Their power and complex manner of fertilising one another.

Eighteen different unions possible.

Mid-styled form eminently feminine in nature.

Lythrum Graefferi likewise trimorphic.

L. thymifolia dimorphic.

L. Hyssopifolia h.o.m.ostyled.

Nesaea verticillata trimorphic.

Lagerstroemia, nature doubtful.

Oxalis, trimorphic species of.

O. Valdiviana.

O. Regnelli, the illegitimate unions quite barren.

O. speciosa.

O. sensitiva.

h.o.m.ostyled species of Oxalis.

Pontederia, the one monocotyledonous genus known to include heterostyled species.

In the previous chapters various heterostyled dimorphic plants have been described, and now we come to heterostyled trimorphic plants, or those which present three forms. These have been observed in three families, and consist of species of Lythrum and of the allied genus Nesaea, of Oxalis and Pontederia. In their manner of fertilisation these plants offer a more remarkable case than can be found in any other plant or animal.

Lythrum salicaria.

(FIGURE 4.10. Diagram of the flowers of the three forms of Lythrum salicaria, in their natural position, with the petals and calyx removed on the near side: enlarged six times.

Top: Long-styled.

Middle: Mid-styled.

Bottom: Short-styled.

The dotted lines with the arrows show the directions in which pollen must be carried to each stigma to ensure full fertility.)

The pistil in each form differs from that in either of the other forms, and in each there are two sets of stamens different in appearance and function. But one set of stamens in each form corresponds with a set in one of the other two forms. Altogether this one species includes three females or female organs and three sets of male organs, all as distinct from one another as if they belonged to different species; and if smaller functional differences are considered, there are five distinct sets of males. Two of the three hermaphrodites must coexist, and pollen must be carried by insects reciprocally from one to the other, in order that either of the two should be fully fertile; but unless all three forms coexist, two sets of stamens will be wasted, and the organisation of the species, as a whole, will be incomplete. On the other hand, when all three hermaphrodites coexist, and pollen is carried from one to the other, the scheme is perfect; there is no waste of pollen and no false co-adaptation. In short, nature has ordained a most complex marriage-arrangement, namely a triple union between three hermaphrodites,--each hermaphrodite being in its female organ quite distinct from the other two hermaphrodites and partially distinct in its male organs, and each furnished with two sets of males.

The three forms may be conveniently called, from the unequal lengths of their pistils, the LONG-STYLED, MID-STYLED, and SHORT-STYLED. The stamens also are of unequal lengths, and these may be called the LONGEST, MID-LENGTH, and SHORTEST.

Two sets of stamens of different length are found in each form. The existence of the three forms was first observed by Vaucher, and subsequently more carefully by Wirtgen ; but these botanists, not being guided by any theory or even suspicion of their functional differences, did not perceive some of the most curious points of difference in their structure. (4/1. Vaucher 'Hist. Phys. des Plantes d'Europe' tome 2 1841 page 371. Wirtgen "Ueber Lythrum salicaria und dessen Formen" 'Verhand. des naturhist. Vereins fur preuss. Rheinl.' 5 Jahrgang 1848 S. 7.) I will first briefly describe the three forms by the aid of Figure 4.10, which shows the flowers, six times magnified, in their natural position, with their petals and calyx on the near side removed.

LONG-STYLED FORM.

This form can be at once recognised by the length of the pistil, which is (including the ovarium) fully one-third longer than that of the mid-styled, and more than thrice as long as that of the short-styled form. It is so disproportionately long, that it projects in the bud through the folded petals.

