Read Ebook: Vegetable Teratology An Account of the Principal Deviations from the Usual Construction of Plants by Masters Maxwell T Maxwell Tylden

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ENATION 443

Excrescences from axile organs, warts, 444--Enation from leaves, 445--From sepals, 448--From petals, 448--Catacorolla, 451--Enation from stamens, 453--From carpels, 453.

ABORTION 455

Abortion of axile organs, 455--Of receptacle, 457--Of leaves, 458--Of perianth, calyx, and corolla, 460--Of stamens, 463--Of pollen, 463--Of pistil, fruit, &c., 464--Of ovules, 466--Depauperated ferns, 466--General remarks, 467.

DEGENERATION 470

Formation of scales, 470--Of hairs, 472--Of glands, 473--Of tendrils, 473.

GENERAL CONCLUSIONS.

General morphology of the leaf and axis, 476.--Homology, 476--Special morphology, 479--Calyx-tube, 480--Androecium, 482--Inferior ovary, 482--Placentation, 483--Structure of the ovule, 484--Leaves of Conifers, 484--Relative position of organs, 484--Law of alternation, 485--Co-relation, 486--Compensation, 488--Teratology and classification, 488.

Double flowers, varieties of, 491--Causes of production, 491--Relation to variegated foliage, 497--List of plants producing, 499.

NOTE 508

INDEX TO SUBJECTS 511

INDEX OF NAMES OF PLANTS 517

FIG. PAGE

INTRODUCTION.

The older botanists, it is true, often made them the basis of satirical allusions to the political or religious questions of the day, especially about the time of the Reformation, and the artists drew largely upon their polemical sympathies in their representations of these anomalies. Linnaeus treated of them to some extent in his 'Philosophia,' but it is mainly to Angustin Pyramus De Candolle that the credit is due of calling attention to the importance of vegetable teratology. This great botanist, not only indirectly, but from his personal research into the nature of monstrosities, did more than any of his predecessors to rescue them from the utter disregard, or at best the contemptuous indifference, of the majority of botanists. De Candolle gave a special impetus to morphology in general by giving in his adhesion to the morphological hypotheses of Goethe. These were no mere figments of the poet's imagination, as they were to a large extent based on the actual investigation of normal and abnormal organisation by Goethe both alone, and also in conjunction with Batsch and Jaeger.

De Candolle's example was contagious. Scarcely a botanist of any eminence since his time but has contributed his quota to the records of vegetable teratology, in proof of which the names of Humboldt, Robert Brown, the De Jussieus, the Saint Hilaires, of Moquin-Tandon, of Lindley, and many others, not to mention botanists still living, may be cited. To students and amateurs the subject seems always to have presented special attractions, probably from the singularity of the appearances presented, and from the fact that in many cases the examination of individual instances of malformation can be carried on, to a large extent, without the lengthened or continuous investigation and critical comparative study required by other departments of botanical science. Be this as it may, teratology owes a very large number of its records to this class of observers.

While the number of scattered papers on vegetable teratology in various European languages is so great as to preclude the possibility of collating them all, there is no general treatise on the subject in the English language, with the exception of Hopkirk's 'Flora Anomala,' a book now rarely met with, and withal very imperfect; and this notwithstanding that Robert Brown early lent his sanction to the doctrines of Goethe, and himself illustrated them by teratological observations. In France, besides important papers of Turpin, Geoffroy de Saint Hilaire, Brongniart, Kirschleger and others, to which frequent allusion is made in the following pages, there is the classic work of Moquin-Tandon, which was translated into German by Schauer. Germany has also given us the monographs of Batsch, Jaeger, Roeper, Engelmann, Schimper, Braun, Fleischer, Wigand, and many others. Switzerland has furnished the treatises of the De Candolles, and of Cramer; Belgium, those of Morren, &c., all of which, as well as many others that might be mentioned, are, with the exception of Moquin-Tandon's 'El?ments,' to be considered as referring to limited portions only and not to the whole subject.

