Sit?? B. 1. Bill ©bran; SJnrtb (Earoltna £>tate (Tiillrg* L55 S00479466 This book is due on the date indicated belo- and is subject to a fine of FIVE CENTS day thereafter. ~"^et*a*«6^ ^^^^■10 JUN*i W/4 DEC 2 4 1976 rJUI fe ™ # ^^^^ • INTRODUCTION BOTANY. London : Printed by A. & R. Spottiswoode, New-Street- Square. AN INTRODUCTION B O T A N BY JOHN LINDLEY, F.R.S. L.S. G.S. MEMBER OF THE IMPERIAL ACADEMY NATURES CURIOSORUM J OF THE BOTANICAL SOCIETY OF RATISBON J OF THE PHYSIOGRAPHICAL SOCIETY OF LUND; OF THE HORTICULTURAL SOCIETY OF BERLIN J HONORARY MEMBER OF THE LYCEUM OF NATURAL HISTORY OF NEW YORK; AS- SISTANT SECRETARY OF THE HORTICULTURAL SOCIETY OF LONDON, ETC. ETC. J AND PROFESSOR OF BOTANY IN THE UNIVERSITY OF LONDON. aiaUitf) &tjc eropper^lPIates ant) numeroiw 2ia3o- PREFACE. A. wo hundred and ninety years have now elapsed since one of the earliest introductions to Botany upon record was published, in four pages folio, by Leonhart Fuchs, a learned physician of Tubingen. At that period Botany was nothing more than the art of distinguishing one plant from another, and of remembering the medical qualities, sometimes real, but more frequently imaginary, which experience, or error, or superstition, had ascribed to them. Little was known of Vegetable Physiology, nothing of Vege- table Anatomy, and even the art of arranging species systematically had still to be discovered; while scarcely a trace existed of those modern views which have raised the science from the mere business of the herb-gatherer to a station among the most intel- lectual branches of natural philosophy. It now comprehends a knowledge not only of the names and uses of plants, but of their external and internal organisation, and of their anatomy and phy- siological phenomena ; it embraces a consideration of the plan upon which those multitudes of vegetable forms that clothe the earth have been created, of the skilful combinations out of which so many various organs have emanated, of the laws that regulate the dispersion and location of species, and of the influ- ence that climate exercises upon their developement ; A 3 PREFACE. and, lastly, from botany as now understood, in its most extensive signification, is inseparable the know- ledge of the various ways in which the laws of vege- table life are applicable to the augmentation of the luxuries and comforts, or to the diminution of the wants and miseries of mankind. It is by no means, as some suppose, a science for the idle philosopher in his closet ; neither is it merely an amusing accom- plishment, as others appear to think ; on the con- trary, its field is in the midst of meadows, and gardens, and forests, on the sides of mountains, and in the depths of mines, — wherever vegetation still flourishes, or wherever it attests by its remains the existence of a former world. It is the science that converts the useless or noxious weed into the nutri- tious vegetable ; which changes a bare volcanic rock, like Ascension, into a green and fertile island ; and which enables the man of science, by the power it gives him of judging how far the productions of one climate are susceptible of cultivation in another, to guide the colonist in his enterprises, and to save him from those errors and losses into which all such per- sons unacquainted with Botany are liable to fall. This science, finally, it is which teaches the physician how to discover in every region the medicines that are best adapted for the maladies that prevail in it ; and which, by furnishing him with a certain clue to the knowledge of the tribes in which particular proper- ties are or are not to be found, renders him as much at ease, alone and seemingly without resources, in a land of unknown herbs, as if he were in the midst of a magazine of drugs in some civilised country. The principles of such a science must necessarily be extremely complicated, and in certain branches, which have only for a short time occupied the atten- PREFACE. Vll tion of observers, or which depend upon obscure and ill-understood evidence, are by no means so clearly defined as could be wished. To explain those prin- ciples, to adduce the evidence by which their truth is supposed to be proved, or the reasoning upon which they are based in cases where direct proof is unattainable ; to show the causes of errors that are now exploded, and the insufficiency of the arguments by which doubtful theories are still defended, — in fine, to draw a distinct line between what is certain and what is doubtful, — are some of the objects of this publication, which is intended for the use of those who, without being willing to occupy themselves with a detailed examination of the vast mass of evi- dence upon which the modern science of botany is founded, are, nevertheless, anxious to acquire a dis- tinct idea of the nature of that evidence. Another and not less important purpose has been to demon- strate, by a series of well-connected proofs, that in no department of natural history are the simplicity and harmony that pervade the universe more strikingly manifest than in the vegetable kingdom, where the most varied forms are produced by the combination of a very small number of distinct organs, and the most important phenomena are distinctly explained by a few simple laws of life and structure. In the execution of these objects, I have followed very nearly the method recommended by the cele- brated Professor De Candolle, than whom no man is entitled to more deference, whether you consider the soundness of his judgment in all that relates to order and arrangement, or the great experience which a long and most successful career of public instruction has necessarily given him. I have begun with what is called Organography a 4 VIU PREFACE. (Book I.) ; or an explanation of the exact structure of plants; a branch of the subject which comprehends all that relates either to the various forms of tissue of which vegetables are constructed, or to the external appearance their elementary organs assume in a state of combination. It is exceedingly desirable that these topics should be well understood, because they form the basis of all other parts of the science. In physiology, every function is executed through the agency of the organs : systematic arrangements de- pendjupon characters arising out of their consider- ation ; and descriptive Botany can have no logical precision without the principles of Organography being first exactly settled. Great difference of opi- nion exists among the most distinguished botanists, upon some points connected with this subject, so that it has been found expedient to enter occasionally into much detail, for the purpose of satisfying the student of the accuracy of the facts and reasonings upon which he is expected to rely. To this succeedsVEGETABLE.PH ysiology (BooklL); or the history of the vital phenonema that have been observed both in plants in general, and in particular species, and also in each of their organs taken sepa- rately. It is that part of the science which has the most direct bearing upon practical objects, and with which the enquirer who would occupy himself more with the utile than the dulce is most likely to be inter- ested. Its laws, however, are either unintelligible, or susceptible of no exact appreciation, without a pre- vious acquaintance with the more important details of Organography. Much of the subject is at present involved in mystery, and the accuracy of many of the conclusions of physiologists is inferred rather than demonstrated ; so that it has been found essential l'KEFACE. ix that the grounds of the more popularly received opinions, whether admitted as true or rejected as erroneous, should be given