The Different Forms of Flowers on Plants of the Same Species Part 5

The two illegitimate unions together: 37 : 48.1.

The most remarkable point in this table is the small average number of seeds from the short-styled flowers when illegitimately fertilised, and the unusually large average number of seeds yielded by the illegitimately fertilised long- styled flowers, relatively in both cases to the product of the legitimately fertilised flowers. (1/18. H. Muller says 'Die Befruchtung' etc. page 352, that the long-styled flowers, when illegitimately fertilised, yield as many seeds as when legitimately fertilised; but by adding up the number of seeds from all the capsules produced by the two methods of fertilisation, as given by him, I arrive at the results shown in Table 1.13. The average number in the long-styled capsules, when legitimately fertilised, is 91.4, and when illegitimately fertilised, 77.5; or as 100 to 85. H. Muller agrees with me that this is the proper manner of viewing the case.) The two legitimate unions compared with the two illegitimate together yield seeds in the ratio of 100 to 61.

H. Muller also tried the effects of illegitimately fertilising the long-styled and short-styled flowers with their own pollen, instead of with that from another plant of the same form; and the results are very striking. For the capsules from the long-styled flowers thus treated contained, on an average, only 15.7 seeds instead of 77.5; and those from the short-styled 6.5, instead of 18.7 seeds per capsule. The number 6.5 agrees closely with Mr. Scott's result from the same form similarly fertilised.

From some observations by Dr. Torrey, Hottonia inflata, an inhabitant of the United States, does not appear to be heterostyled, but is remarkable from producing cleistogamic flowers, as will be seen in the last chapter of this volume.

Besides the genera Primula and Hottonia, Androsace (vel Gregoria, vel Aretia) vitalliana is heterostyled. Mr. Scott fertilised with their own pollen 21 flowers on three short-styled plants in the Edinburgh Botanic Gardens, and not one yielded a single seed; but eight of them which were fertilised with pollen from one of the other plants of the same form, set two empty capsules. (1/19.

See also Trevira.n.u.s in 'Botanische Zeitung' 1863 page 6 on this plant being dimorphic.) He was able to examine only dried specimens of the long-styled forms. But the evidence seems sufficient to leave hardly a doubt that Androsace is heterostyled. Fritz Muller sent me from South Brazil dried flowers of a Statice which he believed to be heterostyled. In the one form the pistil was considerably longer and the stamens slightly shorter than the corresponding organs in the other form. But as in the shorter-styled form the stigmas reached up to the anthers of the same flower, and as I could not detect in the dried specimens of the two forms any difference in their stigmas, or in the size of their pollen-grains, I dare not rank this plant as heterostyled. From statements made by Vaucher I was led to think that Soldanella alpina was heterostyled, but it is impossible that Kerner, who has closely studied this plant, could have overlooked the fact. So again from other statements it appeared probable that Pyrola might be heterostyled, but H. Muller examined for me two species in North Germany, and found this not to be the case.

CHAPTER II.

HYBRID PRIMULAS.

The oxlip a hybrid naturally produced between Primula veris and vulgaris.

The differences in structure and function between the two parent-species.

Effects of crossing long-styled and short-styled oxlips with one another and with the two forms of both parent-species.

Character of the offspring from oxlips artificially self-fertilised and cross- fertilised in a state of nature.

Primula elatior shown to be a distinct species.

Hybrids between other heterostyled species of Primula.

Supplementary note on spontaneously produced hybrids in the genus Verbasc.u.m.

The various species of Primula have produced in a state of nature throughout Europe an extraordinary number of hybrid forms. For instance, Professor Kerner has found no less than twenty-five such forms in the Alps. (2/1. "Die Primulaceen-Bastarten" 'Oesterr. Botanische Zeitschrift' Jahr 1875 Numbers 3, 4 and 5. See also G.o.dron on hybrid Primulas in 'Bull. Soc. Bot. de France' tome 10 1853 page 178. Also in 'Revue des Sciences Nat.' 1875 page 331.) The frequent occurrence of hybrids in this genus no doubt has been favoured by most of the species being heterostyled, and consequently requiring cross-fertilisation by insects; yet in some other genera, species which are not heterostyled and which in some respects appear not well adapted for hybrid-fertilisation, have likewise been largely hybridised. In certain districts of England, the common oxlip--a hybrid between the cowslip (P. veris, vel officinalis) and the primrose (P.

