pollen, and the seed separately sown. From the former, 22 plants were raised, and all were yellow and long-styled. From the latter or the purple-flowered plants, 24 long-styled plants were raised, of which 17 were purple and 7 yellow. In this last case we have an instance of reversion in colour, without the possibility of any cross, to the grandparents or more distant progenitors of the plants in question. Altogether 23 plants in the first generation and 46 in the second generation were raised; and the whole of these 69 illegitimate plants were long-styled!

Eight purple-flowered and two yellow-flowered plants of the first illegitimate generation were fertilised in various ways with their own pollen and with that of the common primrose; and the seeds were separately counted, but as I could detect no difference in fertility between the purple and yellow varieties, the results are run together in Table 5.32.

TABLE 5.32. Primula vulgaris.

Column 1: Nature of plant experimented on, and kind of union. Column 2: Number of flowers fertilised. Column 3: Number of capsules produced. Column 4: Average Number of seeds per capsule. Column 5: Maximum Number of seeds in any one capsule. Column 6: Minimum Number of seeds in any one capsule.

Purple- and yellow-flowered illegitimate long-styled plants, ILLEGITIMATELY fertilised with pollen from the same plant : 72 : 11 : 11.5 : 26 : 5.

Purple- and yellow-flowered illegitimate long-styled plants, ILLEGITIMATELY fertilised with pollen from the common long-styled primrose : 72 : 39 : 31.4 : 62 : 3.

Or, if the ten poorest capsules, including less than 15 seeds, be rejected, we get: 72 : 29 : 40.6 : 62 : 18.

Purple- and yellow-flowered illegitimate long-styled plants, LEGITIMATELY fertilised with pollen from the common short-styled primrose : 26 : 18 : 36.4 : 60 : 9.

Or, if the two poorest capsules, including less than 15 seeds, be rejected, we get: 26 : 16 : 41.2 : 60 : 15.

The long-styled form of the common primrose ILLEGITIMATELY fertilised with pollen from the long-styled illegitimate purple- and yellow-flowered plants: 20 : 14 : 15.4 : 46 : 1.

Or, if the three poorest capsules be rejected, we get: 20 : 11 : 18.9 : 46 : 8.

The short-styled form of the common primrose LEGITIMATELY fertilised with pollen from the long-styled illegitimate purple- and yellow-flowered plants: 10 : 6 : 30.5 : 61 : 6.

If we compare the figures in this table with those given in the first chapter, showing the normal fertility of the common primrose, we shall see that the illegitimate purple- and yellow-flowered varieties are very sterile. For instance, 72 flowers were fertilised with their own pollen and produced only 11 good capsules; but by the standard they ought to have produced 48 capsules; and each of these ought to have contained on an average 52.2 seeds, instead of only 11.5 seeds. When these plants were illegitimately and legitimately fertilised with pollen from the common primrose, the average numbers were increased, but were far from attaining the normal standards. So it was when both forms of the common primrose were fertilised with pollen from these illegitimate plants; and this shows that their male as well as their female organs were in a deteriorated condition. The sterility of these plants was shown in another way, namely, by their not producing any capsules when the access of all insects (except such minute ones as Thrips) was prevented; for under these circumstances the common long-styled primrose produces a considerable number of capsules. There can, therefore, be no doubt that the fertility of these plants was greatly impaired. The loss is not correlated with the colour of the flower; and it was to ascertain this point that I made so many experiments. As the parent-plant growing in Edinburgh was found by Mr. Scott to be in a high degree sterile, it may have transmitted a similar tendency to its offspring, independently of their illegitimate birth. I am, however, inclined to attribute some weight to the illegitimacy of their descent, both from the analogy of other cases, and more especially from the fact that when the plants were LEGITIMATELY fertilised with pollen of the common primrose they yielded an average, as may be seen in the table, of only 5 more seeds than when ILLEGITIMATELY fertilised with the same pollen. Now we know that it is eminently characteristic of the illegitimate offspring of Primula Sinensis that they yield but few more seeds when legitimately fertilised than when fertilised with their own-form pollen.

Primula veris, Brit. Fl.

Var. officinalis of Linn., P. officinalis OF Jacq.

Seeds from the short-styled form of the cowslip fertilised with pollen from the same form germinate so badly that I raised from three successive sowings only fourteen plants, which consisted of nine short-styled and five long-styled plants. Hence the short-styled form of the cowslip, when self-fertilised, does not transmit the same form nearly so truly as does that of P. Sinensis. From the long-styled form, always fertilised with its own-form pollen, I raised in the first generation three long-styled plants,—from their seed 53 long-styled grandchildren,—from their seed 4 long-styled great-grandchildren,—from their seed 20 long-styled great-great-grandchildren,—and lastly, from their seed 8 long-styled and 2 short-styled great-great-great-grandchildren. In this last generation short-styled plants appeared for the first time in the course of the six generations,—the parent long-styled plant which was fertilised with pollen from another plant of the same form being counted as the first generation. Their appearance may be attributed to atavism. From two other long-styled plants, fertilised with their own-form pollen, 72 plants were raised, which consisted of 68 long-styled and 4 short-styled. So that altogether 162 plants were raised from illegitimately fertilised long-styled cowslips, and these consisted of 156 long-styled and 6 short-styled plants.

We will now turn to the fertility and powers of growth possessed by the illegitimate plants. From a short-styled plant, fertilised with its own-form pollen, one short-styled and two long-styled plants, and from a long-styled plant similarly fertilised three long-styled plants were at first raised. The fertility of these six illegitimate plants was carefully observed; but I must premise that I cannot give any satisfactory standard of comparison as far as the number of the seeds is concerned; for though I counted the seeds of many legitimate plants fertilised legitimately and illegitimately, the number varied so greatly during successive seasons that no one standard will serve well for illegitimate unions made during different seasons. Moreover the seeds in the same capsule frequently differ so much in size that it is scarcely possible to decide which ought to be counted as good seed. There remains as the best standard of comparison the proportional number of fertilised flowers which produce capsules containing any seed.

First, for the one illegitimate short-styled plant. In the course of three seasons 27 flowers were illegitimately fertilised with pollen from the same plant, and they yielded only a single capsule, which, however, contained a rather large number of seeds for a union of this nature, namely, 23. As a standard of comparison I may state that during the same three seasons 44 flowers borne by legitimate short-styled plants were self-fertilised, and yielded 26 capsules; so that the fact of the 27 flowers on the illegitimate plant having produced only one capsule proves how sterile it was. To show that the conditions of life were favourable, I will add that numerous plants of this and other species of Primula all produced an abundance of capsules whilst growing close by in the same soil with the present and following plants. The sterility of the above illegitimate short-styled plant depended on both the male and female organs being in a deteriorated condition. This was manifestly the case with the pollen; for many of the anthers were shrivelled or contabescent. Nevertheless some of the anthers contained pollen, with which I succeeded in fertilising some flowers on the illegitimate long-styled plants immediately to be described. Four flowers on this same short-styled plant were likewise LEGITIMATELY fertilised with pollen from one of the following long-styled plants; but only one capsule was produced, containing 26 seeds; and this is a very low number for a legitimate union.