at length. No particular chapter is assigned to the practical application of physiological principles, because this has been con- stantly taken as illustrative of the separate functions of individual parts. Next follows Taxonomy (Book III.); or some account of the Principles of Classification ; — a very important subject, comprehending not only an account of the various methods of arrangement employed by botanists in their systematic works, but an explan- ation of the principles by which the limits of genera and species are determined. It also explains the mode of obtaining a correct view of vegetation, of con- ducting the examinations of unknown plants with precision, of avoiding errors in consequence of acci- dental aberrations from the ordinary structure, and of forming a just estimate of the mutual relation that one part of the vegetable kingdom bears to another. After this I have taken Glossology (Book IV.) ; or, as it was formerly called, Terminology ; restrict- ing it absolutely to the definition of the adjective terms, which are either used exclusively in Botany, or which are used in that science in some particular and unusual sense. The key to this book, and also to the substantive terms explained in Organography, will be found in a copious index at the end of the volume. These four topics exhaust the science considered only with reference to first principles ; there are, how- ever, a few others which it has been thought advis- able to append, on account of their practical value. These are, firstly, Phytography (Book V.) ; or an exposition of the rules to be observed in describing X PREFACE. and naming plants. As the great object of descrip- tions in natural history, is to enable every person to recognise a known species, after its station has been discovered by classification, and also to put those who have not had the opportunity of examining a plant themselves into possession of all the facts necessary to acquire a just notion of its structure and affinities ; it is indispensable that the principles of making descriptions should be clearly understood, both to prevent their being too general to answer the intended purpose, or more prolix than is really requisite. It is the want of a knowledge of these rules that renders the short descriptions of the classical writers of antiquity, and the longer ones of many a modern traveller, equally vague and unintelligible. In this place are inserted a few notes upon the form, ation of an herbarium. After this, has been introduced (Book VI.) a summary of the little which is known of the laws that regulate the distribution of plants upon the surface of the earth ; a question which, however indefinite and unsatisfactory our information may at present be, has begun to assume such an appearance as to justify the expectation, that future discoveries will explain the causes of the characters of vegetation being determined, as they surely are, by climate. Finally, the work is concluded by an exposition of what is called Morphology ; a subject which is in the vegetable what Comparative Anatomy is in the animal kingdom, and which is by far the most im- portant branch of study after Elementary Anatomy and Vegetable Physiology. Organography itself is in all respects an exposition of thg doctrines of Mor- phology ; but the novelty of the subject, and a per- suasion that it would be better understood if treated PREFACE. xi separately, has induced me to make it the subject of particular consideration. Unknown before the time of Linnaeus, and first placed in its true light by the venerable poet Gothe, it lay neglected for nearly thirty years, until, having been revived by Du Petit Thouars, De Candolle, Brown, and others, it has come to be considered the basis of all scientific knowledge of vegetable structure. It has been my wish to bring every subject that I have introduced down, as nearly as possible, to the state in which it is found at the present day; but, alas ! tasks that are infinite can never be accomplished by finite means. While the MSS. has been going- through the press, my table has been covered with works illustrating the fundamental principles of the science, of scarcely any of which has it been possible to make the slightest use. This I regret the more, as some of them are of high interest*, especially with regard to Vegetable Anatomy. In the statements I have made, I have uniformly endeavoured to render due credit to all persons for the discoveries by which they may severally have contributed to the advancement of the science ; and * The more important of these works are, — Agardh, C. A., Lehrbuch der Botanik. Copenhagen, 1831. Kunth, K. S., Handbuch der Botanik. Berlin, 1831. Meyer, F. J. F., Phytotomie. Berlin, 1830. De Candolle, A. P., Physiologie Vegetale, &c. Paris, 1832. Martius, Palmae. The part containing the Anatomy. Arnott, G. W. The article Botany in the Edinburgh Encyclo- paedia. Bischoff, G. W., Handbuch der Botanischen Terminologie und Systemkunde. Nuremberg, 1830. Annales des Sciences Naturelles, par MM. Audouin, Ad. Brongniart, et Dumas. Several Numbers. Mirbel, C.B.. Memoire sur le Marchantia. Paris, 1832. I'll E FACE. if I have on any occasion either omitted to do so, or assumed to myself observations which belong to others, it has been unknowingly or inadvertently. It is, however, impracticable, and if practicable it would not be worth while, to remember upon all occasions from what particular sources information may have been derived. Discoveries, when once communi- cated to the world, become public property : they are thrown into the common stock for mutual benefit; and it is only in the case of debatable opinions, or of any recent and unconfirmed observations, that it really interests the world that authorities should be quoted at all. In the language of a highly valued friend, when writing upon another subject : — " The advanced state of a science is but the accumulation of the discoveries and inventions of many : to refer each of these to its author is the business of the history of science, but does not belong to a work which professes merely to give an account of the science as it is j all that is generally acknowledged must pass current from author to author." * London, Sept. 10. 1832. * Brett's Principles of Astronomy, p. v. CONTENTS. BOOK I. Page Organography; or, of the Structure of Plants I Chap. I. Of the Elementary Organs - - - - ib. 1. Of Cellular Tissue - - 3 2. Of Woody Fibre - - - - 12 5. Of Vascular Tissue - - - - 17 4. Of Spurious Elementary Organs : 1. Intercellular Passages - - -26 2. Receptacles of Secretion - - - 27 3. Air Cells - - ib. 4. Raphides - - 29 Chap. II. Of the Compound Organs in Flowering Plants - - 31 1. Of the Cuticle and its Appendages: 1. Cuticle - - ib. 2. Stomata - ------33 3. Hairs - - - - - - 38 4. Scales - - - - 41 5. Glands - - - - 42 6. Prickles - - 44 2. Of the Stem or Ascending Axis : 1. Of its Parts - ----- 45 2. Of its External Modifications - - 54 3. Of its Internal Modifications - - - 58 1. Of the Exogenous Structure - - 59 2. Of the Endogenous Structure - - 72 3. Of the Root or Descending Axis - - 76 4. Of the Appendages of the Axis - 78 1. Leaf - - 79 2. Stipulae - 98 3. Bracteae - - - - - 100 XIV CONTENTS. Page 4. Flower - - - - - - 105 5. Inflorescence - - - - - 106 6. Calyx - 112 7. Corolla - - - - - - 117 8. Stamens - - - - - 122 9. Disk - - - - - - 137 10. Pistillum - - - - - - 138 11. Receptacle - - - - 152 12. Ovulum - - - - - 153 13. Fruit - - - - - - 160 14. Seed - - - ... 181 15. Naked Seeds ... .... 194 CHAr. III. Of the Compound Organs in Flowerless Plants - - 195 1. Ferns ... _ j£, 2. Equisetaceae - - -- --197 3. Lycopodiacese - - - - - ib. 4. Marsileaceas - - - - 198 5. Mosses ------ 200 6. Hepaticae - 202 7. Lichens - - - ... 