Delphi Collected Works of Grant Allen, page 743
Fig. 35. — Single flower of Woodrush.
Among the rushes themselves, again, there are various levels of retrogressive development — retrogressive, that is to say, if we regard the lily family as an absolute standard; for the various alterations undergone by the different flowers are themselves adaptive to their new condition, though that condition is itself decidedly lower than the one from which they started. The common rush and its immediate congeners resemble the lilies from which they spring in having several seeds in each of the three cells which compose their pistil. But there is an interesting group of small grass-like plants, known as wood-rushes, which combine all the technical characteristics of the true rushes with a general character extremely like that of the grasses. They have long, thin, grass-like blades in the place of leaves; and what is still more important, as indicating an approach to the essentially one-seeded grass tribe, they have only three seeds in the flower, one to each cell of the capsule. These seeds are comparatively large, and are richly stored with food-stuffs for the supply of the young plantlet. One such richly supplied embryo is worth many little unsupported grains, since it stands a much better chance than they do of surviving in the struggle for existence. The wood-rushes may thus be regarded as some of the earliest plants among the great trinary class to adopt those tactics of storing gluten, starch, and other food-stuffs along with the embryo, which have given the cereals their acknowledged superiority as producers of human food. They are closely connected with the rushes, on the one hand, by sundry intermediate species which possess thin leaves instead of cylindrical pithy blades; and they lead on to the grasses, on the other, by reason of their very grass-like foliage, and their reduced number of large, well-furnished, starchy seeds.
In another particular, the rush family supplies us with a useful hint in tracing out the pedigree of the grasses and cereals. Their flowers are for the most part crowded together in large tufts or heads, each containing a considerable number of minute separate blossoms. Even among the true lilies we find some cases of such crowding in the hyacinths and the squills, or still better in the onion and garlic tribe. But with the wind-fertilised rushes, the grouping together of the flowers has important advantages, because it enables the pollen more easily to fix upon one or other of the sensitive surfaces, as the stalks sway backward and forward before a gentle breeze. Among yet more developed or degraded wind-fertilised plants, this crowding of the blossoms becomes even more conspicuous. A common American rush-like water-plant, known as eriocaulon, helps us to bridge over the gap between the rushes and such compound flowers as the sedges and grasses. Eriocaulon and its allies have always one seed only in each cell of the pistil: and they have also generally a very delicate corolla and calyx, of from four to six pieces, representing the original three sepals and three petals of the lilies and rushes. But their minute blossoms are closely crowded together in globular heads, the stamens and pistils being here divided in separate flowers, though both kinds of flowers are combined in each head. From an ancestral form not unlike this, but still more like the wood rushes, we must get both our sedges and our grasses. And though the sedges themselves do not stand in the direct line of descent to wheat and the other cereals, they are yet so valuable as an illustration from their points of analogy and of difference that we must turn aside for a moment to examine the gradual course of their evolution.
Fig. 36.
Fig. 37.
Single flower of Scirpus.
Male and Female flower of Carex.
The simplest and most primitive sedges now surviving, though very degenerate in type, yet retain some distinct traces of their derivation from earlier rush-like and lily-like ancestors. In the earliest existing type, known as scirpus, the calyx and petals which were brightly coloured in the lilies, and which were reduced to six brown scales in the rushes, have undergone a further degradation to the form of six small dry bristles, which now merely remain as rudimentary relics of a once useful and beautiful structure. In some species of scirpus, too, the number of these bristles is reduced from six to four or three. There is still one whorl of three stamens, however; but the second whorl has disappeared; while the pistil now contains only one seed instead of three; though it still retains some trace of the original three cells in the fact that there are three sensitive surfaces, united together at their base into one stalk or style. Each such diminution in the number of seeds is always accompanied by an increase in the effectiveness of those which remain; the difference is just analogous to that between the myriad ill-provided eggs of the cod, whose young fry are for the most part snapped up as soon as hatched, and the two or three eggs of birds, which watch their brood with such tender care, or the single young of cows, horses, and elephants, which guard their calves or foals almost up to the age of full maturity. What the bird or the animal effects by constant feeding with worms or milk, the plant effects by storing its seed with assorted food stuffs for the sprouting embryo.
