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 classified them. My son did the same in Hampshire, and here is the result:—
TABLE 4.22. Lythrum salicaria. Classification 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 passage 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.