In some instances the a.s.sumption of a scale-like form by any organ is attended by a change in texture, the organs becoming dry and scarious, or fleshy. Moquin cites in ill.u.s.tration of the first phenomenon the flower of a _Vicia_, in which the petals were thick and fleshy, like the scales of a bulb; and of the second the leaves of a _Chrysanthemum_, which were replaced by small, glossy scales, like those which invest ordinary leaf-buds. Sometimes the entire flower is replaced by acc.u.mulations of small, acute, green scales. Cases of this kind, wherein the flowers of a pea and of the foxglove were replaced by collections of small ovate green scales packed one over the other till they resembled the strobile of a hop, have been already alluded to. Most of these scales are represented as having had other acc.u.mulations of scales in their axils.

Similar collections of scales may frequently be met with in the birch and in the oak, and probably represent abortive leaf-buds. Other cases of a like kind in _Gentiana Amarella_, where the scales are coloured, are mentioned elsewhere.

In some kinds of _Campanula_ a similar change is not uncommon.

=Formation of hairs, spines, &c.=--The advent.i.tious production of hairs is likewise frequently due to an arrested growth, in some cases arising from pressure impeding the proper development of the organ. In other cases the formation of hair seems to accompany the diminished development of some organ, as on the barren pedicels of the wig plant, _Rhus Cotinus_. A similar production of hair may be noticed in many cases where the development of a branch or of a flower is arrested, and this occurs with especial frequency where the arrest in growth is due to the puncture of an insect, or to the formation of a gall. In such cases the hairs are mere excrescences from the epidermis.

p.r.i.c.kles differ but little from hairs save in their more woody texture, but true spines or thorns are modifications either of a leaf or of a branch. Their presence seems often dependent on the soil in which the plants grow, or on other external circ.u.mstances.

They occur normally in the sepals of _Paronychia serpyllifolia_ and other plants.

=Formation of glands.=--Under this name are a.s.sociated a number of (generally) rudimentary organs very different in their morphological nature and significance, and also in their functions. Some are truly glandular or secreting organs, while others have no visible office.

Anything like a complete account of these structures would be out of place, and reference is only made to them here on account of the occasional existence of intermediate forms, which throw light on the morphological significance of these structures. Thus, in _Pa.s.siflora_ and _Viburnum Opulus_, the so-called glands on the sides of the petiole appear to represent leaflets, and are not unfrequently developed as such.

M. Dunal observed a flower of _Cistus v.a.g.i.n.atus_ in which some of the stamens were replaced by an hypogynous disc.[547] Moquin has seen similar instances in the flowers of a Rose, _Hyperic.u.m_, and Poppy.

M. Planchon[548] gives an account of some very curious malformations in _Drosera intermedia_, which go to show that the ovules are h.o.m.ologous with the glandular hairs on the margins of the leaves of these plants, an opinion corroborated by the researches of MM. Gronland and Trecul.[549]

Dr. Hooker shows that the pitcher of _Nepenthes_ is due to a modification of a gland placed at the extremity of the midrib.[550]

=Formation of tendrils.=--These are of very varied morphological import; sometimes they are degenerated peduncles, as in pa.s.sion-flowers, or vines; at other times they are of foliar origin; or, again, they may proceed from the segments of the perianth, as in _Hodgsonia_ and some other cucurbitaceous plants. From their very different origin in different plants it is necessary to study the development in each case, and not apply to the generality what may be peculiar to one. In any case this formation in question generally belongs more to general morphology than to teratology.[551]

Kirschleger, however, has recorded the existence of a cirrhose sepal in _Cucurbita Pepo_.[552]

FOOTNOTES:

[547] 'Consid. Org. Fleur.,' p. 44, pl. ii, fig. 23.

[548] 'Ann. Sc. Nat.,' 3 ser., Bot. ix, pl. 6, ff. 1, 2.

[549] 'Ann. Sc. Nat.,' 3 ser., Bot. 1855, pp. 297, et 303.

[550] 'Trans. Linn. Soc.,' xxii, p. 415.

[551] See Darwin, "On Climbing Plants," 'Journal of Linnean Society,'

vol. ix, p. 1.

[552] 'Flora,' 1845, p. 615.

GENERAL CONCLUSIONS.

At the end of many of the preceding sections, and whenever the requirements of the case demanded it, a brief summary of the main facts and of the inferences to be derived from them has been given. It may be useful to give in conclusion a few general remarks on the whole subject.

It will be seen from the numerous facts herein cited, that the so-called monstrous formations (excluding morbid growths the result of disease or injury) present no peculiarities absolutely foreign to the normal organisation of plants. The difference between the natural and monstrous development is one of degree and frequency of occurrence, not of kind.