In the more advanced or more degenerate sedges we get still further differentiation for the special function of wind-fertilisation. Take, as an example of these most developed types on this line of development, the common English group of carices (fig. 37). In these, the flowers have absolutely lost all trace of a perianth (that is to say, of the calyx and petals), for they do not possess even the six diminutive bristles which form the last relics of those organs in their allies, the scirpus group. Each flower is either male or female — that is to say, it consists of stamens or ovaries alone. The male flowers are represented by a single scale or bract, inclosing three stamens; and in some species even the stamens are reduced to a pair, so that all trace of the original trinary arrangement is absolutely lost. The female flowers are represented by a single ovary, inclosed in a sort of loose bag, which may perhaps be the final rudiment of a tubular bell-shaped corolla like that of the hyacinth. This ovary contains a single seed, but its shape is often triangular, and it has usually three stigmas or sensitive surfaces, thus dimly pointing back to the three distinct cells of its lily-like ancestors, and the three separate ovaries of its still earlier alisma-like progenitors. In many species, however, even this last souvenir of the trinary type has been utterly obliterated, the ovary having only two stigmas, and assuming a flattened two-sided shape. In all the carices, the flowers are loosely arranged in compact spikes and spikelets, with their mobile stamens hanging out freely to the breeze, and their feathery stigmas prepared to catch the slightest grain of pollen which may happen to be wafted their way by any passing breath of air. The varieties in their arrangement, however, are almost as infinite among the different species as those of the grasses themselves; sometimes the male and female flowers are produced on separate plants; sometimes they grow in separate spikes on the same plant; sometimes the same spike has male flowers at the top and female at the bottom; sometimes the various flowers are mixed up with one another at top and bottom in a regular hotch-potch of higgledy-piggledy confusion. But all the sedges alike are very grass-like in their aspect, with thin blades by way of leaves, and blossoms on tall heads as in the grasses. In fact, the two families are never accurately distinguished by any except technical botanists; to the ordinary observer, they are all grasses together, without petty distinctions of genus and species. Like the grasses, too, the sedges are mostly plants of the open wind-swept plains or marshy levels, where the facilities for wind-fertilisation are greatest and most constantly present.
And now, from this illustrative digression, let us hark back again to the junction point of the rushes, whence alike the sedges and the grasses appear to diverge. In order to understand the nature of the steps by which the cereals have been developed from rush-like ancestors, it will be necessary to look shortly at the actual composition of the flower in grasses, which is the only part of their organism differing appreciably from the ordinary lily type. The blossoms of grasses, in their simplest form, consist of several little green florets, arranged in small clusters, known as spikelets, along a single common axis. Of this arrangement, the head of wheat itself offers a familiar and excellent example. If we pull to pieces one of the spikelets composing such a head, we find it to consist of four or five distinct florets. Omitting special features and unnecessary details, we may say that each floret is made up of two chaffy scales (e, d), known as pales, and representing the calyx, together with a pair of small white petals (c) known as lodicules, three stamens (b), and an ovary with two feathery styles (a). Moreover, the two pales or calyx-pieces are not similar and symmetrical, for the outer one (e) is simple and convex, while the inner one (d) is apparently double, being made up of two pieces rolled into one, and still possessing two green midribs, which show distinctly like ribs on its flat outer surface. Here, it will immediately be apparent, the traces of the original trinary arrangement are very slight indeed.
Fig. 38.
Fig. 39.
Details of flower of Wheat.
Flower of Wheat (glumes removed).
But when we come to inquire into the rationale and genesis of these curiously one-sided flowers, it is not difficult to see that they have been ultimately derived from trinary blossoms of the rush-like type. The first and most marked divergence from that type, for which the analogy of the sedges has already prepared us, is the reduction of the ovary to a single one-seeded cell, whose ripe fruity form is known as a grain. At one time, we may feel pretty sure, there must have existed a group of nascent grasses, which only differed from the wood-rush genus in having a single-celled ovary instead of a three-celled pistil with one seed in each cell; and even the ovary of this primitive grass must have retained one mark of its trinary origin in its possession of three styles to its one grain, thus pointing back (as most sedges still do) to its earlier rush-like origin. That hypothetical form must have had three sepals, three petals, six stamens, and one three-styled ovary. But the peculiar shape of modern grass-flowers is clearly due to their very spiky arrangement along the edge of the axis. In the wood-rushes and the sedges, we see some approach to this condition; but in the grasses, the crowding is far more marked, and the one-sidedness has accordingly become far more conspicuous. Suppose we begin to crowd a number of wind-fertilised lily-like flowers along an axis in this manner, taking care that the stamens and the sensitive feathery styles are always turned outward to catch the breeze (for otherwise they will die out at once), what sort of result shall we finally get?
In the first place, the calyx, consisting of three pieces, will stand towards the crowded stem or axis in such a fashion that one piece will be free and exterior, while two pieces will be interior and next the stem, thus —
O a a a
Now, the effect of constant crushing in this direction will be that the two inner calyx-pieces will be slowly dwarfed, and will tend to coalesce with one another; and this is what has actually happened with the inner pale of wheat and of other grasses, though the midribs of the two originally separate pieces still show on the compound pale, like dark green lines down its centre. Thus, in the fully developed grasses, in place of a trinary calyx, we get two chaffy scales or pales, the outer one representing a single sepal, and the inner one, which has been dwarfed by pressure against the stem, representing two sepals rolled into one, with two midribs still remaining as evidence of their original distinctness.