The drawings, where not otherwise stated, have been executed either from the author's own rough sketches, or from the actual specimens, by Mr. E. M. Williams. A large number of woodcuts have also been kindly placed at the disposal of the author by the proprietors of the 'Gardeners' Chronicle.'

As it is impossible to frame any but a purely arbitrary definition of teratology or to trace the limits between variation and malformation, it may suffice to say that vegetable teratology comprises the history of the irregularities of growth and development in plants, and of the causes producing them. These irregularities differ from variations mainly in their wider deviation from the customary structure, in their more frequent and more obvious dependence on external causes rather than on inherent tendency, in their more sudden appearance, and lastly in their smaller liability to be transmitted by inheritance.

What may be termed normal morphology includes the study of the form, arrangement, size and other characteristic attributes of the several parts of plants, their internal structure, and the precise relation one form bears to another. In order the more thoroughly to investigate these matters it is necessary to consider the mode of growth, and specially the plan of evolution or development of each organ. This is the more needful owing to the common origin of things ultimately very different one from the other, and to the presence of organs which, in the adult state, are identical or nearly so in aspect, but which nevertheless are very unlike in the early stages of their existence. Following Goethe, these changes in the course of development are sometimes called metamorphoses. In this way Agardh admits three kinds of metamorphosis, which he characterises as: 1st. Successive metamorphoses, or those changes in the course of evolution which each individual organ undergoes in its passage from the embryonic to the adult condition, or from the simple and incomplete to the complex and perfect. 2. Ascending metamorphoses, including those changes of form manifested in the same adult organism by the several parts of which it consists--those parts being typically identical or homologous, such as the parts of the flower, or, in animals, the vertebrae, &c. 3. Collateral metamorphoses, comprising those permutations of form and function manifested in homologous organs in the different groups of organisms, classes, orders, genera, species, &c.

Thus, in the first instance, we have a comparative examination of the form of each or any separate part of the same individual at different epochs in its life-history; in the second we have a similar comparison instituted between the several parts of the same organism which originally were identical in appearance, but which have in course of evolution altered in character. In the third form we have the comparative view not of one organ at different times, nor of the several parts of one organism, but of the constituent elements pertaining to those aggregates of individuals to which naturalists apply the terms classes, orders, &c.

In successive metamorphosis we have a measure of the amount of change and of the perfection of structure to which each separate organ attains.

In ascending metamorphosis we have a gauge of the extent of alteration that may take place in the several homologous organs under existing circumstances.

In collateral metamorphosis, in the same way, we have an illustration of the degree of change possible in aggregates of organisms under existing circumstances.

Now it is clear that from an investigation of all three classes just mentioned, we shall be able to gain an idea of those points which are common to all parts, to all individuals or to all aggregates, and those that are peculiar to some of them, and, by eliminating the one from the other, we shall arrive at conclusions which will be more or less generally accurate or applicable, according to the ability of the student and the extent to which the comparative analysis is earned. It is thus that morphologists have been enabled to frame types or standards of reference, and systematists to collocate the organisms they deal with into groups. These standards and groups are more or less artificial in proportion to the amount of knowledge possessed by their framers, and the use they make of it.

From this point of view teratological metamorphosis of all three kinds demands as much attention as that which is called normal. We can have no thorough knowledge of an organ, of an individual which is an aggregate of organs, or of an aggregate of individuals of whatever degree, unless we know approximately, at least, what are the limits of each. It is not possible to trace these limits accurately in the case of natural science, but the larger our knowledge and the wider our generalisations, the closer will be our approach to the truth.