Deviations from the customary form have been shown to arise from excessive or diminished growth, or from arrested or exalted development.

Even in those instances where, for convenience' sake, the term perverted development has been used, it must be understood as applying only to the particular plant or organ under consideration, as the form a.s.sumed is perfectly in accordance with the ordinary conformation of some other plant or group of plants.

The period at which malformations occur is a matter of some importance; this is, indeed, implied in the term arrest of development; evolution goes on with growth up to a certain point and is then stopped, and thus changes are brought about in the part affected of a different nature from those dependent on non-development or suppression.

Some malformations are congenital, therefore, while others are acquired--in the former instance the disturbance is coeval in origin, and contemporaneous in its growth and development, with those of the affected part; in the latter case the organ may have attained its ordinary degree of perfection, or at least may have advanced some way towards it, before any deviation shows itself. True chorisis or fission, for instance, is usually a congenital affection, arising at a very early period of development, while enation takes place from structures which are all but complete as to their organisation, even though they may not have attained their full dimensions. The date of appearance is also of consequence in determining the true nature of some changes; it does not always follow, for instance, that because one organ occupies the position of another, it is of the same nature as the one whose place it fills. The presence of anthers on petals or on such organs as the corona of _Narcissus_ does not necessarily const.i.tute those parts actual stamens, but rather staminodes. The true stamens are either wanting, or if present, they are in advance of their imitators as regards their development.

=General morphology of the leaf and axis. h.o.m.ology.= Since the time when Goethe's generalisations were adopted by A. P. De Caudolle, special attention has been given to the form and mode of development of the leaf-organ; for as it was well said by Wolff, if once the course of evolution and the structure of the leaf were known, those of the parts of the flower would follow as a matter of course.

It is not necessary, in this place, to pursue the subject of the development and construction of the leaf further than they are ill.u.s.trated by ordinary teratological phenomena.

From this point of view perhaps the most interesting circ.u.mstance is the part that the sheath of the leaf plays.[553] In many cases of so-called metamorphosis, it is the sheath of the leaf that is represented and not the blade. In normal anatomy the sepals, petals, carpels, and even the stamens, as a general rule, correspond to the sheath rather than to the blade of the leaf, as may be seen by the arrangement of the veins. The blade of the leaf seems to be set apart for special respiratory and absorbent offices, while the sheath is in structure, if not in office, more akin to the stem. It would not be easy apart from their position to distinguish between a tubular sheathing leaf and a hollow stem. The development of advent.i.tious growths by chorisis or enation has been frequently alluded to in the foregoing pages, and many ill.u.s.trations have been given of the power that leaves have of branching in more than one plane, owing to the projection of secondary growing-points from the primary organ. These new centres of development are closely connected with the fibro-vascular system of the leaf, so that no sooner does a new growing point originate, than vessels are formed to connect the new growth with the general fibrous cord, see pp. 355, 445. This leads M.

Casimir De Candollo to consider the entire leaf as a composite structure. The morphological unit, says he, is the cellular protrusion or growing point (_saillie_) and its corresponding fibro-vascular bundle.[554]

The ident.i.ty, in a morphological point of view, of the leaves and the lateral parts of the flower is so thoroughly recognised that little need be said on that score, save to repeat that the h.o.m.ology of the floral organs is usually not so much with the entire leaf as with its sheath.

The most singular instances of morphological ident.i.ty are those relating to the s.e.xual organs. We have seen the gradual transition of stamens to pistils, and of pistils to stamens, the development of ovules on the edges of the anther, the co-existence of pollen with ovules on an antheroid body, and, stranger still, the actual development of pollen within the tissues of the ovule itself! From such facts, in addition to what we know of the relative position, internal structure, and mode of development of the organs, it is impossible to avoid coming to the conclusion that, however distinctly these parts may, under ordinary circ.u.mstances, be set apart for the performance of distinct functions, morphologically they are h.o.m.ologous.

These ideas may be carried yet farther--the same sort of evidence, which is adduced in support of the morphological ident.i.ty of leaves with the parts of the flower, may be advanced in confirmation of the opinion, that, morphologically, there is no distinction between axis and leaf.

The leaf, according to this view, is a specialised portion of the axis set apart to do certain work, just as the petals, stamens, &c., are leaves told off for distinct uses. It is unnecessary to refer to the intermediate productions linking the leaf-form to that of the axis, all that is requisite here is to point out the facts that teratology lends in support of these views. These may be summed up by the statement that almost all those attributes which morphologists recognise as peculiar to one or the other organ respectively, may be and are manifested by both.