Next, in the case of the petals, which alternate with the sepals of the calyx, the relation to the stem is exactly reversed; for we have here two petals free and exterior, with one interior petal crowded closely against the axis, thus —
O a a a
Here, then, the two external petals will be saved, exactly as the one external sepal was saved in the case of the calyx; and these two petals are represented by the very small white lodicules under the outer pale in our existing wheats and grasses. On the other hand, the inner petal, jammed in between the grain and the inner pale (with the stem at its back), has been utterly crushed out of existence, partly because of its very small size, partly because of its functional uselessness, and partly because it had no other part with which to coalesce, and so to save itself as the inner sepals had managed to do. Moreover, it must be remembered that the sepals do still perform a useful service in protecting the young flower before it opens, and in keeping out noxious insects during the kerning or swelling of the grain; whereas the lodicules or rudimentary petals are now apparently quite functionless; and so we may congratulate ourselves that they are there at all, to preserve for us the true ground-plan of the floral architecture in grasses. Indeed, they have not survived by any means in all grasses: among the smaller and more degraded kinds they are often wholly wanting, having been quite crushed out between the calyx and the grain. It is only the larger and more primitive types that still exhibit them in any great perfection. On the other hand, one group of very large exotic grasses, the bamboos, has three regular petals, thus clearly showing the descent of the family as a whole from rush-like ancestors, and also obviously suggesting that the obsolescence of the inner petal in the other grasses is due to their small size and their closely packed minute flowers.
Among the stamens, one-sidedness has not notably established itself, for in wind-fertilised plants they must necessarily hang out freely to the breeze, and therefore they do not get much crowded between the other parts. A few grasses still even retain their double row of stamens, having six to each floret; but most of them have only one whorl of three. In some of the lower and more degraded forms, however, even the stamens have lost their trinary order, and only two now survive. This is the case in our own very degenerate little sweet-vernal-grass, the plant which imparts its delicious fragrance to new-mown hay. But in the cereals and in most other large species the three stamens still remain in undiminished effectiveness to the present day.
Finally we come to the most important part of all, the ovary. This part, alternating with the stamens, has the same arrangement of styles relatively to the axis as in the case of the petals; and it has undergone precisely the same sort of abortive distortion. The two outer styles, hanging freely out of the calyx, have been preserved like the two outer lodicules; but the inner one, pressed between the grain and the inner pale (with the stem behind it) has been simply crushed out of existence, like its neighbour the inner lodicule.
Thus the final result is that the whole inner portion of the flower (except as regards stamens) has been distorted or rendered abortive by close pressure against the stem (due to the crowding of the florets in the spiky form), while the whole outer portion remains normal and fully developed. We have an outer pale representing a single normal sepal, and an inner pale representing two dwarfed and united sepals; we have two normal outer lodicules or petals, and a blank where the inner petal ought to be; we have three stamens, symmetrically arranged, among the faithless faithful only found; and we have finally two normal outer styles, with a blank in place of the absent inner style. The accompanying diagram, compared with that of the primitive monocotyledon (fig. 32), will make this perfectly clear.
Fig. 40. — Diagram of Wheat flower.
Here, a represents the outer pale or normal sepal, while a and a represent the inner pale composed of the two united sepals. Again, b and b stand for the two lodicules or surviving petals, while b marks the place of the lost petal, now found in the bamboos alone. The stamens are lettered c, c, and c. The two existing styles are shown by d and d, while d marks the abortive inner style, now not even present in a rudimentary condition. It will be observed at once that all the outer side is normal, and all the inner side more or less abortive through pressure against the axis.
Thus it will be seen that the line of links which connects the grasses and cereals with the lilies is absolutely unbroken, and that it consists throughout of one continuous course of degradation. At the same time, by this one-sided and spiky arrangement, the grasses secured for themselves an exceptional advantage in the struggle for existence. No other race of small wind-fertilised plants could compete with them for the possession of the open wind-swept plains; and over all these they spread far and wide, rapidly differentiating themselves into a vast number of divergent genera and species, each adaptively specialised for some peculiar habitat, soil, or climate. At the present time, the grasses number their kinds by thousands; they extend over the whole world from the poles to the equator; and they form the general sward or carpet of greenery over by far the larger portion of the terrestrial globe. Even in Britain alone, with our poor little insular flora, a mere fragment of that belonging to the petty European continent, we number no less than forty-two genera of grasses, distributed into more than one hundred species. In fact, what may fairly be called degradation from one point of view may fairly be called adaptation from another. The organisation of the grasses is certainly lower than that of the lilies, but it fits them better for that station of life to which it has pleased nature to assign them.