It stands out considerably beyond the mid-length stamens; its terminal portion depends a little, but the stigma itself is slightly upturned. The globular stigma is considerably larger than that of the other two forms, with the papillae on its surface generally longer. The six mid-length stamens project about two-thirds the length of the pistil, and correspond in length with the pistil of the mid-styled form. Such correspondence in this and the two following forms is generally very close; the difference, where there is any, being usually in a slight excess of length in the stamens. The six shortest stamens lie concealed within the calyx; their ends are turned up, and they are graduated in length, so as to form a double row. The anthers of these stamens are smaller than those of the mid-length ones. The pollen is of the same yellow colour in both sets. H. Muller measured the pollen-grain in all three forms, and his measurements are evidently more trustworthy than those which I formerly made, so I will give them. (4/2. 'Die Befruchtung der Blumen' 1873 page 193.) The numbers refer to divisions of the micrometer equalling 1/300 millimetres. The grains, distended with water, from the mid-length stamens are 7 to 7 1/2, and those from the shortest stamens 6 to 6 1/2 in diameter, or as 100 to 86. The capsules of this form contain on an average 93 seeds: how this average was obtained will presently be explained. As these seeds, when cleaned, seemed larger than those from the mid-styled or short-styled forms, 100 of them were placed in a good balance, and by the double method of weighing were found to equal 121 seeds of the mid-styled or 142 of the short-styled; so that five long-styled seeds very nearly equal six mid-styled or seven short-styled seeds.

MID-STYLED FORM.

The pistil occupies the position represented in Figure 4.10, with its extremity considerably upturned, but to a variable degree; the stigma is seated between the anthers of the longest and the shortest stamens. The six longest stamens correspond in length with the pistil of the long-styled form; their filaments are coloured bright pink; the anthers are dark-coloured, but from containing bright-green pollen and from their early dehiscence they appear emerald-green.

Hence in general appearance these stamens are remarkably dissimilar from the mid-length stamens of the long-styled form. The six shortest stamens are enclosed within the calyx, and resemble in all respects the shortest stamens of the long-styled form; both these sets correspond in length with the short pistil of the short-styled form. The green pollen-grains of the longest stamens are 9 to 10 in diameter, whilst the yellow grains from the shortest stamens are only 6; or as 100 to 63. But the pollen-grains from different plants appeared to me, in this case and others, to be in some degree variable in size. The capsules contain on an average 130 seeds; but perhaps, as we shall see, this is rather too high an average. The seeds themselves, as before remarked, are smaller than those of the long-styled form.

SHORT-STYLED FORM.

The pistil is here very short, not one-third of the length of that of the long- styled form. It is enclosed within the calyx, which, differently from that in the other two forms, does not enclose any anthers. The end of the pistil is generally bent upwards at right angles. The six longest stamens, with their pink filaments and green pollen, resemble the corresponding stamens of the mid-styled form. But according to H. Muller, their pollen-grains are a little larger, namely 9 1/2 to 10 1/2, instead of 9 to 10 in diameter. The six mid-length stamens, with their uncoloured filaments and yellow pollen, resemble in the size of their pollen-grains and in all other respects the corresponding stamens of the long-styled form. The difference in diameter between the grains from the two sets of anthers in the short-styled form is as 100 to 73. The capsules contain fewer seeds on an average than those of either of the preceding forms, namely 83.5; and the seeds are considerably smaller. In this latter respect, but not in number, there is a gradation parallel to that in the length of the pistil, the long-styled having the largest seeds, the mid-styled the next in size, and the short-styled the smallest.

We thus see that this plant exists under three female forms, which differ in the length and curvature of the style, in the size and state of the stigma, and in the number and size of the seed. There are altogether thirty-six males or stamens, and these can be divided into three sets of a dozen each, differing from one another in length, curvature, and colour of the filaments--in the size of the anthers, and especially in the colour and diameter of the pollen-grains.

Each form bears half-a-dozen of one kind of stamens and half-a-dozen of another kind, but not all three kinds. The three kinds of stamens correspond in length with the three pistils: the correspondence is always between half of the stamens in two of the forms with the pistil of the third form. Table 4.a of the diameters of the pollen-grains, after immersion in water, from both sets of stamens in all three forms is copied from H. Muller; they are arranged in the order of their size:--

TABLE 4.a. Lythrum salicaria. Diameters of pollen-grains after immersion in water.

Column 1: Source of Pollen-grains.

Column 2: Minimum diameter.

Column 3: Maximum diameter.

Longest stamens of short-styled form : 9 1/2 : 10 1/2.

Longest stamens of mid-styled form : 9 : 10.

Mid-length stamens of long-styled form : 7 : 7 1/2.

Mid-length stamens of short-styled form : 7 : 7 1/2.

Shortest stamens of long-styled form : 6 : 6 1/2.

Shortest stamens of mid-styled form : 6 : 6.