204 8. Algae ------ 206 9. Fungi - ----- 208 BOOK II. Physiology ; or, Plants considered in a State of Action - 210 Chap. I. Elementary Organs ----- 221 II. Root - - - - 227 III. Sap ... ... . 230 IV. Pith, Wood, and Bark - - - - 239 V. Leaves - - - 247 VI. Bracteae, Calyx, Corolla, and Disk - - - 257 VII. Stamens and Pistillum - - - - - 262 VIII. Fruit - - - - - 266 IX. Seeds ------- 270 X. Colour, Smell, and Taste - 274 XL Directions - - - - - 277 XII. Irritability - - - - - 288 XIII. Poisons ... - . 293 XIV. Diseases - ..... 297 XV. Hybrid Plants - - - - - - 501 XVI. Flowerless Plants ..... 305 CONTENTS. XV BOOK III. Taxonomy; or, of the Principles of Classification. Pago Chap. I. General Objects of Classification - - 306 II. Artificial Arrangements - ... 509 III. Natural System .... 31s IV. Speculative Modes of Arrangement - - - 324 V. Value of Characters ..... 549 VI. Species, Varieties, Genera, Orders, and Classes - - 365 BOOK IV. Glossology ; or, of the adjective Terms used in Botany - 369 Class I. Of Individual Terms - - - - 371 1 . Of Individual Absolute Terms : - - - 572 1. Figure - ib. 2. Division ..... 586 3. Surface ..... 593 4. Texture or Substance ... 397 5. Size - - - - - 599 6. Duration ..... 401 7. Colour ..... 402 8. Variegation or Marking ... 407 9. Veining - . - - ... 408 2. Of Individual Relative Terms : 1. ^Estivation - - - - 409 2. Direction - - - - - -411 5. Insertion, or Origin - - 415 Class II. Of Collective Terms : 1. Of Arrangement - - - - 417 2. Of Number ... . 420 Class III. Of Terms of Qualification - - 422 Signs - - - ib- Abbreviations - ... 425 XVI tONTENTS. HOOK V. Page Phytography; on. of the Rules to be observed in describing AND NAMING Pl.ANTS ...... 432 Chap. I. Diagnoses ; or, Generic and Specific Characters - 433 II. Descriptions - ... 440 III. Punctuation - - 4.52 IV. Nomenclature and Terminology - - - 454 V. Synonyms - - - - - 460 VI. Herbaria - - 463 VII. Botanical Drawings - 469 BOOK VI. Geografhy; or, the Distribution of Plants upon the Surface of the Globe - - - - . 470 BOOK VII. Morphology; or, of the Metamorphosis of Organs - - 504 Chap. I. Regular Metamorphosis .... 509 II. Irregular Metamorphosis - - - - 521 Explanation of the Plates - ... 527 Index : 1. Substantives - - . . 537 2. Adjectives - 546 INTRODUCTION TO BOTANY. BOOK I. ORGANOGRAPHY ; OR, OF THE STRUCTURE OF PLANTS. CHAPTER I. OF THE ELEMENTARY ORGANS. If plants are considered with reference to their internal organisation, they appear at first sight to consist of a vast multitude of exceedingly minute cavities, separated by a membranous substance ; more exactly examined, it is found that these cavities have a variety of different figures, and that each is closed up from those that surround it; and if the enquiry is carried still farther, it will be discovered that the partitions between the cavities are all double, and that by maceration in water, or by other processes, the cavities with their enclosing membrane may be separated from each other into distinct bodies. These bodies constitute what is called Vegetable Tissue, or Elementary Organs : they are the Simi- lar^ parts of Grew; the Tissu organique of Mirbel ; and the Parties elementaires, or Parties similaires, of De Candolle. The chemical basis of the elementary organs has been found to be oxygen, hydrogen, and carbon, with occasionally a little nitrogen or azote, combined in various proportions : their organic basis is membrane and fibre. The latter only are here to be considered. 2 ORGANOGRAPHY'. BOOK I. It is a common opinion that membrane only is the basis of the tissue of plants, and that fibre is itself a form of mem- brane. But as we find both developed in many of the most imperfectly organised plants, such as Scleroderma and other fungi ; and as it is difficult to conceive how that can be a mere modification of membrane which is generated inde- pendently of it, which has no external resemblance to it, and which is obviously something superadded, it will be better to consider both membrane and fibre as the organic bases of vegetable tissue, rather than the former only. The membrane varies in degree of transparency, being occasionally so exceedingly thin as to be scarcely discover- able, except by the little particles that stick to it, or by its refraction of light, and sometimes having a perceptible green colour, and a thickness which is considerable if compared with the diameter of the cavity it encloses. It generally tears readily, as if its component atoms do not cohere with greater force in one direction than another ; but I have met with a remarkable instance to the contrary of this in Bromelia nudi- caulis, in which the membrane of the cuticle breaks into little teeth of nearly equal width when torn. (Plate I. fig. 6.) It is in almost all cases destitute of visible pores ; although as it is readily permeable by fluids, it must necessarily be furnished with invisible passages. An opinion to the contrary of this has been held by some botanists, who have described the existence of holes or pores in the membrane of tissue, and have even thought they saw a distinct rim to them ; but this idea, which probably originated in imperfect observation with ill-constructed glasses, is now generally abandoned. The supposed pores, with their rim, have been ascertained to be nothing but grains of semi-transparent matter sticking to the membrane : this has been proved by Dutrochet, who found that boiling them in hot nitric acid rendered them opaque, and that dipping them in a solution of caustic potash re- stored their transparency, — a property incompatible with a perforation ; and any one furnished with a good modern microscope may satisfy himself upon the point, without going through Dutrochet's process ; by simple movement in water the grains may be often detached. It however occasionally CHAF. I. OF THE ELEMENTARY ORGANS. 3 happens that holes do exist in the membrane, of which men- tion will be made hereafter. Fibre may be compared to hair of inconceivable fineness, its diameter often not exceeding the -j^o of an inch. It has frequently a greenish colour, but is more commonly trans- parent and colourless. It appeal's to be sometimes capable of extension with the same rapidity as the membrane among which it lies, and to which it usually adheres ; but occasionally elongates less rapidly, when it is broken into minute portions, and carried along by the growing membrane. In direction it is variable (Plates I. and II.) ; sometimes it is straight, and attains a considerable length, as in some fungi ; some- times it is short and straight, but hooked at the apex, as in the lining of the anther of Campanula; occasionally it is straight, and adheres to the side of membrane, as in the same part in Digitalis purpurea ; but its most common direction is spiral. Whether it is solid or hollow has not been fully de- monstrated ; Dr. Purkinje asserts that it is hollow, as will be hereafter mentioned ; but there can be no doubt that it is also, at least sometimes, solid, as in the fibrous cellules of the leaf of Oncidium altissimum ; it is the opinion of many that it is hollow in the case of spiral vessels. Fibre has a constant tendency to anastomosing, in consequence of which reticulated appearances are occasionally found in tissue. The forms under which the elementary organs are seen are, — 1. Cellular tissue; 2. Woody Jibre ; and, 3. Vascular tissue. Sect. I. Of Cellular Tissue. Cellular tissue [Contextus cellulosus or Tela cellulosa, Lat. ; Ptdpa, Parenchyma, or pithy part, of old writers; Zellcngcwebe, Germ.;) generally consists of little bladders or