vulgaris, vel acaulis)--is frequently found, and it occurs occasionally almost everywhere. Owing to the frequency of this intermediate hybrid form, and to the existence of the Bardfield oxlip (P. elatior), which resembles to a certain extent the common oxlip, the claim of the three forms to rank as distinct species has been discussed oftener and at greater length than that of almost any other plant. Linnaeus considered P. veris, vulgaris and elatior to be varieties of the same species, as do some distinguished botanists at the present day; whilst others who have carefully studied these plants do not doubt that they are distinct species. The following observations prove, I think, that the latter view is correct; and they further show that the common oxlip is a hybrid between P. veris and vulgaris.

The cowslip differs so conspicuously in general appearance from the primrose, that nothing need here be said with respect to their external characters. (2/2.

The Reverend W.A. Leighton has pointed out certain differences in the form of the capsules and seed in 'Annals and Magazine of Natural History' 2nd series volume 2 1848 page 164.) But some less obvious differences deserve notice. As both species are heterostyled, their complete fertilisation depends on insects.

The cowslip is habitually visited during the day by the larger humble-bees (namely Bombus muscorum and hortorum), and at night by moths, as I have seen in the case of Cucullia. The primrose is never visited (and I speak after many years' observation) by the larger humble-bees, and only rarely by the smaller kinds; hence its fertilisation must depend almost exclusively on moths. There is nothing in the structure of the flowers of the two plants which can determine the visits of such widely different insects. But they emit a different odour, and perhaps their nectar may have a different taste. Both the long-styled and short-styled forms of the primrose, when legitimately and naturally fertilised, yield on an average many more seeds per capsule than the cowslip, namely, in the proportion of 100 to 55. When illegitimately fertilised they are likewise more fertile than the two forms of the cowslip, as shown by the larger proportion of their flowers which set capsules, and by the larger average number of seeds which the capsules contain. The difference also between the number of seeds produced by the long-styled and short-styled flowers of the primrose, when both are illegitimately fertilised, is greater than that between the number produced under similar circ.u.mstances by the two forms of the cowslip. The long-styled flowers of the primrose when protected from the access of all insects, except such minute ones as Thrips, yield a considerable number of capsules containing on an average 19.2 seeds per capsule; whereas 18 plants of the long-styled cowslip similarly treated did not yield a single seed.

The primrose, as every one knows, flowers a little earlier in the spring than the cowslip, and inhabits slightly different stations and districts. The primrose generally grows on banks or in woods, whilst the cowslip is found in more open places. The geographical range of the two forms is different. Dr.

Bromfield remarks that "the primrose is absent from all the interior region of northern Europe, where the cowslip is indigenous." (2/3. 'Phytologist' volume 3 page 694.) In Norway, however, both plants range to the same degree of north lat.i.tude. (2/4. H. Lecoq 'Geograph. Bot. de l'Europe' tome 8 1858 pages 141, 144. See also 'Annals and Magazine of Natural History' 9 1842 pages 156, 515.

Also Boreau 'Flore du centre de la France' 1840 tome 2 page 376. With respect to the rarity of P. veris in western Scotland, see H.C. Watson 'Cybele Britannica'

2 page 293.)

The cowslip and primrose, when intercrossed, behave like distinct species, for they are far from being mutually fertile. Gartner crossed 27 flowers of P.