We here see that the largest pollen-grains come from the longest stamens, and the least (smallest) from the shortest; the extreme difference in diameter between them being as 100 to 60.

The average number of seeds in the three forms was ascertained by counting them in eight fine selected capsules taken from plants growing wild, and the result was, as we have seen, for the long-styled (neglecting decimals) 93, mid-styled 130, and short-styled 83. I should not have trusted in these ratios had I not possessed a number of plants in my garden which, owing to their youth, did not yield the full complement of seed, but were of the same age and grew under the same conditions, and were freely visited by bees. I took six fine capsules from each, and found the average to be for the long-styled 80, for the mid-styled 97, and for the short-styled 61. Lastly, legitimate unions effected by me between the three forms gave, as may be seen in the following tables, for the long- styled an average of 90 seeds, for the mid-styled 117, and for the short-styled 71. So that we have good concurrent evidence of a difference in the average production of seed by the three forms. To show that the unions effected by me often produced their full effect and may be trusted, I may state that one mid- styled capsule yielded 151 good seeds, which is the same number as in the finest wild capsule which I examined. Some artificially fertilised short- and long- styled capsules produced a greater number of seeds than was ever observed by me in wild plants of the same forms, but then I did not examine many of the latter.

This plant, I may add, offers a remarkable instance, how profoundly ignorant we are of the life-conditions of a species. Naturally it grows "in wet ditches, watery places, and especially on the banks of streams," and though it produces so many minute seeds, it never spreads on the adjoining land; yet, when planted in my garden, on clayey soil lying over chalk, and which is so dry that a rush cannot be found, it thrives luxuriantly, grows to above 6 feet in height, produces self-sown seedlings, and (which is a severer test) is as fertile as in a state of nature. Nevertheless it would be almost a miracle to find this plant growing spontaneously on such land as that in my garden.

According to Vaucher and Wirtgen, the three forms coexist in all parts of Europe. Some friends gathered for me in North Wales a number of twigs from separate plants growing near one another, and cla.s.sified them. My son did the same in Hampshire, and here is the result:--

TABLE 4.22. Lythrum salicaria. Cla.s.sification according to form of flower.

Column 1: Place of origin.

Column 2: Long-styled.

Column 3: Mid-styled.

Column 4: Short-styled.

Column 5: Total.

North Wales : 95 : 97 : 72 : 264.

Hampshire : 53 : 38 : 38 : 129.

Total : 148 : 135 : 110 : 393.

If twice or thrice the number had been collected, the three forms would probably have been found nearly equal; I infer this from considering the above figures, and from my son telling me that if he had collected in another spot, he felt sure that the mid-styled plants would have been in excess. I several times sowed small parcels of seed, and raised all three forms; but I neglected to record the parent-form, excepting in one instance, in which I raised from short-styled seed twelve plants, of which only one turned out long-styled, four mid-styled, and seven short-styled.

Two plants of each form were protected from the access of insects during two successive years, and in the autumn they yielded very few capsules and presented a remarkable contrast with the adjoining uncovered plants, which were densely covered with capsules. In 1863 a protected long-styled plant produced only five poor capsules; two mid-styled plants produced together the same number; and two short-styled plants only a single one. These capsules contained very few seeds; yet the plants were fully productive when artificially fertilised under the net.