vesicles of various figures, adhering together in masses. Occasionally it is composed of fibre only, unconnected by membrane. It is transparent, and in all cases colourless : when it appears otherwise, its colour is always caused by matter contained within it. n 2 ORGANOGRAPHY. BOOK I. If a thin slice of the pith of elder, or any other plant, be examined with a microscope, it will be found to have a sort of honeycomb appearance, as if there were a number of hexagonal l cavities, separated by partitions {fig. 1.). These little cavities are the inside of cellules of cellular tissue ; and the partitions are caused by the cohesion of their sides, as may be easily proved by boiling the pith a short time, when the cellules readily separate from each other. In pulpy fruits, or in those which have their cellular tissue in a loose, dry state when ripe, the cellules may be readily separated from each other without boiling. It was formerly thought that cellular tissue might be compared to the air bubbles in a lather of soap and water, while by some it has been supposed to be formed by the doublings and foldings of a membrane in various di- rections : on both these suppositions, the partitions between the cells would be simple, and not composed of two mem- branes in a state of cohesion. But the facility with which, as has been just stated, the cellules may be separated, sufficiently disproves these opinions. It is probable, however, that al- though the double nature of the partitions in cellular tissue may be demonstrated, yet that the cellules usually grow so firmly together, that their sides really form in their union but one membrane. Cellules are destitute of all perforation or visible pores, so that each is completely closed up from its neighbour, as far as we can see ; although, as they have the power of filter- ing fluids with rapidity, it is certain that they must abound in invisible pores, and that they are not impermeable, as if they were made of glass. An opinion different from this has been and is still entertained by some observers, who have described and figured perforations of the membrane in various plants. CHAP. I. OF THE ELEMENTARY ORGANS. 5 Mirbel states that " the sides of the cellules are sometimes 2 riddled full of holes (Jig. 2.), the aperture of which does not exceed the -jfo of a millimetre (or of half a line) ; or are less frequently pierced with transverse slits, which are occasionally 3 so numerous as to transform the cellules into a real articulated tissue, as in the pith of the Nelumbium (fig. 3.)." This statement is now so well known to have been founded upon inaccurate observation, and such pores or slits are so universally admitted to be small portions of amylaceous matter sticking to the walls, that no additional disproof seems necessary. A good microscope is alone suf- ficient, generally, to show the real nature of these supposed pores, or if not, the test used by Dutrochet, and mentioned at page 2., is in all cases sufficient. It may also be observed, that cellules often contain air- bubbles, which appear to have no direct means of escape, and that the limits of colour are often very accurately defined in petals, as, for instance, in the stripes of tulips and carnations, which could not be the case if cellular tissue were perforated by such holes as have been described ; for in that case colours would necessarily run together. One of the most striking instances with which I am ac- quainted, of cellular tissue having the appearance of pores is in Calycanthus, where it was pointed out to me by Mr. Va- renne. (Plate I. fig. 1.) But even in this, a careful ex- amination with glasses of different magnifying powers shows that the apparent pores are certainly not so, but composed of a solid subsance which may be distinctly seen by varying the direction of the rays of the transmitted light with which it is viewed. Sometimes they appear like luminous points ; by a little alteration of light they acquire a brownish tint ; and if seen with the highest powers of a compound microscope, where there is a great loss of light, they become perfectly opaque. Cellular tissue is always transparent and colourless, or at most only slightly tinged with green. The brilliant colours of vegetable matter, the white, blue, yellow, scarlet, and other hues of the corolla, and the green of the bark and leaves, is b 3 6 ORGANOGRAPHY. BOOK I. not owing to any difference in the colour of the cellules, but to colouring matter of different kinds which they contain. In the stem of Impatiens balsamina, a single cell is frequently red in the midst of others that are colourless. Examine the red cellule, and you will find it filled with a colouring matter of which the rest are destitute. The bright satiny appearance of many richly coloured flowers depends upon the colourless quality of the tissue. Thus, in Thysanotus fascicularis, the flowers of which are of a deep brilliant violet, with a remark- able satiny lustre, that appearance will be found to arise from each particular cellule containing a single drop of coloured fluid which gleams through the white shining membrane of the cellules, and produces the flickering lustre that is per- ceived. The colouring matter of the cellular tissue is fre- quently fluid, but is in the leaves and other parts more commonly composed of granules of various sizes ; this is particularly the case in all green parts ; in which the granules lie amongst greenish liquid, the latter of which, as they grow older, dries up, while the granules themselves gradually change to olive green, and finally to brown. Kieser distinguishes three sorts of globules among tissue : — 1. Round extremely transparent bodies, of a more or less regular figure, found principally in young plants and in coty- ledons, and soluble in boiling water : it is these that constitute starch or faecula. 2. Globules of a small size, a more irre- gular figure, and coloured either green or some other tint. They are not soluble in water, but are so in alcohol; but when dissolved, their matter is not precipitated by the addition of water, on which account they are distinguishable from resinous substances. 3. Extremely small round bodies, vary- ing in colour, and found floating in the proper juices of vegetables. The green granules are what M. Turpin calls Globuline. He believes them to be young cellules, and that it is from them that new tissue is developed. There does not, however, appear to be any evidence of this, which must be considered, at present, a gratuitous hypothesis, if, indeed, it were not rather said to be an untenable one. No one has ever seen the granules passing through the sides of the cellules ; no CHAP. I. OF THE ELEMENTARY ORGANS. 