vulgaris with pollen of P. veris, and obtained 16 capsules; but these did not contain any good seed. (2/5. 'b.a.s.t.a.r.derzeugung' 1849 page 721.) He also crossed 21 flowers of P. veris with pollen of P. vulgaris; and now he got only five capsules, containing seed in a still less perfect condition. Gartner knew nothing about heterostylism; and his complete failure may perhaps be accounted for by his having crossed together the same forms of the cowslip and primrose; for such crosses would have been of an illegitimate as well as of a hybrid nature, and this would have increased their sterility. My trials were rather more fortunate. Twenty-one flowers, consisting of both forms of the cowslip and primrose, were intercrossed legitimately, and yielded seven capsules (i.e. 33 per cent), containing on an average 42 seeds; some of these seeds, however, were so poor that they probably would not have germinated. Twenty-one flowers on the same cowslip and primrose plants were also intercrossed illegitimately, and they likewise yielded seven capsules (or 33 per cent), but these contained on an average only 13 good and bad seeds. I should, however, state that some of the above flowers of the primrose were fertilised with pollen from the polyanthus, which is certainly a variety of the cowslip, as may be inferred from the perfect fertility inter se of the crossed offspring from these two plants. (2/6. Mr.

Scott has discussed the nature of the polyanthus ('Proceedings of the Linnean Society' 8 Botany 1864 page 103), and arrives at a different conclusion; but I do not think that his experiments were sufficiently numerous. The degree of infertility of a cross is liable to much fluctuation. Pollen from the cowslip at first appears rather more efficient on the primrose than that of the polyanthus; for 12 flowers of both forms of the primrose, fertilised legitimately and illegitimately with pollen of the cowslip gave five capsules, containing on an average 32.4 seeds; whilst 18 flowers similarly fertilised by polyanthus-pollen yielded only five capsules, containing only 22.6 seeds. On the other hand, the seeds produced by the polyanthus-pollen were much the finest of the whole lot, and were the only ones which germinated.) To show how sterile these hybrid unions were I may remind the reader that 90 per cent of the flowers of the primrose fertilised legitimately with primrose-pollen yielded capsules, containing on an average 66 seeds; and that 54 per cent of the flowers fertilised illegitimately yielded capsules containing on an average 35.5 seeds per capsule. The primrose, especially the short-styled form, when fertilised by the cowslip, is less sterile, as Gartner likewise observed, than is the cowslip when fertilised by the primrose. The above experiments also show that a cross between the same forms of the primrose and cowslip is much more sterile than that between different forms of these two species.

The seeds from the several foregoing crosses were sown, but none germinated except those from the short-styled primrose fertilised with pollen of the polyanthus; and these seeds were the finest of the whole lot. I thus raised six plants, and compared them with a group of wild oxlips which I had transplanted into my garden. One of these wild oxlips produced slightly larger flowers than the others, and this one was identical in every character (in foliage, flower- peduncle, and flowers) with my six plants, excepting that the flowers of the latter were tinged of a dingy red colour, from being descended from the polyanthus.

We thus see that the cowslip and primrose cannot be crossed either way except with considerable difficulty, that they differ conspicuously in external appearance, that they differ in various physiological characters, that they inhabit slightly different stations and range differently. Hence those botanists who rank these plants as varieties ought to be able to prove that they are not as well fixed in character as are most species; and the evidence in favour of such instability of character appears at first sight very strong. It rests, first, on statements made by several competent observers that they have raised cowslips, primroses, and oxlips from seeds of the same plant; and, secondly, on the frequent occurrence in a state of nature of plants presenting every intermediate gradation between the cowslip and primrose.

The first statement, however, is of little value; for, heterostylism not being formerly understood, the seed-bearing plants were in no instance protected from the visits of insects (2/7. One author states in the 'Phytologist' volume 3 page 703 that he covered with bell-gla.s.ses some cowslips, primroses, etc., on which he experimented. He specifies all the details of his experiment, but does not say that he artificially fertilised his plants; yet he obtained an abundance of seed, which is simply impossible. Hence there must have been some strange error in these experiments, which may be pa.s.sed over as valueless.); and there would be almost as much risk of an isolated cowslip, or of several cowslips if consisting of the same form, being crossed by a neighbouring primrose and producing oxlips, as of one s.e.x of a dioecious plant, under similar circ.u.mstances, being crossed by the opposite s.e.x of an allied and neighbouring species. Mr. H.C. Watson, a critical and most careful observer, made many experiments by sowing the seeds of cowslips and of various kinds of oxlips, and arrived at the following conclusion, namely, "that seeds of a cowslip can produce cowslips and oxlips, and that seeds of an oxlip can produce cowslips, oxlips, and primroses." (2/8. 'Phytologist' 2 pages 217, 852; 3 page 43.) This conclusion harmonises perfectly with the view that in all cases, when such results have been obtained, the unprotected cowslips have been crossed by primroses, and the unprotected oxlips by either cowslips or primroses; for in this latter case we might expect, by the aid of reversion, which notoriously comes into powerful action with hybrids, that the two parent-forms in appearance pure, as well as many intermediate gradations, would be occasionally produced.