In a state of nature the flowers are incessantly visited for their nectar by hive- and other bees, various Diptera and Lepidoptera. (4/3. H. Muller gives a list of the species 'Die Befruchtung der Blumen' page 196. It appears that one bee, the Cilissa melanura, almost confines its visits to this plant.) The nectar is secreted all round the base of the ovarium; but a pa.s.sage is formed along the upper and inner side of the flower by the lateral deflection (not represented in the diagram) of the basal portions of the filaments; so that insects invariably alight on the projecting stamens and pistil, and insert their proboscides along the upper and inner margin of the corolla. We can now see why the ends of the stamens with their anthers, and the ends of the pistils with their stigmas, are a little upturned, so that they may be brushed by the lower hairy surfaces of the insects' bodies. The shortest stamens which lie enclosed within the calyx of the long- and mid-styled forms can be touched only by the proboscis and narrow chin of a bee; hence they have their ends more upturned, and they are graduated in length, so as to fall into a narrow file, sure to be raked by the thin intruding proboscis. The anthers of the longer stamens stand laterally farther apart and are more nearly on the same level, for they have to brush against the whole breadth of the insect's body. In very many other flowers the pistil, or the stamens, or both, are rectangularly bent to one side of the flower. This bending may be permanent, as with Lythrum and many others, or may be effected, as in Dictamnus fraxinella and others, by a temporary movement, which occurs in the case of the stamens when the anthers dehisce, and in the case of the pistil when the stigma is mature; but these two movements do not always take place simultaneously in the same flower. Now I have found no exception to the rule, that when the stamens and pistil are bent, they bend to that side of the flower which secretes nectar, even though there be a rudimentary nectary of large size on the opposite side, as in some species of Corydalis. When nectar is secreted on all sides, they bend to that side where the structure of the flower allows the easiest access to it, as in Lythrum, various Papilionaceae, and others. The rule consequently is, that when the pistils and stamens are curved or bent, the stigma and anthers are thus brought into the pathway leading to the nectary.

There are a few cases which seem to be exceptions to this rule, but they are not so in truth; for instance, in the Gloriosa lily, the stigma of the grotesque and rectangularly bent pistil is brought, not into any pathway from the outside towards the nectar-secreting recesses of the flower, but into the circular route which insects follow in proceeding from one nectary to the other. In Scrophularia aquatica the pistil is bent downwards from the mouth of the corolla, but it thus strikes the pollen-dusted breast of the wasps which habitually visit these ill-scented flowers. In all these cases we see the supreme dominating power of insects on the structure of flowers, especially of those which have irregular corollas. Flowers which are fertilised by the wind must of course be excepted; but I do not know of a single instance of an irregular flower which is thus fertilised.

Another point deserves notice. In each of the three forms two sets of stamens correspond in length with the pistils in the other two forms. When bees suck the flowers, the anthers of the longest stamens, bearing the green pollen, are rubbed against the abdomen and the inner sides of the hind legs, as is likewise the stigma of the long-styled form. The anthers of the mid-length stamens and the stigma of the mid-styled form are rubbed against the under side of the thorax and between the front pair of legs. And, lastly, the anthers of the shortest stamens and the stigma of the short-styled form are rubbed against the proboscis and chin: for the bees in sucking the flowers insert only the front part of their heads into the flower. On catching bees, I observed much green pollen on the inner sides of the hind legs and on the abdomen, and much yellow pollen on the under side of the thorax. There was also pollen on the chin, and, it may be presumed, on the proboscis, but this was difficult to observe. I had, however, independent proof that pollen is carried on the proboscis; for a small branch of a protected short-styled plant (which produced spontaneously only two capsules) was accidentally left during several days pressing against the net, and bees were seen inserting their proboscides through the meshes, and in consequence numerous capsules were formed on this one small branch. From these several facts it follows that insects will generally carry the pollen of each form from the stamens to the pistil of corresponding length; and we shall presently see the importance of this adaptation. It must not, however, be supposed that the bees do not get more or less dusted all over with the several kinds of pollen; for this could be seen to occur with the green pollen from the longest stamens. Moreover a case will presently be given of a long-styled plant producing an abundance of capsules, though growing quite by itself, and the flowers must have been fertilised by their own kinds of pollen; but these capsules contained a very poor average of seed. Hence insects, and chiefly bees, act both as general carriers of pollen, and as special carriers of the right sort.

Wirtgen remarks on the variability of this plant in the branching of the stem, in the length of the bracteae, size of the petals, and in several other characters. (4/4. 'Verhand. des naturhist. Vereins fur Pr. Rheinl.' 5 Jahrgang 1848 pages 11, 13.) The plants which grew in my garden had their leaves, which differed much in shape, arranged oppositely, alternately, or in whorls of three.