The most satisfactory classification of malformations would be one founded upon the nature of the causes inducing the several changes. Thus, in all organised beings, there is a process of growth, mere increase in bulk as it were, and a process of evolution or metamorphosis, in accordance with which certain parts assume a different form from the rest, in order the better to fit them for the performance of different offices. Should growth and development be uniform and regular, that is in accordance with what is habitual in any particular species, there is no monstrosity, but if either growth or development be in any way irregular, malformation results. Hence, theoretically, the best way of grouping cases of malformation would be according as they are the consequences of:--1st. Arrest of Growth; 2ndly. Excessive Growth; 3rdly. Arrest of Development; 4thly, of Excessive or Irregular Development.

In practice, however, there are so many objections to this plan that it has not been found practicable to carry it out. The inability arises to a great extent from our ignorance of what should be attributed to arrest of growth, what to excess of development, and so on. Moreover, a student with a malformed plant before him must necessarily ascertain in what way it is malformed before he can understand how it became so, and for this purpose any scheme that will enable him readily to detect the kind of monstrosity he is examining, even though it be confessedly artificial and imperfect will be better than a more philosophical arrangement which circumstances prevent him from employing.

The plan followed in this volume is a slight modification of that adopted by Moquin-Tandon, and with several additions. In it the aim is to place before the student certain salient and easily recognisable points by reference to which the desired information can readily be found. Under each subdivision will be found general explanatory remarks, illustrative details, and usually a summary of the more important facts and the inferences to be derived from them. Bibliographical references and lists of the plants most frequently affected with particular malformations are also given. In reference to both these points it must be remembered that absolute completeness is not aimed at; had such fullness of detail been possible of attainment it would have necessitated for its publication a much larger volume than the present. It is hoped that both the lists of books and of plants are sufficiently full for all general purposes.

Teratological alterations are rarely isolated phenomena, far more generally they are associated with other and often compensatory changes. Hence it is often necessary, in studying any given malformation, to refer to two or more subdivisions, and in this way a certain amount of repetition becomes unavoidable. The details of the several cases of malformation given in these pages are generally arranged according to their apparent degree of importance. Thus, in a case of prolification associated with multiplication of the petals, the former change is a greater deviation from the customary form than the latter, hence reference should be made, in the first instance, to the sections treating on prolification, and afterwards to those on multiplication. To facilitate such research, numerous cross references are supplied.

It has already been said that an arrangement like that here considered as typical is natural to some flowers in their adult state, and to a vast number in their immature condition. It would be no extravagant hypothesis to surmise that this was the primitive structure of the flower in the higher plants. Variations from it may have arisen in course of time, owing to the action of an inherent tendency to vary, or from external circumstances and varied requirements which may have induced corresponding adaptations, and which may have been transmitted in accordance with the principle of hereditary transmission. This hypothesis necessarily implies a prior simplicity of organisation, of which, indeed, there is sufficient proof; many cases of malformation can thus be considered as so many reversions to the ancestral form.

Thus, teratology often serves as an aid in the study of morphology in general, and also in that of special groups of plants, and hence may even be of assistance in the determination of affinities. In any case the data supplied by teratology require to be used with caution and in conjunction with those derived from the study of development and from analogy. It is even possible that some malformations, especially when they acquire a permanent nature and become capable of reproducing themselves by seed, may be the starting-point of new species, as they assuredly are of new races, and between a race and a species he would be a bold man who would undertake to draw a hard and fast line.

Discredit has been cast on teratology because it has been incautiously used. At one time it was made to prove almost everything; what wonder that by some, now-a-days, it is held to prove nothing. True the evidence it affords is sometimes negative, often conflicting, but it is so rather from imperfect interpretation than from any intrinsic worthlessness. If misused the fault lies with the disciple, not with Nature.

Teratology as a guide to the solution of morphological problems has been especially disparaged in contrast with organogeny, but unfairly so. There is no reason to exalt or to disparage either at the expense of the other. Both should receive the attention they demand. The study of development shows the primitive condition and gradual evolution of parts in any given individual or species; it carries us back some stages further in the history of particular organisms, but so also does teratology. Many cases of arrest of development show the mode of growth and evolution more distinctly, and with much greater ease to the observer, than does the investigation of the evolution of organs under natural circumstances. Organogeny by no means necessarily, or always, gives us an insight into the principles regulating the construction of flowers in general. It gives us no archetype except in those comparatively rare cases where primordial symmetry and regularity exist. When an explanation of the irregularity of development in these early stages of the plant's history is required, recourse must be had to the inferences and deductions drawn from teratological investigations and from the comparative study of allied forms precisely as in the case of adult flowers.