7 rupture of the sides of the cellules, caused by such a passage, has ever been detected ; and yet it seems inconceivable how the granules are to be constantly developing as new tissue, without some such trace of their passage being observable. Those who are curious to know the exact nature of this speculation, should consult the memoir of the author, in the Memoir rs du Museum, vol.18, p. 212. The mode in which cellular or any other tissue is really formed, is buried in mystery. It has been suspected by Mr. Valentine, and I be- lieve the same idea has also occurred to Mr. Bauer, that it may be caused by the extrication of gaseous matter among mucus ; but it is obvious that there are many difficulties in the way of this supposition. Amici says the cellules pullulate. The cellules develope, in some cases, with great rapidity. I have seen Lupinus polyphyllus grow in length, at the rate of an inch and a half a day. The leaf of Urania speciosa has been found by Mulder to lengthen at the rate of from one and a half to three and a half lines per houi', and even as much as from four to five inches per day. This may be computed to equal the developement of at least 4000 or 5000 cellules per hour. But the most remarkable instances of this sort are to be found in the mushroom tribe, which in all cases develope with surprising ra- pidity. It is stated by Junghuns, that he has known the Bovista giganteum, in damp warm weather, grow in a single night from the size of a mere point to that of a huge gourd. We are not further informed of the dimensions of this specimen ; but sup- posing its cellules to be not less than the ^o? of an inch in diameter, and I suspect they are nearer the ^o* it may be fairly estimated to have consisted, when full grown, of about 47,000,000,000 cellules ; so that, supposing it to have grown in the course of twelve hours, its cellules must have developed at the rate of near 4,000,000,000 per hour, or of more than sixty-six millions in a minute. The cellules of cellular tissue are always very small, but are exceedingly variable in size. The largest are generally found in the gourd tribe (Cucurbitaceae), or in pith, or in aquatic plants ; and of these some are as much as the 3-V of an inch in diameter ; the ordinary size is about the ^foj or the ^J^, and they are sometimes not more than the tt)W» Kieser has b 4 8 ORGANOGRAPHY. BOOK I. computed that in the garden pink more than 5,100 are con- tained in half a cubic line. Cellular tissue is found in two essentially different states, the membranous and the Jibrous. Membranous Cellular Tissue is that in which the sides consist of membrane only, without any trace of fibre ; it is the most common, and was, till lately, supposed to be the only kind that exists. This sort of tissue is to be considered the basis of vegetable structure, and the only form indispen- sable to a plant. Many plants consist of nothing else ; and while numberless vegetables are destitute of all other kinds of tissue, the membranous cellular tissue is never absent. It constitutes the whole of Mosses, Algae, Fungi, Lichens, and the like ; it forms all the pulpy parts, the parenchyma of leaves, the pith, medullary rays, and principal part of the bark, in the stem of exogenous plants, the soft substance of the stem of endogenous plants, the delicate membranes of flowers and their appendages, and both the hard and soft parts of fruits and seeds. It appears that the spheroid is the figure which should be considered normal or typical in this kind of tissue ; for that is the form in which cellules are always found when they are generated separately, without exercising any pressure upon each other ; as, for example, is visible in the leaf of the white lily, and in the pulp of the strawberry or of other soft fruits, or in the dry berry of the jujube. All other forms of the cellules are considered to be caused by the compression or extension of such spheroids. When a* mass of spheroidal cellules is pressed together equally in all directions, rhomboidal dodecahedrons are pro- duced, which, if cut across, exhibit the appearance of hexa- gons. (Plate I. fig. 12.) This is the state in which the tissue is found in the pith of all plants ; and the rice paper, sold in the shops for making artificial flowers, and for drawing upon, which is really the pith of a Chinese plant, is an excellent illustration of it. If the force of extension or compression be greater in one direction than another, various other forms are produced, of which the following have been observed : — 1. The oblong ; in the stem of Orchis latifolia, and in the inside of many leaves. (Plate I. fig. 9.) CHAP. I. OF THE ELEMENTARY ORGANS. 9 2. The lobed (Plate I. fig. 2.f) ; in the inside of the leaf of Nuphar luteum, Lilium candidum, Vicia Faba, &c. : in this form of cellular tissue the vesicles are sometimes oblong with a sort of leg or projecting lobe towards one end ; and some- times irregularly triangular, with the sides pressed in and the angles truncated. They are well represented in the plates of M. Adolphe Brongniart's memoir upon the Organisation of Leaves, in the Annales des Sciences, vol. xxi. 3. The square ,• in the cuticle of some leaves, in the bark of many herbaceous plants, and frequently in pith. (Plate I. fig. 13.) 4. The prismatical ; in some pith, in liber, and in the vicinity of vessels of any sort. (Plate 1. fig. 6.) 5. The cylindrical (Plate I. fig. 8. a) ; in Chara; this has been seen by Amici so large, that a single cellule measured four inches in length and one third of a line in diameter. (Ann. des Sciences, vol. ii. p. 246.) 6. The fusiform or the oblong pointed at each end ; in wood, and in the membrane that surrounds the seed of a Gourd. These are what Dutrochet calls clostres. (Plate II. fig. 19. 8.; Plate I. fig. 5.) 7. The muriform ; in the medullary rays. This consists of parallelopiped cellules compressed between woody fibre or vessels, with their principal diameter horizontal, and in the direction of the radii of the stem. It is so arranged that when viewed laterally it resembles the bricks in a wall ; whence its name. (Plate I. fig. 7.) 8. The compressed; in the cuticle of all plants. The cellules are often so compressed as to appear to be only a single membrane. (Plate I. fig. 2. a; Plate III. fig. 3, 4, &c.) 9. The irregular : in the testa of many seeds, as Casuarina : here the form of the cellules is so very irregular that they can be reduced to no certain form. 10. The sinuous ; in the cuticle, and also sometimes beneath it, as in the leaf of Lilium candidum. (Plate III. fig. 5.) Cellular tissue is frequently called Parenchyma. Professor Link distinguishes both Parenchyma and Prosenchyma ; re- ferring to the former all tissue in which the cellules are applied by their plane faces (Plate I. fig. 1. 3. 6, 7, &c), and 10 ORGANOGRAPHY. BOOK I. which consequently have truncated extremities ; and to the latter, forms of tissue in which the cellules taper to each end, and, consequently, overlap each other at their extremities. (Plate II. fig. 8. 19.) Fibrous cellular or Fibro-cellular Tissue is that in which the sides are composed either of both membrane and fibre together, or of fibre only. It is only lately that this kind has been recognised. The first observation with which I am acquainted is that of Mol- denhauer, who, in 1779, described the leaves of Sphagnum as marked by fibres twisted spirally. (Fig. 1. a, p. 4.) Link afterwards stated, that the supposed fibres were nothing but the lines where small cells contained in a larger one unite to- gether ; and his opinion was received. It is nevertheless cer- tain, that the tissue of Sphagnum is as Moldenhauer described it. In November, 1827, I described the tissue of Maurandya Barclayana as consisting of cellules formed of spiral threads crossing each other, interlaced from the base to the apex, and connected by a membrane. A few other solitary cases of the observation of this kind of tissue had subsequently oc- curred, when the admirable investigation of a modern anato- mist suddenly threw an entirely new light upon the subject, Instead of being very rare, cellular tissue of this kind appears to be found in various parts ; it has been already mentioned as existing in the leaves of Sphagnum ; it is also found in the pith of Rubus odoratus. I originally discovered it in the parenchyma of the leaves of Oncidium altissimum, and in the testa of various seeds. Mr. Griffiths has detected it abundantly in the aerial roots of Orchideous plants, observ- ations since confirmed by Mr. Brown ; and Dr. Purkinje has shown, by a series of excellent observations and drawings, that it forms the lining of the valyes of almost all anthers. The forms under which it exists in these parts are far more various than those of membranous cellular tissue. The principal varieties are these : — A. Membrane and Fibre combined. 