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13
ANGIOSPERMS

albuminous Dicotyledons the cotyledons act as the absorbents of the reserve-food of the seed and are commonly brought above ground (epigeal), either withdrawn from the seed-coat or carrying it upon them, and then they serve as the first green organs of the plant. The part of the stem below the cotyledons (hypocotyl) commonly plays the greater part in bringing this about. Exalbuminous Dicotyledons usually store reserve-food in their cotyledons, which may in germination remain below ground (hypogeal). In albuminous Monocotyledons the cotyledon itself, probably in consequence of its terminal position, is commonly the agent by which the embryo is thrust out of the seed, and it may function solely as a feeder, its extremity developing as a sucker through which the endosperm is absorbed, or it may become the first green organ, the terminal sucker dropping off with the seed-coat when the endosperm is exhausted. Exalbuminous Monocotyledons are either hydrophytes or strongly hygrophilous plants and have often peculiar features in germination.

Distribution by seed appears to satisfy so well the requirements of Angiosperms that distribution by vegetative buds is only an occasional process. At the same time every bud on a Vegetative reproduction. shoot has the capacity to form a new plant if placed in suitable conditions, as the horticultural practice of propagation by cuttings shows; in nature we see plants spreading by the rooting of their shoots, and buds we know may be freely formed not only on stems but on leaves and on roots. Where detachable buds are produced, which can be transported through the air to a distance, each of them is an incipient shoot which may have a root, and there is always reserve-food stored in some part of it. In essentials such a bud resembles a seed. A relation between such vegetative distribution buds and production of flower is usually marked. Where there is free formation of buds there is little flower and commonly no seed, and the converse is also the case. Viviparous plants are an illustration of substitution of vegetative buds for flower.

The position of Angiosperms as the highest plant-group is unassailable, but of the point or points of their origin from the general stem of the plant kingdom, and of the path Phylogeny and taxonomy. or paths of their evolution, we can as yet say little. Until well on in the Mesozoic period geological history tells us nothing about Angiosperms, and then only by their vegetative organs. We readily recognize in them now-a-days the natural classes of Dicotyledons and Monocotyledons distinguished alike in vegetative and in reproductive construction, yet showing remarkable parallel sequences in development; and we see that the Dicotyledons are the more advanced and show the greater capacity for further progressive evolution. But there is no sound basis for the assumption that the Dicotyledons are derived from Monocotyledons; indeed, the palaeontological evidence seems to point to the Dicotyledons being the older. This, however, does not entitle us to assume the origin of Monocotyledons from Dicotyledons, although there is manifestly a temptation to connect helobic forms of the former with ranal ones of the latter. There is no doubt that the phylum of Angiosperms has not sprung from that of Gymnosperms.

Within each class the flower-characters as the essential feature of Angiosperms supply the clue to phylogeny, but the uncertainty regarding the construction of the primitive angiospermous flower gives a fundamental point of divergence in attempts to construct progressive sequences of the families. Simplicity of flower-structure has appeared to some to be always primitive, whilst by others it has been taken to be always derived. There is, however, abundant evidence that it may have the one or the other character in different cases. Apart from this, botanists are generally agreed that the concrescence of parts of the flower-whorls—in the gynaeceum as the seed-covering, and in the corolla as the seat of attraction, more than in the androecium and the calyx—is an indication of advance, as is also the concrescence that gives the condition of epigyny. Dorsiventrality is also clearly derived from radial construction, and anatropy of the ovule has followed atropy. We should expect the albuminous state of the seed to be an antecedent one to the exalbuminous condition, and the recent discoveries in fertilization tend to confirm this view. Amongst Dicotyledons the gamopetalous forms are admitted to be the highest development and a dominant one of our epoch. Advance has been along two lines, markedly in relation to insect-pollination, one of which has culminated in the hypogynous epipetalous bicarpellate forms with dorsiventral often large and loosely arranged flowers such as occur in Scrophulariaceae, and the other in the epigynous bicarpellate small-flowered families of which the Compositae represent the most elaborate type. In the polypetalous forms progression from hypogyny to epigyny is generally recognized, and where dorsiventrality with insect-pollination has been established, a dominant group has been developed as in the Leguminosae. The starting-point of the class, however, and the position within it of apetalous families with frequently unisexual flowers, have provoked much discussion. In Monocotyledons a similar advance from hypogyny to epigyny is observed, and from the dorsiventral to the radial type of flower. In this connexion it is noteworthy that so many of the higher forms are adapted as bulbous geophytes, or as aerophytes to special xerophilous conditions. The Gramineae offer a prominent example of a dominant self-pollinated or wind-pollinated family, and this may find explanation in a multiplicity of factors.