The study of development is of the highest importance in the examination of plants as individuals, but in regard to comparative anatomy and morphology, and specially in its relation to the study of vegetable homology it has no superiority over teratology. Those who hold the contrary opinion do so, apparently, because they overlook the fact that there is no distinction, save of degree, to be drawn between the laws regulating normal organisation, and those by which so-called abnormal formations are regulated.

It is sometimes said, and not wholly without truth, that teratology, as it stands at present, is little more than a record of facts, but in proportion as the laws that regulate normal growth are better understood, so will the knowledge of those that govern the so-called monstrous formations increase. Sufficient has been already said to prove that there is no intrinsic difference between the laws of growth in the two cases. As our knowledge increases we shall be enabled to ascertain approximately of what extent of variation a given form is capable, under given conditions, and to refer all formations now considered anomalous to a few well-defined forms. Already teratology has done much towards showing the erroneous nature of many morphological statements that still pass current in our text-books, though their fallacy has been demonstrated again and again. Thus organs are said to be fused which were never separate, disjunctions and separations are assigned to parts that were never joined, adhesions and cohesions are spoken of in cases where, from the nature of things, neither adhesion nor cohesion could have existed. Some organs are said to be atrophied which were never larger and more fully developed than they now are, and so on. So long as these expressions are used in a merely conventional sense and for purposes of artificial classification or convenience, well and good, but let us not delude ourselves that we are thus contributing to the philosophical study either of the conformation of plants or of the affinities existing between them. What hope is there that we shall ever gain clear conceptions as to the former, as long as we tie ourselves down to formulas which are the expressions of facts as they appear to be, rather than as they really are? What chance is there of our attaining to comprehensive and accurate views of the genealogy and affinities of plants as long as we are restricted by false notions as to the conformation and mutual relation of their parts?

That teratology may serve the purposes of systematic botany to a greater extent than might at first be supposed becomes obvious from a consideration of such facts as are mentioned under the head of Peloria, while the presence of rudimentary organs, or the occasional appearance of additional parts, or other changes, may, and often do, afford a clue to the relationship existing between plants--a relationship that might otherwise be unsuspected. So, too, some of the alterations met with appear susceptible of no other explanations than that they are reversions to some pre-existing form, or, at any rate, that they are manifestations of a phase of the plant affected different from that which is habitual, and due, as it were, to a sort of allotropism.

The mutations and perversions of form, associated as they commonly are with corresponding changes of function, show the connection between teratology and physiology--a connection which is seen to be the more intimate when viewed in the light afforded by the writings and experiments of Gaertner, Sprengel, and St. Hilaire, and, in our own times, especially by the writings and experiments of Mr. Darwin, whose works on the 'Origin of Species,' and particularly on the 'Variation of Animals and Plants under Domestication' comprise so large a collection of facts for the use of students in most departments of biology. It will suffice to allude, in support of these statements, to the writings of Mr. Darwin on such subjects as rudimentary organs, the use or disuse of certain parts according to circumstances, the frequently observed tendency of some flowers to become structurally unisexual, the liability of other flowers perfectly organised to become functionally imperfect, at least so far as any reciprocal action of the organs of the same flower is concerned, reversions, classification, general morphology, and other subjects handled at once with such comprehensive breadth and minute accuracy of detail by our great physiologist.

In the following pages alterations of function, unless attended by corresponding alterations of form, are either only incidentally alluded to, or are wholly passed over; such, for instance, as alterations in the period of flowering, in the duration of the several organs, and so forth. Pathological changes, lesions caused by insect puncture or other causes, also find no place in this book, unless the changes are of such a character as to admit of definite comparison with normal conformation. Usually such changes are entirely heteromorphous, and, as it were, foreign to the natural organisation.