Nevertheless the two following statements offer considerable difficulty. The Reverend Professor Henslow raised from seeds of a cowslip growing in his garden, various kinds of oxlips and one perfect primrose; but a statement in the same paper perhaps throws light on this anomalous result. (2/9. Loudon's 'Magazine of Natural History' 3 1830 page 409.) Professor Henslow had previously transplanted into his garden a cowslip, which completely changed its appearance during the following year, and now resembled an oxlip. Next year again it changed its character, and produced, in addition to the ordinary umbels, a few single- flowered scapes, bearing flowers somewhat smaller and more deeply coloured than those of the common primrose. From what I have myself observed with oxlips, I cannot doubt that this plant was an oxlip in a highly variable condition, almost like that of the famous Cytisus adami. This presumed oxlip was propagated by offsets, which were planted in different parts of the garden; and if Professor Henslow took by mistake seeds from one of these plants, especially if it had been crossed by a primrose, the result would be quite intelligible. Another case is still more difficult to understand: Dr. Herbert raised, from the seeds of a highly cultivated red cowslip, cowslips, oxlips of various kinds, and a primrose. (2/10. 'Transactions of the Horticultural Society' 4 page 19.) This case, if accurately recorded, which I much doubt, is explicable only on the improbable a.s.sumption that the red cowslip was not of pure parentage. With species and varieties of many kinds, when intercrossed, one is sometimes strongly prepotent over the other; and instances are known of a variety crossed by another, producing offspring which in certain characters, as in colour, hairiness, etc., have proved identical with the pollen-bearing parent, and quite dissimilar to the mother-plant (2/11. I have given instances in my work 'On the Variation of Animals and Plants under Domestication' chapter 15 2nd edition volume 2 page 69.); but I do not know of any instance of the offspring of a cross perfectly resembling, in a considerable number of important characters, the father alone. It is, therefore, very improbable that a pure cowslip crossed by a primrose should ever produce a primrose in appearance pure. Although the facts given by Dr. Herbert and Professor Henslow are difficult to explain, yet until it can be shown that a cowslip or a primrose, carefully protected from insects, will give birth to at least oxlips, the cases. .h.i.therto recorded have little weight in leading us to admit that the cowslip and primrose are varieties of one and the same species.

Negative evidence is of little value; but the following facts may be worth giving:--Some cowslips which had been transplanted from the fields into a shrubbery were again transplanted into highly manured land. In the following year they were protected from insects, artificially fertilised, and the seed thus procured was sown in a hotbed. The young plants were afterwards planted out, some in very rich soil, some in stiff poor clay, some in old peat, and some in pots in the greenhouse; so that these plants, 765 in number, as well as their parents, were subjected to diversified and unnatural treatment; but not one of them presented the least variation except in size--those in the peat attaining almost gigantic dimensions, and those in the clay being much dwarfed.

I do not, of course, doubt that cowslips exposed during SEVERAL successive generations to changed conditions would vary, and that this might occasionally occur in a state of nature. Moreover, from the law of a.n.a.logical variation, the varieties of any one species of Primula would probably in some cases resemble other species of the genus. For instance I raised a red primrose from seed from a protected plant, and the flowers, though still resembling those of the primrose, were borne during one season in umbels on a long foot-stalk like that of a cowslip.