With respect to the five illegitimate long-styled plants of the first generation, derived from the above self-fertilised short-styled and long-styled parents, their fertility was observed during the same three years. These five plants, when self-fertilised, differed considerably from one another in their degree of fertility, as was the case with the illegitimate long-styled plants of Lythrum salicaria; and their fertility varied much according to the season. I may premise, as a standard of comparison, that during the same years 56 flowers on legitimate long-styled plants of the same age and grown in the same soil, were fertilised with their own pollen, and yielded 27 capsules; that is, 48 per cent. On one of the five illegitimate long-styled plants 36 flowers were self- fertilised in the course of the three years, but they did not produce a single capsule. Many of the anthers on this plant were contabescent; but some seemed to contain sound pollen. Nor were the female organs quite impotent; for I obtained from a LEGITIMATE cross one capsule with good seed. On a second illegitimate long-styled plant 44 flowers were fertilised during the same years with their own pollen, but they produced only a single capsule. The third and fourth plants were in a very slight degree more productive. The fifth and last plant was decidedly more fertile; for 42 self-fertilised flowers yielded 11 capsules. Altogether, in the course of the three years, no less than 160 flowers on these five illegitimate long-styled plants were fertilised with their own pollen, but they yielded only 22 capsules. According to the standard above given, they ought to have yielded 80 capsules. These 22 capsules contained on an average 15.1 seeds. I believe, subject to the doubts before specified, that with legitimate plants the average number from a union of this nature would have been above 20 seeds. Twenty-four flowers on these same five illegitimate long-styled plants were legitimately fertilised with pollen from the above-described illegitimate short-styled plant, and produced only 9 capsules, which is an extremely small number for a legitimate union. These 9 capsules, however, contained an average of 38 apparently good seeds, which is as large a number as legitimate plants sometimes yield. But this high average was almost certainly false; and I mention the case for the sake of showing the difficulty of arriving at a fair result; for this average mainly depended on two capsules containing the extraordinary numbers of 75 and 56 seeds; these seeds, however, though I felt bound to count them, were so poor that, judging from trials made in other cases, I do not suppose that one would have germinated; and therefore they ought not to have been included. Lastly, 20 flowers were legitimately fertilised with pollen from a legitimate plant, and this increased their fertility; for they produced 10 capsules. Yet this is but a very small proportion for a legitimate union.

There can, therefore, be no doubt that these five long-styled plants and the one short-styled plant of the first illegitimate generation were extremely sterile. Their sterility was shown, as in the case of hybrids, in another way, namely, by their flowering profusely, and especially by the long endurance of the flowers. For instance, I fertilised many flowers on these plants, and fifteen days afterwards (namely on March 22nd) I fertilised numerous long-styled and short- styled flowers on common cowslips growing close by. These latter flowers, on April 8th, were withered, whilst most of the illegitimate flowers remained quite fresh for several days subsequently; so that some of these illegitimate plants, after being fertilised, remained in full bloom for above a month.

We will now turn to the fertility of the 53 illegitimate long-styled grandchildren, descended from the long-styled plant which was first fertilised with its own pollen. The pollen in two of these plants included a multitude of small and shrivelled grains. Nevertheless they were not very sterile; for 25 flowers, fertilised with their own pollen, produced 15 capsules, containing an average of 16.3 seeds. As already stated, the probable average with legitimate plants for a union of this nature is rather above 20 seeds. These plants were remarkably healthy and vigorous, as long as they were kept under highly favourable conditions in pots in the greenhouse; and such treatment greatly increases the fertility of the cowslip. When these same plants were planted during the next year (which, however, was an unfavourable one), out of doors in good soil, 20 self-fertilised flowers produced only 5 capsules, containing extremely few and wretched seeds.

Four long-styled great-grandchildren were raised from the self-fertilised grandchildren, and were kept under the same highly favourable conditions in the greenhouse; 10 of their flowers were fertilised with own-form pollen and yielded the large proportion of 6 capsules, containing on an average 18.7 seeds. From these seeds 20 long-styled great-great-grandchildren were raised, which were likewise kept in the greenhouse. Thirty of their flowers were fertilised with their own pollen and yielded 17 capsules, containing on an average no less than 32, mostly fine seeds. It appears, therefore, that the fertility of these plants of the fourth illegitimate generation, as long as they were kept under highly favourable conditions, had not decreased, but had rather increased. The result, however, was widely different when they were planted out of doors in good soil, where other cowslips grew vigorously and were completely fertile; for these illegitimate plants now became much dwarfed in stature and extremely sterile, notwithstanding that they were exposed to the visits of insects, and must have been legitimately fertilised by the surrounding legitimate plants. A whole row of these plants of the fourth illegitimate generation, thus freely exposed and legitimately fertilised, produced only 3 capsules, containing on an average only 17 seeds. During the ensuing winter almost all these plants died, and the few survivors were miserably unhealthy, whilst the surrounding legitimate plants were not in the least injured.

The seeds from the great-great-grandchildren were sown, and 8 long-styled and 2 short-styled plants of the fifth illegitimate generation raised. These whilst still in the greenhouse produced smaller leaves and shorter flower-stalks than some legitimate plants with which they grew in competition; but it should be observed that the latter were the product of a cross with a fresh stock,—a circumstance which by itself would have added much to their vigour. (5/11. For full details of this experiment, see my 'Effects of Cross and Self- fertilisation' 1876 page 220.) When these illegitimate plants were transferred to fairly good soil out of doors, they became during the two following years much more dwarfed in stature and produced very few flower-stems; and although they must have been legitimately fertilised by insects, they yielded capsules, compared with those produced by the surrounding legitimate plants, in the ratio only of 5 to 100! It is therefore certain that illegitimate fertilisation, continued during successive generations, affects the powers of growth and fertility of P. veris to an extraordinary degree; more especially when the plants are exposed to ordinary conditions of life, instead of being protected in a greenhouse.


Mr. Scott has described a plant of this kind growing in the Botanic Garden of Edinburgh. (5/12. 'Proceedings of the Linnean Society' volume 8 1864 page 105.) He states that it was highly self-fertile, although insects were excluded; and he explains this fact by showing, first, that the anthers and stigma are in close apposition, and that the stamens in length, position and size of their pollen-grains resemble those of the short-styled form, whilst the pistil resembles that of the long-styled form both in length and in the structure of the stigma. Hence the self-union of this variety is, in fact, a legitimate union, and consequently is highly fertile. Mr. Scott further states that this variety yielded very few seeds when fertilised by either the long- or short- styled common cowslip, and, again, that both forms of the latter, when fertilised by the equal-styled variety, likewise produced very few seeds. But his experiments with the cowslip were few, and my results do not confirm his in any uniform manner.