1. Fibres twisted spirally, adhering to a spheroidal or angu- lar membrane, and often anastomosing irregularly, without the CHAP. I. OF THE ELEMENTARY ORGANS. 11 spires touching each other. (Plate I. fig. 12.) This is what is found in Oncidium altissimum leaves, in the aerial roots of some Orchideous plants, in the lining of many anthers, and is what Mohl has figured ( Ueber die Poren, <$~c. tab. i. fig. 9.), from the pith of Rubus odoratus. It approaches very nearly to the nature of spiral vessels, hereafter to be described, and ap- pears only to be distinguishable by the spires of the fibres not being in contact, being incapable of unrolling, having no elasticity or tenacity ; and by not being cylindrical and taper- ing to each end, but spheroidal. 2. Fibres crossing each other spirally, and forming a reticu- lated appearance by their anastomosing in oblong or botuli- form cells. Of this nature are the reticulated cells of the testa of Maurandya Barclayana, Wightia gigantea, and the like. (Plate I. fig. 1 1.) 3. Fibres running straight along the sides of truncated cylindrical cells in the anthers of Calla eethiopica and many other plants. (Plate I. fig. 13.) 4. Fibres running transversely in parallel lines round three of the sides of prismatical right-angled cells, in the anthers of Nymphseaceae, &c. 5. Fibres very short, attached to the sides of cells of various figures, to which they give a sort of toothed appearance, as in the anther of Phlomis fruticosa and other Labiatae. (Plate I. fig. 15.) The last three were first noticed by Dr. Purkinje. 6. The fibre twisted spirally, in the open membranous tubes that form the elaters of Jungermannia, apparently con- stitutes another form of tissue of this order. (Plate I. fig. 17.) B. Fibre tsoitJiout Membrane. 7. Spiral fibres repressed by mucus, but having sufficient elasticity to uncoil when the mucus is dissolved, and then breaking up into rings. (Plate I. fig. 16.) These are what are found in the testa of Collomia linearis. They approach spiral vessels so very nearly, that when I originally discovered them I mistook them for such. They are known by their roundish or depressed figure when at rest, and by the want <>f an inclosing membrane, and by their brittleness when un- coiled. I '2 ORGANOGRAPHY. BOOK I. 8. Fibres short, straight, and radiating, so as to form little starlike appearances, found in the lining of the anthers of Polygala Chamaebuxus, &c. by Dr. Purkinje. (Plate I. fig. 19.) 9. Fibres originating in a circle, curving upwards into a sort of dome, and uniting at the summit, observed by the same anatomist in the anthers of Veronica perfoliata, &c. 10. Fibres standing in rows, each distinct from its neigh- bour, and having its point hooked, so that the whole has some resemblance to the teeth of a currycomb, in the anthers of Campanula; first noticed by Dr. Purkinje. (Plate I. fig 18.) 1 1. Fibres forming distinct arches, as seen in the anthers of Linaria cymbalaria, &c. by Dr. Purkinje. (Plate I. fig. 4.)* In the centre of some of the cellules of the cellular tissue of many plants there is a roundish nucleus, apparently consisting of granular matter, the nature of which is unknown. It was originally remarked by Mr. Francis Bauer, in the cellules of the stigma of Phaius Tankervilliae. A few other vegetable anatomists subsequently noticed its existence ; and Mr. Brown, in his recent Memoir on the mode of impregnation in Orchi- deae and Asclepiadeae, has made it the subject of more ex- tended observation. According to this gentleman, such nuclei not only occasionally appear on the cuticle of some plants (Plate III. fig. 9.), in the pubescence of Cypripedium and others, and in the internal tissue of the leaves, but also in the cells of the ovulum before impregnation. It would also seem that Mr. Brown considers stomata to be formed by the juxta- position of two of these nuclei. Sect. II. Of Woody Fibre. This (Vasa fibrosa, Lat. ; Petits tubes, Mirb. ; Tissu cellu- laire allonge or ligneiuc, Fr. ; Vaisseaux propres fasciculaires, Mirb. ; Lignecsfistulce, Malpighi ; Fasergefdsse, or Bastrbhren, * According to the last mentioned author, the fibres themselves are generally tubular, and either perfectly round or somewhat compressed, or even three or four sided. He considers it proved, that they are hollow, by their appearance when compressed, by their occasionally containing bubbles of air, and by the difference between their state when dried and when recent. CHAP. I. OF THE ELEMENTARY ORGANS. l^J Germ.; perhaps the Vital vessels of Schultz;) consists of very slender transparent membranous tubes, tapering acutely to each end, lying in bundles, and, like the cellular tissue, having no direct communication with each other, except by invisible pores. (Plate II. fig. 1. a, b ,■ 2. 5. a, &c.) Many vegetable anatomists consider it a mere form of cellular tissue, in an elongated state; but scarcely with justice; for if this mode of viewing the subject were pushed a little farther, it would be necessary to refer every modification of tissue to the cellular, which would be obviously improper.* Woody fibre may be at all times known by its elongated figure and extremely attenuated character ; usually it has no sort of markings upon its surface, except occasionally a par- ticle or two of greenish matter in its inside ; but sometimes it is covered with spots that have been mistaken for pores, and that give it a peculiar character (Plate II. fig. 3. and 4.); and I have remarked an instance, in Oncidium altissimum, of its having tubercles on its surface. (Plate II. fig. 2.) Gene- rally, while cellular tissue is brittle, and has little or no cohe- sion, woody fibre has great tenacity and strength ; whence its capability of being manufactured into linen. Every thing pre- pared from flax, hemp, and the like, is composed of woody fibre. That even the most delicate of it consists of tubes, may be readily seen by examining it with a high magnifying power, and also by the occasional detection of particles of greenish matter in its inside. (Plate II. fig. 2. b.) A very different opinion has nevertheless been held by some physiologists, who have thought that the woody fibre is capable of endless divisibility. " When," says Duhamel, " I have examined under the microscope one of the principal fibres of a pear tree, it seemed to me to consist of a bundle of yet finer fibres ; and when I have detached one of those fibres, and submitted it to a more powerful magnifying power than the first, it has still * The distinction between cellular tissue and woody fibre is more pro- nounced in the long club-shaped aerial radicle of Rhizophora Candelaria, than in any plant with which I am acquainted. It there consists of large, very long, transparent tubes, lying imbedded in fine brownish granular matter, which is minute cellular tissue. 