With regard to the second cla.s.s of facts in support of the cowslip and primrose being ranked as mere varieties, namely, the well-ascertained existence in a state of nature of numerous linking forms (2/12. See an excellent article on this subject by Mr. H.C. Watson in the 'Phytologist' volume 3 page 43.):--If it can be shown that the common wild oxlip, which is intermediate in character between the cowslip and primrose, resembles in sterility and other essential respects a hybrid plant, and if it can further be shown that the oxlip, though in a high degree sterile, can be fertilised by either parent-species, thus giving rise to still finer gradational links, then the presence of such linking forms in a state of nature ceases to be an argument of any weight in favour of the cowslip and primrose being varieties, and becomes, in fact, an argument on the other side. The hybrid origin of a plant in a state of nature can be recognised by four tests: first, by its occurrence only where both presumed parent-species exist or have recently existed; and this holds good, as far as I can discover, with the oxlip; but the P. elatior of Jacq., which, as we shall presently see, const.i.tutes a distinct species, must not be confounded with the common oxlip. Secondly, by the supposed hybrid plant being nearly intermediate in character between the two parent-species, and especially by its resembling hybrids artificially made between the same two species. Now the oxlip is intermediate in character, and resembles in every respect, except in the colour of the corolla, hybrids artificially produced between the primrose and the polyanthus, which latter is a variety of the cowslip. Thirdly, by the supposed hybrids being more or less sterile when crossed inter se: but to try this fairly two distinct plants of the same parentage, and not two flowers on the same plant, should be crossed; for many pure species are more or less sterile with pollen from the same individual plant; and in the case of hybrids from heterostyled species the opposite forms should be crossed. Fourthly and lastly, by the supposed hybrids being much more fertile when crossed with either pure parent-species than when crossed inter se, but still not as fully fertile as the parent-species.

For the sake of ascertaining the two latter points, I transplanted a group of wild oxlips into my garden. They consisted of one long-styled and three short- styled plants, which, except in the corolla of one being slightly larger, resembled each other closely. The trials which were made, and the results obtained, are shown in tables 2.14, 2.15, 2.16, 2.17 and 2.18. No less than twenty different crosses are necessary in order to ascertain fully the fertility of hybrid heterostyled plants, both inter se and with their two parent-species.

In this instance 256 flowers were crossed in the course of four seasons. I may mention, as a mere curiosity, that if any one were to raise hybrids between two trimorphic heterostyled species, he would have to make 90 distinct unions in order to ascertain their fertility in all ways; and as he would have to try at least 10 flowers in each case, he would be compelled to fertilise 900 flowers and count their seeds. This would probably exhaust the patience of the most patient man.

TABLE 2.14. Crosses inter se between the two forms of the common Oxlip.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled oxlip: 20 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled oxlip: 10 flowers fertilised, did not produce one capsule.

Column 3: Illegitimate union.

Long-styled oxlip, by its own pollen: 24 flowers fertilised, produced five capsules, containing 6, 10, 20, 8, and 14 seeds. Average 11.6.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled oxlip: 10 flowers fertilised, did not produce one capsule.

TABLE 2.15. Both forms of the Oxlip crossed with Pollen of both forms of the Cowslip, P. veris.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled cowslip: 18 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled cowslip: 18 flowers fertilised, produced three capsules, containing 7, 3, and 3 wretched seeds, apparently incapable of germination.

Column 3: Illegitimate union.

Long-styled oxlip, by pollen of long-styled cowslip: 11 flowers fertilised, produced one capsule, containing 13 wretched seeds.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled cowslip: 5 flowers fertilised, produced two capsules, containing 21 and 28 very fine seeds.

TABLE 2.16. Both forms of the Oxlip crossed with Pollen of both forms of the Primrose, P. vulgaris.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled primrose: 34 flowers fertilised, produced two capsules, containing 5 and 12 seeds.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled primrose: 26 flowers fertilised, produced six capsules, containing 16, 20, 5, 10, 19, and 24 seeds. Average 15.7.

Many of the seeds very poor, some good.

Column 3: Illegitimate union.

Long-styled oxlip, by pollen of long-styled primrose: 11 flowers fertilised, produced four capsules, containing 10, 7, 5, and 6 wretched seeds. Average 7.0.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled primrose: 5 flowers fertilised, produced five capsules, containing 26, 32, 23, 28, and 34 seeds. Average 28.6.

TABLE 2.17. Both forms of the Cowslip crossed with Pollen of both forms of the Oxlip.