I raised twenty plants from self-fertilised seed sent me by Mr. Scott; and they all produced red flowers, varying slightly in tint. Of these, two were strictly long-styled both in structure and in function; for their reproductive powers were tested by crosses with both forms of the common cowslip. Six plants were equal-styled; but on the same plant the pistil varied a good deal in length during different seasons. This was likewise the case, according to Mr. Scott, with the parent-plant. Lastly, twelve plants were in appearance short-styled; but they varied much more in the length of their pistils than ordinary short- styled cowslips, and they differed widely from the latter in their powers of reproduction. Their pistils had become short-styled in structure, whilst remaining long-styled in function. Short-styled cowslips, when insects are excluded, are extremely barren: for instance, on one occasion six fine plants produced only about 50 seeds (that is, less than the product of two good capsules), and on another occasion not a single capsule. Now, when the above twelve apparently short-styled seedlings were similarly treated, nearly all produced a great abundance of capsules, containing numerous seeds, which germinated remarkably well. Moreover three of these plants, which during the first year were furnished with quite short pistils, on the following year produced pistils of extraordinary length. The greater number, therefore, of these short-styled plants could not be distinguished in function from the equal- styled variety. The anthers in the six equal-styled and in the apparently twelve short-styled plants were seated high up in the corolla, as in the true short- styled cowslip; and the pollen-grains resembled those of the same form in their large size, but were mingled with a few shrivelled grains. In function this pollen was identical with that of the short-styled cowslip; for ten long-styled flowers of the common cowslip, legitimately fertilised with pollen from a true equal-styled variety, produced six capsules, containing on an average 34.4 seeds; whilst seven capsules on a short-styled cowslip illegitimately fertilised with pollen from the equal-styled variety, yielded an average of only 14.5 seeds.

As the equal-styled plants differ from one another in their powers of reproduction, and as this is an important subject, I will give a few details with respect to five of them. First, an equal-styled plant, protected from insects (as was done in all the following cases, with one stated exception), spontaneously produced numerous capsules, five of which gave an average of 44.8 seeds, with a maximum in one capsule of 57. But six capsules, the product of fertilisation with pollen from a short-styled cowslip (and this is a legitimate union), gave an average of 28.5 seeds, with a maximum of 49; and this is a much lower average than might have been expected. Secondly, nine capsules from another equal-styled plant, which had not been protected from insects, but probably was self-fertilised, gave an average of 45.2 seeds, with a maximum of 58. Thirdly, another plant which had a very short pistil in 1865, produced spontaneously many capsules, six of which contained an average of 33.9 seeds, with a maximum of 38. In 1866 this same plant had a pistil of wonderful length; for it projected quite above the anthers, and the stigma resembled that of the long-styled form. In this condition it produced spontaneously a vast number of fine capsules, six of which contained almost exactly the same average number as before, namely 34.3, with a maximum of 38. Four flowers on this plant, legitimately fertilised with pollen from a short-styled cowslip, yielded capsules with an average of 30.2 seeds. Fourthly another short-styled plant spontaneously produced in 1865 an abundance of capsules, ten of which contained an average of 35.6 seeds, with a maximum of 54. In 1866 this same plant had become in all respects long-styled, and ten capsules gave almost exactly the same average as before, namely 35.1 seeds, with a maximum of 47. Eight flowers on this plant, legitimately fertilised with pollen from a short-styled cowslip, produced six capsules, with the high average of 53 seeds, and the high maximum of 67. Eight flowers were also fertilised with pollen from a long-styled cowslip (this being an illegitimate union), and produced seven capsules, containing an average of 24.4 seeds, with a maximum of 32. The fifth and last plant remained in the same condition during both years: it had a pistil rather longer than that of the true short-styled form, with the stigma smooth, as it ought to be in this form, but abnormal in shape, like a much-elongated inverted cone. It produced spontaneously many capsules, five of which, in 1865, gave an average of only 15.6 seeds; and in 1866 ten capsules still gave an average only a little higher, namely of 22.1, with a maximum of 30. Sixteen flowers were fertilised with pollen from a long-styled cowslip, and produced 12 capsules, with an average of 24.9 seeds, and a maximum of 42. Eight flowers were fertilised with pollen from a short-styled cowslip, but yielded only two capsules, containing 18 and 23 seeds. Hence this plant, in function and partially in structure, was in an almost exactly intermediate state between the long-styled and short-styled form, but inclining towards the short-styled; and this accounts for the low average of seeds which it produced when spontaneously self-fertilised.

The foregoing five plants thus differ much from one another in the nature of their fertility. In two individuals a great difference in the length of the pistil during two succeeding years made no difference in the number of seeds produced. As all five plants possessed the male organs of the short-styled form in a perfect state, and the female organs of the long-styled form in a more or less complete state, they spontaneously produced a surprising number of capsules, which generally contained a large average of remarkably fine seeds. With ordinary cowslips LEGITIMATELY FERTILISED, I once obtained from plants cultivated in the greenhouse the high average, from seven capsules, of 58.7 seeds, with a maximum in one capsule of 87 seeds; but from plants grown out of doors I never obtained a higher average than 41 seeds. Now two of the equal- styled plants, grown out of doors and spontaneously SELF-FERTILISED, gave averages of 44 and 45 seeds; but this high fertility may perhaps be in part attributed to the stigma receiving pollen from the surrounding anthers at exactly the right period. Two of these plants, fertilised with pollen from a short-styled cowslip (and this in fact is a legitimate union), gave a lower average than when self-fertilised. On the other hand, another plant, when similarly fertilised by a cowslip, yielded the unusually high average of 53 seeds, with a maximum of 67. Lastly, as we have just seen, one of these plants was in an almost exactly intermediate condition in its female organs between the long- and short-styled forms, and consequently, when self-fertilised, yielded a low average of seed. If we add together all the experiments which I made on the equal-styled plants, 41 spontaneously self-fertilised capsules (insects having been excluded) gave an average of 34 seeds, which is exactly the same number as the parent-plant yielded in Edinburgh. Thirty-four flowers, fertilised with pollen from the short-styled cowslip (and this is an analogous union), produced 17 capsules, containing an average of 33.8 seeds. It is a rather singular circumstance, for which I cannot account, that 20 flowers, artificially fertilised on one occasion with pollen from the same plants yielded only ten capsules, containing the low average of 26.7 seeds.