We have the stem acquiring the characters of the leaf, and the leaf those of the stem. Thus we have seen leaves, leaf-buds, branches, and flower-buds springing from leaves or leaf-organs;[555] see pp. 174, 177, 445, &c. The structure that we are apt to a.s.sociate exclusively with one is found to pertain to the other. The arrangement of the vascular cords in the leaf-organ finds its counterpart in the axis, generally, it is true, modified to suit altered circ.u.mstances or diverse purposes. In some cases the disposition is absolutely indistinguishable in the two organs. It may then be said that the distinctions usually drawn between axis and leaf are not absolute, and that, however necessary such a separation may be for descriptive or physiological purposes, morphologically the two organs are identical. Again, it may be said that leaf and axis are two phases of the same organ,--an organ capable of existing in its undifferentiated state in the form of a thallus among Cryptogams, but which in the higher groups of plants becomes marked out into separate portions, each portion having its own distinct functions to fulfil for the common benefit of the whole organisation.[556]

=Special morphology.=--Under this heading brief reference may be made to some of the organs whose morphological nature has been, and still is, much contested. It is clear that for the due elucidation of these matters, development and the comparative investigation of similar structures in different plants must be studied. Teratological data by themselves can no more be trusted to give a correct solution of any particular question, than the evidence furnished by other departments of botanical science taken separately. With this statement by way of caution, allusion may be made to some of the organs whose morphological construction is ill.u.s.trated by the facts recorded in the present volume.

=Calyx-tube.=--In descriptive botany it is the common practice to speak of a calyx-tube, by which is meant a tubular or sheathing portion at the base of the flower, below the sepals or calyx-lobes, and distinct or inseparable from the ovary. The question morphology has to solve is whether this tubular structure is to be considered as a portion of the axis, or whether it is to be regarded as composed of the confluent bases of the sepals.

Mr. Bentham, who has recently reviewed the evidence as to the nature of the calyx-tube in his paper on _Myrtaceae_,[557] still holds to the notion that the "calyx-tube" or "hypanthium" is formed from the concretion of the basal portions of the sepals. He founds his conclusions upon such facts as the following: the circ.u.mstance that the point of origin of the leaf is not always the same as the point of disarticulation or separation from the axis, inasmuch as the basal portion of the leaf is often adherent to the stem for some distance, though still recognisable as foliar not axial in its nature. In the same manner, the corolla and androecium may be concrete at the base, so that the stamens are for convenience' sake described as inserted into the tube of the corolla, though it is generally admitted that both stamens and petals are really hypogynous, and it is not usual to consider the corolla-tube up to the divergence of the stamens as part of the receptacle. A similar remark applies to the carpels and placentas.

Mr. Bentham further considers that the gradual disconnection of the various whorls, that may be traced in many plants, is a further proof of concretion, rather than of expansion of the axis, but this argument may fairly be met by the consideration that the several whorls emerge at different heights.[558]

Organs originally free and distinct become ultimately combined at the base by the gradual protrusion from the receptacle of a ring or tube under them, as in the stamens of _Leguminosae_; yet, says Mr. Bentham, no one would propose to describe the staminal tube of monadelphous _Leguminosae_ as part of the receptacle and not of the stamens. Perhaps not, for descriptive purposes, but morphologically it would not be easy to separate such a tube from the receptacle. The princ.i.p.al kinds of malformation which have a bearing on this subject are mentioned at pp.

77-81 and 247, from which it may be seen that the evidence furnished by teratology is conflicting. It would seem, indeed, that while in some families of plants there may be a real calyx-tube, in others the tubular portion is a sheath-like prolongation of the axis. In _Primula_ or _Pedicularis_, where the venation is clearly laminar, the tubular portion is distinctly calycine. In other cases the so-called calyx-tube seems as certainly to be an expansion of the receptacle, as in _Rosaceae_, _Myrtaceae_, _Melastomaceae_, _Pa.s.siflora_,[559] &c.

Where the petals and stamens are described as being inserted into the throat of the calyx, or are perigynous, it may be a.s.sumed as a general rule, subject to but few exceptions, that the so-called calyx-tube is really a portion of the receptacle.[560] After all, this is very much a question of words, and for the following reasons,--very often the base of the calyx does evidently form a tube, and no one can say where the calyx ends and the receptacle begins. Again, many leaves are known to originate in the form of a ring-like protrusion from the axis, and from this primary ring originate secondary developments. Thus the a.s.serted difference between a leaf, with such a history of development, and an axial structure becomes obliterated. From this point of view, peltate leaves like those of _Tropaeolum_ or _Nelumbium_ become very significant.