14 ORGANOGRAPHY. BOOK I. appeared to be formed of a great number of yet more delicate fibres." {Physique des Arbres, i. 57.) To this opinion Du Petit Thouars assents, conceiving the tenuity of a fibre to be infinite, as well as its extensibility. (Essais sur la Vegetation^ p. 150.) These vjjews have doubtless arisen from the use of very imperfect microscopes, under low powers of which such appearances as Duhamel describes are visible ; but with modern glasses, and after maceration in nitric acid, or even in pure water, each particular fibre can be separated with the greatest facility. Their diameter is often very much less than that of the finest human hair ; the tubes of hemp, for example, when completely separated, are nearly six times smaller. It must, however, be observed, that the fibres of this plant, as used in linen making, are by no means in a state of final separation, each of the finest fibres that meet the naked eye being in reality a bundle of tubes. While, however, some are of this extremely small size, others have a diameter as considerable as that of ordinary cellular tissue itself; in Coniferae the tubes are often ybts or -j^-q of an inch in diameter, and in the Lime they average about tto« Link states (Elementa, p. 85.) that they are very large in trees of hot countries, as, for instance, the Brazilian coffee. It has been asserted by some writers, that the tubes of the woody fibre are occasionally divided internally by transverse septa or partitions ; but the fact is denied by Link, who de- clares that " ejusmodi septa non existunt." It is no doubt true that, in general, there is no trace of such septa ; but I think it is impossible to deny their existence in the tissue of the Lime tree, at least. There are three distinct kinds of woody fibre : — 1. That in which the walls are not occupied with either granules or glands sticking to them, or in which the former are of very rare occurrence. (Plate II. fig. 1.) This is the finest and the commonest of all ; and is also the most genuine state of woody fibre. 2. That in which the walls have uniformly considerable numbers of granules of regular size sticking to them in a scattered manner. (Plate II. fig. 3, 4, 5.) These granules have been, and are still considered by many anatomists as CHAP. I. OF THE ELEMENTARY ORGANS. 15 pores in the sides of the tissue. They have been, in parti- cular, so described and represented lately by Mons. Brong- niart in Cycadese, in which the tubes are large, and the appearance very conspicuous. (Annates des Sciences, vol. xvi. tab. 21.) But I think it possible to demonstrate that this is an optical deception, and that the supposed perforations are of the same nature as the similar punctuations in cel- lular tissue, viz. semitransparent granules. In the first place, no colourless light passes through the supposed pores in any case ; on the contrary, they are dark, and have a solid appearance at all times, except when, at a certain distance out of the focus of the microscope, they become luminous. Secondly, if they were holes, they would, at least, be seen open when the tissue is dry and contracted, although they might close up when it becomes swollen with moisture. That, however, they never are: on the contrary, they are more opaque when dry than when wet. Thirdly, they be- come more and more opaque as the magnifying power with which they are viewed is increased ; a circumstance which seems incompatible with perforations. Finally, and it is this which will possibly be regarded most conclusive, if the tissue of Zamia be allowed to remain macerating for some time in dilute nitric acid, the apparent pores disappear : that is to say, the granules that cause the appearance of perforations are dissolved. It has been thought that such appearances as these were confined to Cycadese and Coniferae ; but I suspect that they are far from uncommon in other families. Such tissue constitutes a considei'able part of the wood of Calycan- thus (Plate II. fig. 4.), as has been already noticed ; and it is abundant in an East Indian genus allied to Trichopus. This kind of tissue might be called granular xwody Jibre : it ap- proaches very nearly to the character of ducts, into which, in Zamia, it seems to pass by almost insensible transitions. It may, however, be known from dotted ducts, either by its very acuminated extremities, or by its granules not being arranged in a spiral manner. 2. The third kind of woody fibre is the glandular. This has hitherto only been noticed in Conifers, in which it is uniformly found in every species. Its dimensions are more considerable than that of either of the last-mentioned forms; 16 ORGANOGRAPHY. BOOK f. f © I® % 1 1® h 0 © © <2>\ P) wu i g7 0 m (S) @ hi kx I (2) © Hn 1 and, like the second, it has been described as perforated with pores. It differs from granular woody fibre in the markings of the tube being vesicular, and usually transparent, with a darkened centre (Plate II. fig. 5, 6. 8.), which last is what has been described as a pore, the vesicle itself being considered a thickened rim. Kieser figures the glands as pores, both in Pine wood (Jig. 4.), and in Ephedra {fg- 5.), and in other cases also. They may be most conveni- ently found by examining a thin shaving of Pinus Strobus with a microscope, when they will be seen in the form of transparent globules, having a dark centre, and placed upon the walls of the tissue. That these globules are not pores, seems to me to be proved thus : they are flaccid when dry, and distend when moistened, which is not the property of a pore ; their centre is more generally opaque than transparent, which is also not the property of a pore ; they may be torn through the middle without any hole becoming visible ; and, finally, they may sometimes be detached from the tissue (Plate II. fig. 7.), or fall away spontaneously. In the latter case they leave a hole in the tissue at the place where they grew ; and holes thus occasioned misled Kieser into the belief that the woody fibre of Ephedra was really pierced with pores of considerable magnitude. An illustration of the manner in which these perforations are caused will be found in Plate II. fig. 7. M. Adolphe Brongniart has rightly stated, that there exists in Gnetum Gnemon a form of tissue exactly the same as in Coniferae. (Voyage de Freycinet.) In a species of that genus collected in Tavoy by Dr. Wallich, the glands are only different from what we commonly meet with in Coniferae, in being arranged side by side, instead of being placed in single rows irregularly one above the other. (Plate II. fig. 5.) Woody fibre constitutes a considerable proportion of the ligneous part of all plants ; it is common in bark, and it forms the principal portion of the veins of leaves, to which it gives stiffness and tenacity. CHAP. I. OF THE ELEMENTARY ORGANS. 17 Of Vascular Tissue. Vascular tissue consists of simple membranous tubes ta- pering to each end, but often ending abruptly, either having a fibre generated spirally in their inside, or having their walls marked by dots or transverse bars arranged in a spiral direction. Such appears to me to be the most accurate mode of de- scribing this kind of tissue, upon the exact nature of which anatomists are, however, much divided in opinion ; some be- lieving that the fibre coheres independently of any membrane, others doubting or denying the mode in which the vessels ter- minate ; some describing the vessels as ramifying ; and a fourth class ascribing to them pores and fissures, as we have already seen has been done in cellular tissue and woody fibre. It will be most convenient to consider all these points separately, along with the varieties into which vascular tissue passes. There are two principal kinds of vascular tissue ; viz. spiral vessels (Plate II. fig. 9. 11.), and ducts (Plate II. fig. 13. 15, 1G. 18. 