The practical applications of teratology deserve the attention of those cultivators who are concerned in the embellishment of our gardens and the supply of our tables. The florist lays down a certain arbitrary standard of perfection, and attempts to make flowers conform to that model. Whether it be in good taste or not to value all flowers, in proportion as they accord with an artificial and comparatively inelastic standard of this kind, we need not stop to enquire; suffice it to say, that taking the matter in its broadest sense, the aim of the florist is to produce large, symmetrical flowers, brightly and purely coloured, or if parti-coloured, the colours must be distinct, harmonious, or contrasted. When all this is done, the flower, in most instances, becomes 'monstrous' of the eyes in the botanist, though all the more interesting to the student of morphology on that account. In like manner the double flowers, the "breaks," the "sports" which the florist cultivates so anxiously, are all of them greater or less deviations from the ordinary form, while the broccolies, the cabbages, and many other products of our kitchen gardens and fields owe the estimation in which they are held entirely to those peculiarities which, by an unhappy application of words, are called monstrous by botanists. Grafting, layering, the "striking" of cuttings, the formation of adventitious roots and buds, processes on which the cultivator so greatly relies for the propagation and extension of his plants, are also matters with which teratology concerns itself. Again the difficulty experienced occasionally in getting vines, strawberries, &c., to set properly, may sometimes be accounted for by that inherent tendency which some plants possess of exchanging an hermaphrodite for a unisexual condition.

For reasons then of direct practical utility, no less than on purely scientific grounds, it is desirable to study these irregularities of growth, their nature, limits, and inducing causes; and to this end it is hoped the present work may, in some degree, contribute.

FOOTNOTES:

An excellent summary of the history of Vegetable Teratology is given in Kirschleger's 'Essai historique de la T?ratologie V?g?tale,' Strasburg, 1845.

In some instances diagrams and formulae are given in explanation of the conformation of monstrous flowers; in general these require no further explanation than is given in the text, unless it be to state that the horizontal line--is intended to indicate the cohesion of the parts over which it is placed, while the vertical line | signifies the adhesion of the organs by whose side it is placed. The formula

shows that the sepals are distinct, the petals coherent, and the stamens adherent to the petals.

Wolff was the first to call attention to the great importance of the study of development. He was followed by Turpin, Mirbel, Schleiden, Payer, and others, and its value is now fully recognised by botanists.

Agardh, "Theoria Syst. Plant.," p. xxiii.

In the memoirs of Hopkirk, Kirschleger, Cramer, Hallier, and others, malformations are arranged primarily according to the organs affected, an arrangement which has only convenience to justify it. It is hoped that the index and the headings to the paragraphs in the present volume will suit the convenience of the reader as well as if the more artificial plan just alluded to had been adopted.

Cryptogamous plants are only incidentally alluded to in these pages, owing to their wide difference in structure from flowering plants. Attention may, also, here be called to a paper of M. de Seynes in a recent number of the Bulletin of the Botanical Society of France, vol. xiv, p. 290, tab. 5 et 6, in which numerous cases of malformation among agarics are recorded. See also same publication, vol. iv, p. 744; vol. v, p. 211; vol. vi, p. 496.

On this subject see a paper of M. Naudin in the 'Comptes Rendus,' 1867, t. 64, pp. 929-933.

It is probable that many terms and expressions calculated to mislead in the way above mentioned are made use of in the following pages. The inconsistency manifested by their use may be excused on the ground of ignorance of the true structure, and by the circumstance that in many cases facts alone are recorded without an explanation of them being offered. Moreover, it is desirable to act in conformity with the usual practice of botanical writers, and not to change established terminology, even if suspected to convey false ideas, until the true condition of affairs be thoroughly well ascertained by organogenetic research or other means.

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