Though best known for his artificial (or sexual) system, Linnaeus was impressed with the importance of elaborating a natural system of arrangement in which plants should be arranged according to their true affinities. In his Philosophia Botanica (1751) Linnaeus grouped the genera then known into sixty-seven orders (fragmenta), all except five of which are Angiosperms. He gave names to these but did not characterize them or attempt to arrange them in larger groups. Some represent natural groups and had in several cases been already recognized by Ray and others, but the majority are, in the light of modern knowledge, very mixed. Well-defined polypetalous and gamopetalous genera sometimes occur in the same order, and even Monocotyledons and Dicotyledons are classed together where they have some striking physiological character in common.

Work on the lines suggested by the Linnaean fragmenta was continued in France by Bernard de Jussieu and his nephew, Antoine Laurent, and the arrangement suggested by the latter in his Genera Plantarum secundum Ordines Naturales disposita (1789) is the first which can claim to be a natural system. The orders are carefully characterized, and those of Angiosperms are grouped in fourteen classes under the two main divisions Monocotyledons and Dicotyledons. The former comprise three classes, which are distinguished by the relative position of the stamens and ovary; the eleven classes of the latter are based on the same set of characters and fall into the larger subdivisions Apetalae, Monopetalae and Polypetalae, characterized respectively by absence, union or freedom of the petals, and a subdivision, Diclines Irregulares, a very unnatural group, including one class only. A. P. de Candolle introduced several improvements into the system. In his arrangement the last subdivision disappears, and the Dicotyledons fall into two groups, a larger containing those in which both calyx and corolla are present in the flower, and a smaller, Monochlamydeae, representing the Apetalae and Diclines Irregulares of Jussieu. The dichlamydeous group is subdivided into three, Thalamiflorae, Calyciflorae and Corolliflorae, depending on the position and union of the petals. This, which we may distinguish as the French system, finds its most perfect expression in the classic Genera Plantarum (1862-1883) of Bentham and Hooker, a work containing a description, based on careful examination of specimens, of all known genera of flowering plants. The subdivision is as follows:-

Dicotyledons.
Polypetalae Thalamiflorae.
Disciflorae.
Calyciflorae.
Gamopetalae Inferae.
Heteromerae.
Bicarpellatae.
Monochlamydeae in eight series.
Monocotyledons in seven series.

Of the Polypetalae, series 1, Thalamiflorae, is characterized by hypogynous petals and stamens, and contains 34 orders distributed in 6 larger groups or cohorts. Series 2, Disciflorae, takes its name from a development of the floral axis which forms a ring or cushion at the base of the ovary or is broken up into glands; the ovary is superior. It contains 23 orders in 4 cohorts. Series 3, Calyciflorae, has petals and stamens perigynous, or sometimes superior. It contains 27 orders in 5 cohorts.

Of the Gamopetalae, series 1, Inferae, has an inferior ovary and stamens usually as many as the corolla-lobes. It contains 9 orders in 3 cohorts. Series 2, Heteromerae, has generally a superior ovary, stamens as many as the corolla-lobes or more, and more than two carpels. It contains 12 orders in 3 cohorts. Series 3, Bicarpellatae, has generally a superior ovary and usually two carpels. It contains 24 orders in 4 cohorts.

The eight series of Monochlamydeae, containing 36 orders, form groups characterized mainly by differences in the ovary and ovules, and are now recognized as of unequal value.

The seven series of Monocotyledons represent a sequence beginning with the most complicated epigynous orders, such as Orchideae and Scitamineae, and passing through the petaloid hypogynous orders (series Coronarieae) of which Liliaceae is the representative to Juncaceae and the palms (series Calycinae) where the perianth loses its petaloid character and thence to the Aroids, screw-pines and