As bearing on inheritance, it may be added that 72 seedlings were raised from one of the red-flowered, strictly equal-styled, self-fertilised plants descended from the similarly characterised Edinburgh plant. These 72 plants were therefore grandchildren of the Edinburgh plant, and they all bore, as in the first generation, red flowers, with the exception of one plant, which reverted in colour to the common cowslip. In regard to structure, nine plants were truly long-styled and had their stamens seated low down in the corolla in the proper position; the remaining 63 plants were equal-styled, though the stigma in about a dozen of them stood a little below the anthers. We thus see that the anomalous combination in the same flower, of the male and female sexual organs which properly exist in the two distinct forms, was inherited with much force. Thirty- six seedlings were also raised from long and short-styled common cowslips, crossed with pollen from the equal-styled variety. Of these plants one alone was equal-styled, 20 were short-styled, but with the pistil in three of them rather too long, and the remaining 15 were long-styled. In this case we have an illustration of the difference between simple inheritance and prepotency of transmission; for the equal-styled variety, when self-fertilised, transmits its character, as we have just seen, with much force, but when crossed with the common cowslip cannot withstand the greater power of transmission of the latter.


I have little to say on this genus. I obtained seeds of P. officinalis from a garden where the long-styled form alone grew, and raised 11 seedlings, which were all long-styled. These plants were named for me by Dr. Hooker. They differed, as has been shown, from the plants belonging to this species which in Germany were experimented on by Hildebrand (5/13. 'Botanische Zeitung' 1865 page 13.); for he found that the long-styled form was absolutely sterile with its own pollen, whilst my long-styled seedlings and the parent-plants yielded a fair supply of seed when self-fertilised. Plants of the long-styled form of Pulmonaria angustifolia were, like Hildebrand's plants, absolutely sterile with their own pollen, so that I could never procure a single seed. On the other hand, the short-styled plants of this species, differently from those of P. officinalis, were fertile with their own pollen in a quite remarkable degree for a heterostyled plant. From seeds carefully self-fertilised I raised 18 plants, of which 13 proved short-styled and 5 long-styled.

Polygonum fagopyrum.

From flowers on long-styled plants fertilised illegitimately with pollen from the same plant, 49 seedlings were raised, and these consisted of 45 long-styled and 4 short-styled. From flowers on short-styled plants illegitimately fertilised with pollen from the same plant 33 seedlings were raised, and these consisted of 20 short-styled and 13 long-styled. So that the usual rule of illegitimately fertilised long-styled plants tending much more strongly than short-styled plants to reproduce their own form here holds good. The illegitimate plants derived from both forms flowered later than the legitimate, and were to the latter in height as 69 to 100. But as these illegitimate plants were descended from parents fertilised with their own pollen, whilst the legitimate plants were descended from parents crossed with pollen from a distinct individual, it is impossible to know how much of their difference in height and period of flowering, is due to the illegitimate birth of the one set, and how much to the other set being the product of a cross between distinct plants.]


It is remarkable how closely and in how many points illegitimate unions between the two or three forms of the same heterostyled species, together with their illegitimate offspring, resemble hybrid unions between distinct species together with their hybrid offspring. In both cases we meet with every degree of sterility, from very slightly lessened fertility to absolute barrenness, when not even a single seed-capsule is produced. In both cases the facility of effecting the first union is much influenced by the conditions to which the plants are exposed. (5/14. This has been remarked by many experimentalists in effecting crosses between distinct species; and in regard to illegitimate unions I have given in the first chapter a striking illustration in the case of Primula veris.) Both with hybrids and illegitimate plants the innate degree of sterility is highly variable in plants raised from the same mother-plant. In both cases the male organs are more plainly affected than the female; and we often find contabescent anthers enclosing shrivelled and utterly powerless pollen-grains. The more sterile hybrids, as Max Wichura has well shown, are sometimes much dwarfed in stature, and have so weak a constitution that they are liable to premature death (5/15. 'Die Bastardbefruchtung im Pflanzenreich' 1865.); and we have seen exactly parallel cases with the illegitimate seedlings of Lythrum and Primula. Many hybrids are the most persistent and profuse flowerers, as are some illegitimate plants. When a hybrid is crossed by either pure parent-form, it is notoriously much more fertile than when crossed inter se or by another hybrid; so when an illegitimate plant is fertilised by a legitimate plant, it is more fertile than when fertilised inter se or by another illegitimate plant. When two species are crossed and they produce numerous seeds, we expect as a general rule that their hybrid offspring will be moderately fertile; but if the parent species produce extremely few seeds, we expect that the hybrids will be very sterile. But there are marked exceptions, as shown by Gartner, to these rules. So it is with illegitimate unions and illegitimate offspring. Thus the mid- styled form of Lythrum salicaria, when illegitimately fertilised with pollen from the longest stamens of the short-styled form, produced an unusual number of seeds; and their illegitimate offspring were not at all, or hardly at all, sterile. On the other hand, the illegitimate offspring from the long-styled form, fertilised with pollen from the shortest stamens of the same form, yielded few seeds, and the illegitimate offspring thus produced were very sterile; but they were more sterile than might have been expected relatively to the difficulty of effecting the union of the parent sexual elements. No point is more remarkable in regard to the crossing of species than their unequal reciprocity. Thus species A will fertilise B with the greatest ease; but B will not fertilise A after hundreds of trials. We have exactly the same case with illegitimate unions; for the mid-styled Lythrum salicaria was easily fertilised by pollen from the longest stamens of the short-styled form, and yielded many seeds; but the latter form did not yield a single seed when fertilised by the longest stamens of the mid-styled form.

Another important point is prepotency. Gartner has shown that when a species is fertilised with pollen from another species, if it be afterwards fertilised with its own pollen, or with that of the same species, this is so prepotent over the foreign pollen that the effect of the latter, though placed on the stigma some time previously, is entirely destroyed. Exactly the same thing occurs with the two forms of a heterostyled species. Thus several long-styled flowers of Primula veris were fertilised illegitimately with pollen from another plant of the same form, and twenty-four hours afterwards legitimately with pollen from a short- styled dark-red polyanthus which is a variety of P. veris; and the result was that every one of the thirty seedlings thus raised bore flowers more or less red, showing plainly how prepotent the legitimate pollen from a short-styled plant was over the illegitimate pollen from a long-styled plant.