In this latter case the stems were hexagonal; those of the other plants being quadrangular. But we are concerned chiefly, with the reproductive organs: the upward bending of the pistil is variable, and especially in the short-styled form, in which it is sometimes straight, sometimes slightly curved, but generally bent at right angles. The stigma of the long-styled pistil frequently has longer papillae or is rougher than that of the mid-styled, and the latter than that of the short-styled; but this character, though fixed and uniform in the two forms of Primula veris, etc., is here variable, for I have seen mid- styled stigmas rougher than those of the long-styled. (4/5. The plants which I observed grew in my garden, and probably varied rather more than those growing in a state of nature. H. Muller has described the stigmas of all three forms with great care, and he appears to have found the stigmatic papillae differing constantly in length and structure in the three forms, being longest in the long-styled form.) The degree to which the longest and mid-length stamens are graduated in length and have their ends upturned is variable; sometimes all are equally long. The colour of the green pollen in the longest stamens is variable, being sometimes pale greenish-yellow; in one short-styled plant it was almost white. The grains vary a little in size: I examined one short-styled plant with the grains from the mid-length and shortest anthers of the same size. We here see great variability in many important characters; and if any of these variations were of service to the plant, or were correlated with useful functional differences, the species is in that state in which natural selection might readily do much for its modification.

ON THE POWER OF MUTUAL FERTILISATION BETWEEN THE THREE FORMS.

Nothing shows more clearly the extraordinary complexity of the reproductive system of this plant, than the necessity of making eighteen distinct unions in order to ascertain the relative fertilising power of the three forms. Thus the long-styled form has to be fertilised with pollen from its own two kinds of anthers, from the two in the mid-styled, and from the two in the short-styled form. The same process has to be repeated with the mid-styled and short-styled forms. It might have been thought sufficient to have tried on each stigma the green pollen, for instance, from either the mid- or short-styled longest stamens, and not from both; but the result proves that this would have been insufficient, and that it was necessary to try all six kinds of pollen on each stigma. As in fertilising flowers there will always be some failures, it would have been advisable to have repeated each of the eighteen unions a score of times; but the labour would have been too great; as it was, I made 223 unions, i.e. on an average I fertilised above a dozen flowers in the eighteen different methods. Each flower was castrated; the adjoining buds had to be removed, so that the flowers might be safely marked with thread, wool, etc.; and after each fertilisation the stigma was examined with a lens to see that there was sufficient pollen on it. Plants of all three forms were protected during two years by large nets on a framework; two plants were used during one or both years, in order to avoid any individual peculiarity in a particular plant. As soon as the flowers had withered, the nets were removed; and in the autumn the capsules were daily inspected and gathered, the ripe seeds being counted under the microscope. I have given these details that confidence may be placed in the following tables, and as some excuse for two blunders which, I believe, were made. These blunders are referred to, with their probable cause, in two footnotes to the tables. The erroneous numbers, however, are entered in the tables, that it may not be supposed that I have in any one instance tampered with the results.

A few words explanatory of the three tables must be given. Each is devoted to one of the three forms, and is divided into six compartments. The two upper ones in each table show the number of good seeds resulting from the application to the stigma of pollen from the two sets of stamens which correspond in length with the pistil of that form, and which are borne by the other two forms. Such unions are of a legitimate nature. The two next lower compartments show the result of the application of pollen from the two sets of stamens, not corresponding in length with the pistil, and which are borne by the other two forms. These unions are illegitimate. The two lowest compartments show the result of the application of each form's own two kinds of pollen from the two sets of stamens belonging to the same form, and which do not equal the pistil in length. These unions are likewise illegitimate. The term own-form pollen here used does not mean pollen from the flower to be fertilised--for this was never used--but from another flower on the same plant, or more commonly from a distinct plant of the same form. The figure "0" means that no capsule was produced, or if a capsule was produced that it contained no good seed. In some part of each row of figures in each compartment, a short horizontal line may be seen; the unions above this line were made in 1862, and below it in 1863. It is of importance to observe this, as it shows that the same general result was obtained during two successive years; but more especially because 1863 was a very hot and dry season, and the plants had occasionally to be watered. This did not prevent the full complement of seed being produced from the more fertile unions; but it rendered the less fertile ones even more sterile than they otherwise would have been. I have seen striking instances of this fact in making illegitimate and legitimate unions with Primula; and it is well known that the conditions of life must be highly favourable to give any chance of success in producing hybrids between species which are crossed with difficulty.

TABLE 4.23. Lythrum salicaria, long-styled form.

TABLE 4.23.1. Legitimate union.