Читать книгу Vegetable Teratology - Maxwell T. Masters - Страница 7

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As full details relating to the disposition or arrangement of the general organs of flowering plants are given in all the ordinary text-books, it is only necessary in this place to allude to the main facts at present known, and which serve as the standard of comparison with which all morphological changes are compared.

Even in the case of the roots, which appear to be very irregular in their ramification, it has been found that, in the first instance at least, the rootlets or fibrils are arranged in regular order one over another, in a certain determinate number of vertical ranks, generally either in two or in four, sometimes in three or in five series. This regularity of arrangement (Rhizotaxy), first carefully studied by M. Clos, is connected with the disposition of the fibro-vascular bundles in the body of the root. This primitive regularity is soon lost as the plant grows.

In the case of the leaves there are two principal modes of arrangement, dependent, as it would seem, on their simultaneous or on their successive development; thus, if two leaves on opposite sides of the stem are developed at the same time, we have the arrangement called opposite; if there are more than two, the disposition is then called verticillate or whorled. On the other hand, if the leaves are developed in succession, one after the other, they are found to emerge from the stem in a spiral direction. In either case the leaves are arranged in a certain regular manner, according to what are called the laws of Phyllotaxis, which need not be entered into fully here; but in order the better to estimate the teratological changes which take place, it may be well to allude to the following circumstances relating to the alternation of parts. The effect of this alternation is such, that no two adjacent leaves stand directly over or in front one of the other, but a little to one side or a little higher up. Now, in the alternate arrangement the successive leaves of each spiral cycle alternate one with another till the coil is completed. For the sake of clearness this may be illustrated thus:—Suppose the spiral cycle to comprise five leaves, numbered 1, 2, 3, 4, 5, then 2 would intervene between 1 and 3, and so on, while the sixth leaf would be the commencement of a new series, and would be placed exactly over 1. This arrangement may be thus formularised:

6 7 8 9 10

1 2 3 4 5

In the verticillate or simultaneous arrangement of leaves the case is somewhat different. Let us suppose a whorl of eight leaves, surmounted by a similar whorl of eight. In such a case it will generally be found that the whorls alternate one with another, as may be represented by this symbol:

9 10 11 12 13 14 15 16

1 2 3 4 5 6 7 8

The simplest illustration of this arrangement is seen in the case of decussate leaves, where those organs are placed in pairs, and the pairs cross one another at right angles. This may be expressed by the following symbol:

7 8

5 6

3 4

1 2

Thus, while in both the annular and the spiral modes of development the individual members of each complete series necessarily alternate one with another, in the former case the series themselves alternate, while in the successive arrangement they are placed directly one over the other. There are, of course, exceptions, but the rule is as has been stated, and the effect is to prevent one leaf from interfering with the development and growth of its neighbours.

In the case of the whorled or simultaneous arrangement the conditions of growth must be uniform on all sides, but in the successive or spiral disposition the conditions influencing growth act with unequal force, on different sides of the stem, at the same time. In the whorl there is an illustration of radiating symmetry, while in the spiral arrangement there is a transition to the bilateral symmetry. There are frequent passages from one to the other even under normal circumstances; thus, while the one arrangement obtains in the ordinary leaves, the parts of the flower may be disposed according to the other method. In the annular disposition it generally happens that the rings are separated one from the other by the development of the stem between them, the internodes between the constituent leaves themselves of course being undeveloped; on the other hand, in the spiral or successive arrangement there is no such alternate growth and arrest of growth of the stem between the leaves, or between successive cycles, but the growth is, under favorable conditions, continuous—leaf is separated from leaf, and cycle from cycle, by the continually elongating stem. Thus, the two modes of growth correspond precisely with those observed in the case of definite and indefinite inflorescence respectively.

Fig. 1.—Diagram showing the arrangement of parts in a complete, regular, pentamerous flower: s, sepals; p, petals; st, stamens; o, ovaries.

The same arrangements, that are observed in the disposition of the leaves, apply equally well to the several parts of the flower; thus, in what is for convenience considered the typical flower, there is a calyx of five or more distinct sepals, equal in size, and arranged in a whorl, a corolla of a similar number of petals alternating with the sepals, five stamens placed in the same position with reference to the petals, and five carpels alternating with the stamens. Throughout this book this arrangement is taken as the standard of reference. Nevertheless the spiral order does occur in the floral leaves as well as in those of the stem; it often happens, especially when the organs are numerous, that they form spiral series; and the same holds good very generally, when the parts of the flower are uneven in number, as in the very common quincuncial arrangement of the sepals, &c.

To these general remarks, intended to show the agreement between the disposition of the leaves of the stem and those of the flower, it is merely necessary to add that the arrangement of the placentas, as well as that of the ovules borne on them, is also definite, and takes place according to methods explained in all the text-books, and on which, therefore, it is not necessary to dilate in this place.

The branches of the stem or axis correspond for the most part in disposition with that of the leaves from the axils of which they originate, subject, however, to numerous disturbing causes, and to alterations from the usual or typical order brought about by the development of buds. These latter organs, as it seems, may be found in almost any situation, though their ordinary position is in the axil of a leaf or at the end of a stem or branch.

The points just mentioned are of primary importance in structural botany, and as such are seized on not only by the morphologist, but by the systematic botanist, who finds in them the characters by which he may separate one group from another. Thanks to the labours of those observers who have devoted their attention to that difficult but most important branch of study, organogeny, or the investigation of the development of the various organs, and to the researches of the students of comparative anatomy or morphology, the main principles regulating the arrangement and form of the organs of flowering plants seem to be fairly well established, though in matters of detail much remains to be cleared up, even in such important points as the share which the axis takes in the construction of the flower and fruit, the nature of the placenta, the construction of the ovules, and other points.

The facts already known justify the adoption of a standard or typical arrangement as just mentioned. The intrinsic value of this type is shown by the facility with which all varieties of form or arrangement may be explained by reference to certain modifications of it. It must, however, be considered as an abstraction, and should be looked on in the light rather of a scaffolding, which enables us to see the building and its several parts, than of the edifice itself, but which latter, from our imperfect knowledge and limited powers, we could not see without some such assistance.

The typical form may be, hypothetically at least, considered as the primitive one transmitted by hereditary descent from generation to generation, and modified to suit the requirements of the individual, or in accordance with circumstances. If it be borne in mind that it is but an artificial contrivance, more or less true—a means to an end, and not the end itself—no harm will arise from its employment; and as knowledge increases, or as circumstances demand, the hypothetical type can be replaced by another more in accordance with the actual state of science.

Teratological changes in the arrangement of organs depend upon arrest of growth, as when parts usually spirally arranged remain verticillate, owing to the non-development of the internodes, or to excessive growth, or development; but in many instances it is impossible, without studying the development of the malformed flower, to ascertain whether the altered arrangement is due to an excessive or to a diminished action. Practically, however, it is of comparatively little importance to know whether, say, the isolation of parts, that are usually combined together, is congenital (i.e. the result of an arrest of growth preventing their union), or whether it be due to a separation of parts primitively undivided; the effect remains the same, though the cause may have been very different.

The principal alterations to be mentioned under this head may therefore be conveniently arranged under the following categories:—Union, Independence, Displacement, Prolification, Heterotaxy, and Heterogamy.