In all the several foregoing points the parallelism is wonderfully close between the effects of illegitimate and hybrid fertilisation. It is hardly an exaggeration to assert that seedlings from an illegitimately fertilised heterostyled plant are hybrids formed within the limits of one and the same species. This conclusion is important, for we thus learn that the difficulty in sexually uniting two organic forms and the sterility of their offspring, afford no sure criterion of so-called specific distinctness. If any one were to cross two varieties of the same form of Lythrum or Primula for the sake of ascertaining whether they were specifically distinct, and he found that they could be united only with some difficulty, that their offspring were extremely sterile, and that the parents and their offspring resembled in a whole series of relations crossed species and their hybrid offspring, he might maintain that his varieties had been proved to be good and true species; but he would be completely deceived. In the second place, as the forms of the same trimorphic or dimorphic heterostyled species are obviously identical in general structure, with the exception of the reproductive organs, and as they are identical in general constitution (for they live under precisely the same conditions), the sterility of their illegitimate unions and that of their illegitimate offspring, must depend exclusively on the nature of the sexual elements and on their incompatibility for uniting in a particular manner. And as we have just seen that distinct species when crossed resemble in a whole series of relations the forms of the same species when illegitimately united, we are led to conclude that the sterility of the former must likewise depend exclusively on the incompatible nature of their sexual elements, and not on any general difference in constitution or structure. We are, indeed, led to this same conclusion by the impossibility of detecting any differences sufficient to account for certain species crossing with the greatest ease, whilst other closely allied species cannot be crossed, or can be crossed only with extreme difficulty. We are led to this conclusion still more forcibly by considering the great difference which often exists in the facility of crossing reciprocally the same two species; for it is manifest in this case that the result must depend on the nature of the sexual elements, the male element of the one species acting freely on the female element of the other, but not so in a reversed direction. And now we see that this same conclusion is independently and strongly fortified by the consideration of the illegitimate unions of trimorphic and dimorphic heterostyled plants. In so complex and obscure a subject as hybridism it is no slight gain to arrive at a definite conclusion, namely, that we must look exclusively to functional differences in the sexual elements, as the cause of the sterility of species when first crossed and of their hybrid offspring. It was this consideration which led me to make the many observations recorded in this chapter, and which in my opinion make them worthy of publication.


The essential character of heterostyled plants. Summary of the differences in fertility between legitimately and illegitimately fertilised plants. Diameter of the pollen-grains, size of anthers and structure of stigma in the different forms. Affinities of the genera which include heterostyled species. Nature of the advantages derived from heterostylism. The means by which plants became heterostyled. Transmission of form. Equal-styled varieties of heterostyled plants. Final remarks.

In the foregoing chapters all the heterostyled plants known to me have been more or less fully described. Several other cases have been indicated, especially by Professor Asa Gray and Kuhn, in which the individuals of the same species differ in the length of their stamens and pistils (6/1. Asa Gray 'American Journal of Science' 1865 page 101 and elsewhere as already referred to. Kuhn 'Botanische Zeitung' 1867 page 67.); but as I have been often deceived by this character taken alone, it seems to me the more prudent course not to rank any species as heterostyled, unless we have evidence of more important differences between the forms, as in the diameter of the pollen-grains, or in the structure of the stigma. The individuals of many ordinary hermaphrodite plants habitually fertilise one another, owing to their male and female organs being mature at different periods, or to the structure of the parts, or to self-sterility, etc.; and so it is with many hermaphrodite animals, for instance, land-snails or earth-worms; but in all these cases any one individual can fully fertilise or be fertilised by any other individual of the same species. This is not so with heterostyled plants: a long-styled, mid-styled or short-styled plant cannot fully fertilise or be fertilised by any other individual, but only by one belonging to another form. Thus the essential character of plants belonging to the heterostyled class is that the individuals are divided into two or three bodies, like the males and females of dioecious plants or of the higher animals, which exist in approximately equal numbers and are adapted for reciprocal fertilisation. The existence, therefore, of two or three bodies of individuals, differing from one another in the above more important characteristics, offers by itself good evidence that the species is heterostyled. But absolutely conclusive evidence can be derived only from experiments, and by finding that pollen must be applied from the one form to the other in order to ensure complete fertility.

In order to show how much more fertile each form is when legitimately fertilised with pollen from the other form (or in the case of trimorphic species, with the proper pollen from one of the two other forms) than when illegitimately fertilised with its own-form pollen, I will append Table 6.33 giving a summary of the results in all the cases hitherto ascertained. The fertility of the unions may be judged by two standards, namely, by the proportion of flowers which, when fertilised in the two methods, yield capsules, and by the average number of seeds per capsule. When there is a dash in the left hand column opposite to the name of the species, the proportion of the flowers which yielded capsules was not recorded.

TABLE 6.33. Fertility of the legitimate unions taken together, compared with that of the illegitimate unions together. The fertility of the legitimate unions, as judged by both standards, is taken as 100.

Column 1: Name of species. Column 2: Illegitimate unions : proportional number of flowers which produced capsules. Column 3: Illegitimate unions : average number of seeds per capsule.

Primula veris : 69 : 65.

Primula elatior : 27 : 75.

Primula vulgaris : 60 : 54.

Primula Sinensis : 84 : 63.

Primula Sinensis (second trial) : 0 : 53.

Primula Sinensis (Hildebrand) : 100 : 42.

Primula auricula (Scott) : 80 : 15.

Primula Sikkimensis (Scott) : 95 : 31.

Primula cortusoides (Scott) : 74 : 66.

Primula involucrata (Scott) : 72 : 48.

Primula farinosa (Scott) : 71 : 44.

Average of the nine species of Primula : 88.4 : 69.

Hottonia palustris (H. Muller) : - : 61.

Linum grandiflorum (the difference probably is much greater) : - : 69.

Linum perenne : - : 20.

Linum perenne (Hildebrand) : 0 : 0.

Pulmonaria officinalis (German stock, Hildebrand) : 0 : 0.

Pulmonaria angustifolia : 35 : 32.

Mitchella repens : 20 : 47.

Borreria, Brazilian sp. : - : 0.

Polygonum fagopyrum : - : 46.

Lythrum salicaria : 33 : 46.

Oxalis Valdiviana (Hildebrand) : 2 : 34.

Oxalis Regnelli : 0 : 0.

Oxalis speciosa : 15 : 49.

The two or three forms of the same heterostyled species do not differ from one another in general habit or foliage, as sometimes, though rarely, happens with the two sexes of dioecious plants. Nor does the calyx differ, but the corolla sometimes differs slightly in shape, owing to the different position of the anthers. In Borreria the hairs within the tube of the corolla are differently situated in the long-styled and short-styled forms. In Pulmonaria there is a slight difference in the size of the corolla, and in Pontederia in its colour. In the reproductive organs the differences are much greater and more important. In the one form the stamens may be all of the same length, and in the other graduated in length, or alternately longer and shorter. The filaments may differ in colour and thickness, and are sometimes nearly thrice as long in the one form as in the other. They adhere also for very different proportional lengths to the corolla. The anthers sometimes differ much in size in the two forms. Owing to the rotation of the filaments, the anthers, when mature, dehisce towards the circumference of the flower in one form of Faramea, and towards the centre in the other form. The pollen-grains sometimes differ conspicuously in colour, and often to an extraordinary degree in diameter. They differ also somewhat in shape, and apparently in their contents, as they are unequally opaque. In the short-styled form of Faramea the pollen-grains are covered with sharp points, so as to cohere readily together or to an insect; whilst the smaller grains of the long-styled form are quite smooth.