In both the leaf-stalk is cylindrical and traversed, as in the case of all cylindrical leaf-stalks, by a circle of fibro-vascular cords, as in a branch, and which radiate in all directions in the blade of the leaf.

Now, if (as often happens to a slight extent) the central portion of the leaf were much depressed, owing to the disproportionate growth of the peripheral, as contrasted with the central portions, we should have a funnel-like or tubular formation, precisely similar to many of the so-called calyx-tubes. And, if we further suppose new growths to originate from the sides of this funnel or tube, by chorisis or enation, we should have the h.o.m.ologue of a tubular calyx, to the inner surface of which are attached petals, stamens, &c. From the consideration of circ.u.mstances such as these just detailed, together with that of the arrangement of the vascular cords, M. Casimir De Candolle arrives at the conclusion that the calyx-tube is a ring-like projection from an axis whose further direct development is arrested. The secondary projections or growing-points correspond to the several fibro-vascular cords of the primary ring, and are ultimately developed into sepals, petals, stamens and ovaries (see pp. 394, 509).

=Androecium.=--The main points of morphological interest relating to the androecium, referred to in this volume, are those concerning the structure of the anther (see p. 292), the compound nature of the stamens in some orders (see pp. 294, 345), and the nature of the androecium in orchids (see p. 380).

=Inferior ovary.=--Is the pistil always foliar in its morphological nature, or is it, in some cases, as Schleiden taught, formed from the axis alone? To a great extent the reply to this question is dependent on the conclusions that may be arrived at as to the true nature of the calyx-tube. Considered from a teratological point of view, there is no reason for considering the inferior ovary to be purely axial. On the contrary, the evidence derived from this source supports the ordinary opinion that the carpels are inv.a.g.i.n.ated within the expanded top of the flower-stalk and more or less adherent to it. Some of the gourds afford good ill.u.s.trations of this, the upper part of the carpels in these fruits projecting beyond the axial portion. But this matter loses much of its importance if the morphological ident.i.ty of axis and leaf-organ be conceded. The carpels in inferior ovaries seldom or never correspond to the lamina of the leaf, and between the v.a.g.i.n.al portion of the carpellary leaf, and the axis who shall draw the distinction?

=Placentation.=--Some botanists have considered the placentas to be portions of the carpel, and have compared the production of ovules on them to the formation of buds on the leaf of _Bryophyllum_. Others have been led to see in each placenta, even when it is, to all outward appearance, a portion of the carpellary leaf, a direct prolongation from the axis, adherent to the leaf. Teratology shows that ovules may be formed indifferently on leaf-organs or on stem-organs. Sutural, parietal, axile, free-central placentation, and, if there be more forms, all may be met with even in the same ovary (see pp. 96, 508). Now, if there were such special tendencies in the axis, as contrasted with the leaf, to produce ovules, it is hardly likely that such anomalous arrangements as those just mentioned would be as frequent as they are.

But as leaves produce other leaves, from their edges or their surfaces, and as they form buds in the same situations, just as axial organs do,[561] there is surely little ground for considering the placentas, or ovuliferous portions of the plant, to be of necessity axial. Here again, much of the difficulty vanishes if the morphological ident.i.ty of the leaf-form and of the stem-form be admitted.

=Structure of the ovule.=--The nature of the ovule and of its coverings has been a fertile source of controversy. The teratological data bearing on this subject have been given at pp. 262-272. These data strongly support the notion of the foliar nature of the coatings, and of the axial nature of the nucleus, taking leaf and axis either in the ordinary sense, or as modifications one of the other. It has been shown that the ovular coats may themselves become carpels, and that ovules may be developed upon ovules, p. 268. Whether the intra-carpellary siliques of _Cheiranthus_, not uncommonly met with (p. 182), are instances of ovular trans.m.u.tation may be open to doubt.

The axial nature of the nucleus has been inferred from its position, mode of growth, and from its occasionally lengthening into a leafy or even a floriferous shoot. Probably it may occasionally be invested by sheathing coats, more a.n.a.logous to tubular processes from the receptacle, than to foliar organs, as is the case in _Welwitschia_. The discussion of this matter, however, pertains rather to normal morphology than to teratology.

=Morphology of conifers.=--The nature of the pseudo-leaves of _Sciadopitys_, and probably of other Conifers, is ill.u.s.trated by teratology, as also is the true const.i.tution of the scale of the cone (see pp. 192, 245, 352), though it must be admitted that little or no light is thrown on that much-contested point--the true nature of the ovule of Gymnosperms.