With respect to the pistil, the style may be almost thrice as long in the one form as in the other. In Oxalis it sometimes differs in hairiness in the three forms. In Linum the pistils either diverge and pass out between the filaments, or stand nearly upright and parallel to them. The stigmas in the two forms often differ much in size and shape, and more especially in the length and thickness of their papillae; so that the surface may be rough or quite smooth. Owing to the rotation of the styles, the papillose surface of the stigma is turned outwards in one form of Linum perenne, and inwards in the other form. In flowers of the same age of Primula veris the ovules are larger in the long-styled than in the short-styled form. The seeds produced by the two or three forms often differ in number, and sometimes in size and weight; thus, five seeds from the long-styled form of Lythrum salicaria equal in weight six from the mid-styled and seven from the short-styled form. Lastly, short-styled plants of Pulmonaria officinalis bear a larger number of flowers, and these set a larger proportional number of fruit, which however yield a lower average number of seed, than the long-styled plants. With heterostyled plants we thus see in how many and in what important characters the forms of the same undoubted species often differ from one another—characters which with ordinary plants would be amply sufficient to distinguish species of the same genus.

As the pollen-grains of ordinary species belonging to the same genus generally resemble one another closely in all respects, it is worth while to show, in Table 6.34, the difference in diameter between the grains from the two or three forms of the same heterostyled species in the forty-three cases in which this was ascertained. But it should be observed that some of the following measurements are only approximately accurate, as only a few grains were measured. In several cases, also, the grains had been dried and were then soaked in water. Whenever they were of an elongated shape their longer diameters were measured. The grains from the short-styled plants are invariably larger than those from the long-styled, whenever there is any difference between them. The diameter of the former is represented in the table by the number 100.

TABLE 6.34. Relative diameter of the pollen-grains from the forms of the same heterostyled species; those from the short-styled form being represented by 100.


Column 1: Name of species. Column 2: From the long-styled form : relative diameter.

Primula veris : 67.

Primula vulgaris : 71.

Primula Sinensis (Hildebrand) : 57.

Primula auricula : 71.

Hottonia palustris (H. Muller) : 61.

Hottonia palustris (self) : 64.

Linum grandiflorum : 100.

Linum perenne (diameter variable) : 100 (?).

Linum flavum : 100.

Pulmonaria officinalis : 78.

Pulmonaria angustifolia : 91.

Polygonum fagopyrum : 82.

Leucosmia Burnettiana : 99.

Aegiphila elata : 62.

Menyanthes trifoliata : 84.

Limnanthemum Indicum : 100.

Villarsia (sp.?) : 75.

Forsythia suspensa : 94.

Cordia (sp.?) : 100.

Gilia pulchella : 100.

Gilia micrantha : 81.

Sethia acuminata : 83.

Erythroxylum (sp.?) : 93.

Cratoxylon formosum : 86.

Mitchella repens, pollen-grains of the long-styled a little smaller.

Borreria (sp.?) : 92.

Faramea (sp.?) : 67.

Suteria (sp.?) (Fritz Muller) : 75.

Houstonia coerulea : 72.

Oldenlandia (sp.?) : 78.

Hedyotis (sp.?) : 88.

Coccocypselum (sp.?) (Fritz Muller) : 100.

Lipostoma (sp.?) : 80.

Cinchona micrantha : 91.


Column 1: Name of species. Column 2: Ratio expressing the extreme differences in diameter of the pollen- grains from the two sets of anthers in the three forms.

Lythrum salicaria : 60.

Nesaea verticillata : 65.

Oxalis Valdiviana (Hildebrand) : 71.

Oxalis Regnelli : 78.

Oxalis speciosa : 69.

Oxalis sensitiva : 84.

Pontederia (sp.?) : 55.

Column 1: Name of species. Column 2: Ratio between the diameters of the pollen-grains of the two sets of anthers in the same form.

Oxalis rosea, long-styled form (Hildebrand) : 83.

Oxalis compressa, short-styled form : 83.

Pontederia (sp.?) short-styled form : 87.

Pontederia other sp. mid-styled form : 86.

We here see that, with seven or eight exceptions out of the forty-three cases, the pollen-grains from one form are larger than those from the other form of the same species. The extreme difference is as 100 to 55; and we should bear in mind that in the case of spheres differing to this degree in diameter, their contents differ in the ratio of six to one. With all the species in which the grains differ in diameter, there is no exception to the rule that those from the anthers of the short-styled form, the tubes of which have to penetrate the longer pistil of the long-styled form, are larger than the grains from the other form. This curious relation led Delpino (as it formerly did me) to believe that the larger size of the grains in the short-styled flowers is connected with the greater supply of matter needed for the development of their longer tubes. (6/2. 'Sull' Opera, la Distribuzione dei Sessi nelle Piante' etc 1867 page 17.) But the case of Linum, in which the grains of the two forms are of equal size, whilst the pistil of the one is about twice as long as that of the other, made me from the first feel very doubtful with respect to this view. My doubts have since been strengthened by the cases of Limnanthemum and Coccocypselum, in which the grains are of equal size in the two forms; whilst in the former genus the pistil is nearly thrice and in the latter twice as long as in the other form. In those species in which the grains are of unequal size in the two forms, there is no close relationship between the degree of their inequality and that of their pistils. Thus in Pulmonaria officinalis and in Erythroxylum the pistil in the long-styled form is about twice the length of that in the other form, whilst in the former species the pollen-grains are as 100 to 78, and in the latter as 100 to 93 in diameter. In the two forms of Suteria the pistil differs but little in length, whilst the pollen-grains are as 100 to 75 in diameter. These cases seem to prove that the difference in size between the grains in the two forms is not determined by the length of the pistil, down which the tubes have to grow. That with plants in general there is no close relationship between the size of the pollen-grains and the length of the pistil is manifest: for instance, I found that the distended grains of Datura arborea were .00243 of an inch in diameter, and the pistil no less than 9.25 inches in length; now the pistil in the small flowers of Polygonum fagopyrum is very short, yet the larger pollen-grains from the short-styled plants had exactly the same diameter as those from the Datura, with its enormously elongated pistil.

Notwithstanding these several considerations, it is difficult quite to give up the belief that the pollen-grains from the longer stamens of heterostyled plants have become larger in order to allow of the development of longer tubes; and the foregoing opposing facts may possibly be reconciled in the following manner. The tubes are at first developed from matter contained within the grains, for they are sometimes exserted to a considerable length, before the grains have touched the stigma; but botanists believe that they afterwards draw nourishment from the conducting tissue of the pistil. It is hardly possible to doubt that this must occur in such cases as that of the Datura, in which the tubes have to grow down the whole length of the pistil, and therefore to a length equalling 3,806 times the diameter of the grains (namely, .00243 of an inch) from which they are protruded. I may here remark that I have seen the pollen-grains of a willow, immersed in a very weak solution of honey, protrude their tubes, in the course of twelve hours, to a length thirteen times as great as the diameter of the grains. Now if we suppose that the tubes in some heterostyled species are developed wholly or almost wholly from matter contained within the grains, while in other species from matter yielded by the pistil, we can see that in the former case it would be necessary that the grains of the two forms should differ in size relatively to the length of the pistil which the tubes have to penetrate, but that in the latter case it would not be necessary that the grains should thus differ. Whether this explanation can be considered satisfactory must remain at present doubtful.

There is another remarkable difference between the forms of several heterostyled species, namely in the anthers of the short-styled flowers, which contain the larger pollen-grains, being longer than those of the long-styled flowers. This is the case with Hottonia palustris in the ratio of 100 to 83. With Limnanthemum Indicum the ratio is as 100 to 70. With the allied Menyanthes the anthers of the short-styled form are a little and with Villarsia conspicuously larger than those of the long-styled. With Pulmonaria angustifolia they vary much in size, but from an average of seven measurements of each kind the ratio is as 100 to 91. In six genera of the Rubiaceae there is a similar difference, either slightly or well marked. Lastly, in the trimorphic Pontederia the ratio is 100 to 88; the anthers from the longest stamens in the short-styled form being compared with those from the shortest stamens in the long-styled form. On the other hand, there is a similar and well-marked difference in the length of the stamens in the two forms of Forsythia suspensa and of Linum flavum; but in these two cases the anthers of the short-styled flowers are shorter than those of the long-styled. The relative size of the anthers was not particularly attended to in the two forms of the other heterostyled plants, but I believe that they are generally equal, as is certainly the case with those of the common primrose and cowslip.

The pistil differs in length in the two forms of every heterostyled plant, and although a similar difference is very general with the stamens, yet in the two forms of Linum grandiflorum and of Cordia they are equal. There can hardly be a doubt that the relative length of these organs is an adaptation for the safe transportal by insects of the pollen from the one form to the other. The exceptional cases in which these organs do not stand exactly on a level in the two forms may probably be explained by the manner in which the flowers are visited. With most of the species, if there is any difference in the size of the stigma of the two forms, that of the long-styled, whatever its shape may be, is larger than that of the short-styled. But here again there are some exceptions to the rule, for in the short-styled form of Leucosmia Burnettiana the stigmas are longer and much narrower than those of the long-styled; the ratio between the lengths of the stigmas in the two forms being 100 to 60. In the three Rubiaceous genera, Faramea, Houstonia and Oldenlandia, the stigmas of the short- styled form are likewise somewhat longer and narrower; and in the three forms of Oxalis sensitiva the difference is strongly marked, for if the length of the two stigmas of the long-styled pistil be taken as 100, it will be represented in the mid- and short-styled forms by the numbers 141 and 164. As in all these cases the stigmas of the short-styled pistil are seated low down within a more or less tubular corolla, it is probable that they are better fitted by being long and narrow for brushing the pollen off the inserted proboscis of an insect.

With many heterostyled plants the stigma differs in roughness in the two forms, and when this is the case there is no known exception to the rule that the papillae on the stigma of the long-styled form are longer and often thicker than those on that of the short-styled. For instance, the papillae on the long-styled stigma of Hottonia palustris are more than twice the length of those in the other form. This holds good even in the case of Houstonia coerulea, in which the stigmas are much shorter and stouter in the long-styled than in the short-styled form, for the papillae on the former compared with those on the latter are as 100 to 58 in length. The length of the pistil in the long-styled form of Linum grandiflorum varies much, and the stigmatic papillae vary in a corresponding manner. From this fact I inferred at first that in all cases the difference in length between the stigmatic papillae in the two forms was one merely of correlated growth; but this can hardly be the true or general explanation, as the shorter stigmas of the long-styled form of Houstonia have the longer papillae. It is a more probable view that the papillae, which render the stigma of the long-styled form of various species rough, serve to entangle effectually the large-sized pollen-grains brought by insects from the short-styled form, thus ensuring its legitimate fertilisation. This view is supported by the fact that the pollen-grains from the two forms of eight species in Table 6.34 hardly differ in diameter, and the papillae on their stigmas do not differ in length.

The species which are at present positively or almost positively known to be heterostyled belong, as shown in Table 6.35, to 38 genera, widely distributed throughout the world. These genera are included in fourteen Families, most of which are very distinct from one another, for they belong to nine of the several great Series, into which phanerogamic plants have been divided by Bentham and Hooker.

TABLE 6.35. List of genera including heterostyled species.



ERYTHROXYLEAE: Erythroxylum. Sethia.

GERANIACEAE: Linum. Oxalis.

LYTHRACEAE: Lythrum. Neseae.

RUBIACEAE: Cinchona. Bouvardia. Manettia. Hedyotis. Oldenlandia. Houstonia. Coccocypselum. Lipostoma. Knoxia. Faramea. Psychotria. Rudgea. Suteria. Mitchella. Diodia. Borreria. Spermacoce.

PRIMULACEAE: Primula. Hottonia. Androsace.

OLEACEAE: Forsythia.

GENTIANACEAE: Menyanthes. Limnanthemum. Villarsia.



BORAGINEAE: Pulmonaria.


POLYGONEAE: Polygonum.

THYMELEAE: Thymelea.



In some of these families the heterostyled condition must have been acquired at a very remote period. Thus the three closely allied genera, Menyanthes, Limnanthemum, and Villarsia, inhabit respectively Europe, India, and South America. Heterostyled species of Hedyotis are found in the temperate regions of North and the tropical regions of South America. Trimorphic species of Oxalis live on both sides of the Cordillera in South America and at the Cape of Good Hope. In these and some other cases it is not probable that each species acquired its heterostyled structure independently of its close allies. If they did not do so, the three closely connected genera of the Menyantheae and the several trimorphic species of Oxalis must have inherited their structure from a common progenitor. But an immense lapse of time will have been necessary in all such cases for the modified descendants of a common progenitor to have spread from a single centre to such widely remote and separated areas. The family of the Rubiaceae contains not far short of as many heterostyled genera as all the other thirteen families together; and hereafter no doubt other Rubiaceous genera will be found to be heterostyled, although a large majority are homostyled. Several closely allied genera in this family probably owe their heterostyled structure to descent in common; but as the genera thus characterised are distributed in no less than eight of the tribes into which this family has been divided by Bentham and Hooker, it is almost certain that several of them must have become heterostyled independently of one another. What there is in the constitution or structure of the members of this family which favours their becoming heterostyled, I cannot conjecture. Some families of considerable size, such as the Boragineae and Verbenaceae, include, as far as is at present known, only a single heterostyled genus. Polygonum also is the sole heterostyled genus in its family; and though it is a very large genus, no other species except P. fagopyrum is thus characterised. We may suspect that it has become heterostyled within a comparatively recent period, as it seems to be less strongly so in function than the species in any other genus, for both forms are capable of yielding a considerable number of spontaneously self-fertilised seeds. Polygonum in possessing only a single heterostyled species is an extreme case; but every other genus of considerable size which includes some such species likewise contains homostyled species. Lythrum includes trimorphic, dimorphic, and homostyled species.

Trees, bushes, and herbaceous plants, both large and small, bearing single flowers or flowers in dense spikes or heads, have been rendered heterostyled. So have plants which inhabit alpine and lowland sites, dry land, marshes and water. (6/3. Out of the 38 genera known to include heterostyled species, about eight, or 21 per cent, are more or less aquatic in their habits. I was at first struck with this fact, for I was not then aware how large a proportion of ordinary plants inhabit such stations. Heterostyled plants may be said in one sense to have their sexes separated, as the forms must mutually fertilise one another. Therefore it seemed worth while to ascertain what proportion of the genera in the Linnean classes, Monoecia, Dioecia and Polygamia, contained species which live "in water, marshes, bogs or watery places." In Sir W.J. Hooker's 'British Flora' 4th edition 1838, these three Linnean classes include 40 genera, 17 of which (i.e. 43 per cent) contain species inhabiting the just-specified stations. So that 43 per cent of those British plants which have their sexes separated are more or less aquatic in their habits, whereas only 21 per cent of heterostyled plants have such habits. I may add that the hermaphrodite classes, from Monandria to Gynandria inclusive, contain 447 genera, of which 113 are aquatic in the above sense, or only 25 per cent. It thus appears, as far as can be judged from such imperfect data, that there is some connection between the separation of the sexes in plants and the watery nature of the sites which they inhabit; but that this does not hold good with heterostyled species.)

When I first began to experimentise on heterostyled plants it was under the impression that they were tending to become dioecious; but I was soon forced to relinquish this notion, as the long-styled plants of Primula which, from possessing a longer pistil, larger stigma, shorter stamens with smaller pollen- grains, seemed to be the more feminine of the two forms, yielded fewer seeds than the short-styled plants which appeared to be in the above respects the more masculine of the two. Moreover, trimorphic plants evidently come under the same category with dimorphic, and the former cannot be looked at as tending to become dioecious. With Lythrum salicaria, however, we have the curious and unique case of the mid-styled form being more feminine or less masculine in nature than the other two forms. This is shown by the large number of seeds which it yields in whatever manner it may be fertilised, and by its pollen (the grains of which are of smaller size than those from the corresponding stamens in the other two forms) when applied to the stigma of any form producing fewer seeds than the normal number. If we suppose the process of deterioration of the male organs in the mid-styled form to continue, the final result would be the production of a female plant; and Lythrum salicaria would then consist of two heterostyled hermaphrodites and a female. No such case is known to exist, but it is a possible one, as hermaphrodite and female forms of the same species are by no means rare. Although there is no reason to believe that heterostyled plants are regularly becoming dioecious, yet they offer singular facilities, as will hereafter be shown, for such conversion; and this appears occasionally to have been effected.

We may feel sure that plants have been rendered heterostyled to ensure cross- fertilisation, for we now know that a cross between the distinct individuals of the same species is highly important for the vigour and fertility of the offspring. The same end is gained by dichogamy or the maturation of the reproductive elements of the same flower at different periods,—by dioeciousness—self-sterility—the prepotency of pollen from another individual over a plant's own pollen,—and lastly, by the structure of the flower in relation to the visits of insects. The wonderful diversity of the means for gaining the same end in this case, and in many others, depends on the nature of all the previous changes through which the species has passed, and on the more or less complete inheritance of the successive adaptations of each part to the surrounding conditions. Plants which are already well adapted by the structure of their flowers for cross-fertilisation by the aid of insects often possess an irregular corolla, which has been modelled in relation to their visits; and it would have been of little or no use to such plants to have become heterostyled. We can thus understand why it is that not a single species is heterostyled in such great families as the Leguminosae, Labiatae, Scrophulariaceae, Orchideae, etc., all of which have irregular flowers. Every known heterostyled plant, however, depends on insects for its fertilisation, and not on the wind; so that it is a rather surprising fact that only one genus, Pontederia, has a plainly irregular corolla.

Why some species are adapted for cross-fertilisation, whilst others within the same genus are not so, or if they once were, have since lost such adaptation and in consequence are now usually self-fertilised, I have endeavoured elsewhere to explain to a certain limited extent. (6/4. 'The Effects of Cross and Self- fertilisation' 1876 page 441.) If it be further asked why some species have been adapted for this end by being made heterostyled, rather than by any of the above specified means, the answer probably lies in the manner in which heterostylism originated,—a subject immediately to be discussed. Heterostyled species, however, have an advantage over dichogamous species, as all the flowers on the same heterostyled plant belong to the same form, so that when fertilised legitimately by insects two distinct individuals are sure to intercross. On the other hand, with dichogamous plants, early or late flowers on the same individual may intercross; and a cross of this kind does hardly any or no good. Whenever it is profitable to a species to produce a large number of seeds and this obviously is a very common case, heterostyled will have an advantage over dioecious plants, as all the individuals of the former, whilst only half of the latter, that is the females, yield seeds. On the other hand, heterostyled plants seem to have no advantage, as far as cross-fertilisation is concerned, over those which are sterile with their own pollen. They lie indeed under a slight disadvantage, for if two self-sterile plants grow near together and far removed from all other plants of the same species, they will mutually and perfectly fertilise one another, whilst this will not be the case with heterostyled dimorphic plants, unless they chance to belong to opposite forms.

It may be added that species which are trimorphic have one slight advantage over the dimorphic; for if only two individuals of a dimorphic species happen to grow near together in an isolated spot, the chances are even that both will belong to the same form, and in this case they will not produce the full number of vigorous and fertile seedlings; all these, moreover, will tend strongly to belong to the same form as their parents. On the other hand, if two plants of the same trimorphic species happen to grow in an isolated spot, the chances are two to one in favour of their not belonging to the same form; and in this case they will legitimately fertilise one another, and yield the full complement of vigorous offspring.


This is a very obscure subject, on which I can throw little light, but which is worthy of discussion. It has been shown that heterostyled plants occur in fourteen natural families, dispersed throughout the whole vegetable kingdom, and that even within the family of the Rubiaceae they are dispersed in eight of the tribes. We may therefore conclude that this structure has been acquired by various plants independently of inheritance from a common progenitor, and that it can be acquired without any great difficulty—that is, without any very unusual combination of circumstances.

It is probable that the first step towards a species becoming heterostyled is great variability in the length of the pistil and stamens, or of the pistil alone. Such variations are not very rare: with Amsinckia spectabilis and Nolana prostrata these organs differ so much in length in different individuals that, until experimenting on them, I thought both species heterostyled. The stigma of Gesneria pendulina sometimes protrudes far beyond, and is sometimes seated beneath the anthers; so it is with Oxalis acetosella and various other plants. I have also noticed an extraordinary amount of difference in the length of the pistil in cultivated varieties of Primula veris and vulgaris.

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