20.). Spiral vessels (Jig.6,7.) {Vasa spiralia, Lat. ; Trachea of many ; Fistula spirales of Malpighi ; Spiralgefdsse or Schrcm- be?igcfasse, Germ. ;) are membranous tubes with conical extre- mities; their inside being occupied by a fibre twisted spirally, and capable of unrolling with elasticity. To the eye they, when at rest, look like wire twisted round a cylinder that is after- c 18 ORGANOGRAPHY. BOOK I. wards removed. For the purpose of finding them for examin- ation, the stalk of a strawberry leaf, or a young shoot of the Cornus alba (common dogwood) may be conveniently used ; in these they may be readily detected by gently pulling the specimen asunder, when they unroll, and appear to the naked eye like a fine cobweb. Very different opinions have been entertained as to the exact structure of spiral vessels. They have been considered to be composed of a fibre only, twisted spirally, without any connecting membrane ; or to have their coils connected by an extremely thin membrane, which is destroyed when the vessel unrolls ; or to consist of a fibre rolled round a membranous cylinder ; or even, and this was Malpighi's idea, to be formed by a spiral fibre kept together as a tube by interlaced fibres. Again, the fibre itself has been by some thought to be a flat strap, by others a tube, and by a third class of observers a kind of gutter formed by a strap having its edges turned a little inwards. Finally, the mode in which they terminate, although formerly stated by Mirbel to be continuous with the cellular tissue, is so little known, that the learned M. De Can- dolle, in his Organographies published in 1827, remarks, " Personne jusqu' ici n'a vu d'une maniere claire, ni I'origine, ni la terminaison d'un vaisseau." (P. 58.) As doubts upon these points arise from the extreme minuteness of the vessels, and from the different degrees of skill that observers employ in the use of the microscope, I can scarcely hope that any ob- servations of mine will have much weight. Nevertheless I may be permitted to state briefly what arguments occur to me in support of the definition of the spiral vessel as given above. With regard to the presence of an external membrane within which the spiral fibre is developed, it must be con- fessed that direct observation is scarcely sufficient to settle that point. It is easy to prove the existence of a membrane, but it is difficult to demonstrate whether it is external or internal with respect to the fibre. The best mode of examin- ation is to separate a vessel entire from the rest of the tissue, which may be done by boiling the subject, and then tearing it in pieces with the points of needles or any delicate sharp CHAP. I. OF THE ELEMENTARY ORGANS. 19 instrument ; the real structure will then become much more apparent than if the vessel be viewed in connection with the surrounding tissue. From some beautiful preparations of this kind by Mr. Valentine and Mr. Griffiths, it appears that the membrane is external : in the root of the Hyacinth, for example, the coils of the spiral vessel touch each other, except towards its extremities ; there they gradually separate, and it is then easy to see that the spiral fibre does not project beyond the membrane, but is bounded externally by the latter, which would not be the case if the membrane were internal : a representation of such a vessel is given at Plate II. fig. 9. Another argument as to the membrane being external may not unfairly be taken from the manifest analogy that a spiral vessel bears to that form of cellular tissue (p.l I.), in which a spiral fibre is generated within a cellule : it is probable that the origin of the fibre is the same in both cases, and that its position with regard to the membrane is also the same. It is much more difficult to determine whether the fibre is solid, or tubular, or flat like a strap ; and Amici has even declared his belief that the question is not capable of solution with such optical instruments as are now in use. When magnified 500 times in diameter, a fibre appears to be transparent in the middle, and more or less opaque at the edges ; a circumstance which has no doubt given rise to the idea that it is a strap or riband, with the edges either thickened, according to M. De Candolle* or rolled inwards according to Mirbel. But it is also the property of a trans- parent cylinder to exhibit this appearance when viewed by transmitted light, as any one may satisfy himself by examin- ing a bit of a thermometer tube. A better mode of judging is, perhaps, to be found in the way in which the fibre bends when the vessel is flattened. If it were a flat thread, there would be no convexity at the angle of flexure, but the external edo-e of the bend would be straight. The fibre, however, always maintains its roundness, whatever the degree of pres- sure that I have been able to apply to it. (Plate II. fig. 10.) This I think conclusive as to the roundness of the fibre ; but it does not determine the question of its being tubular or solid. I should have been induced to think, with Dr. Bischoff, c 2 20 ORGANOGRAPHY. BOOK I. who has investigated the nature of spiral vessels with singular skill [De vera Vasorum Plantarum spiralium Stmctura et Fwic- tione Commentation 1829) that it is solid, if it did not appear to have been ascertained by Hedvvig that, when coloured fluids rise in spiral vessels, they follow the direction of the spires. This fact may, however, be explained upon the sup- position that they rise in the channels formed by the approxi- mation of cylindrical fibres, and not in the fibres them- selves ; in which case there could be little doubt that the fibres are really solid. The nature of the termination of spiral vessels is now placed beyond all doubt, by the preparations of Mr. Valentine, above alluded to, and by some observations of my own. It is stated by Professor Nees von Esenbeck, in his Handbuch der Botanik, published in 1820, that they terminate in a conical manner; and in 1824 M. Dutrochet asserts, that they end in conical spires, the point of which becomes very acute ; but one would not suppose, judging from the figure given by the latter writer, that he had seen the terminations very clearly. It is, however, certain that the statement of Nees von Esenbeck is correct, and that the spiral vessel generally terminates in a cone. If the point of such a vessel in the Hyacinth (Plate II. fig. 9.) be examined, it will be seen that the end of the spiral fibre lies just within the acute point of the vessel, and that the spires become gradually more and more relaxed as they approach the extremity, as if their power of extension gradually diminished, and the mem- brane acquired its pointed figure by the diminution of elasti- city and extensibility in the fibre. It is not, however, always in a distinct membrane that the spiral vessel ends. In Ne- penthes the fibres terminate in a blunt cone, in which no membrane is discoverable. (Plate II. fig. 11.)* * A singular change occurs in the appearance of the spiral vessels of Nepenthes, after long maceration in dilute nitric acid, or caustic potash ; the extremities cease to be conical and spirally fibrous, but become little transparent oblong sacs, in which the spires of the fibres gradually lose themselves. This alteration, which is a very likely cause of deception, is perhaps owing to the extremities of the vessels being more soluble than the other part, the sac being the confluent dissolved fibres. This is in some measure confirmed by the subsequent disappearance of all trace of fibres in any part of the vessels, under the influence of those powerful solvents. CHAP. I. OF THE ELEMENTARY ORGANS. 21 A spiral vessel is formed by the convolutions either of a single spire, or of many. In the former case it is called simple, in the latter compound. The simple is the most com- mon. (Plate II. fig. 9.) Kieser finds from two to nine fibres in the Banana ; M. de la Chesnaye as many as twenty-two in the same plant. There are four in Nepenthes. (Plate II. fig. 11.) In general, compound spiral vessels are thought to be almost confined to Monocotyledonous plants, where they are very common in certain families, especially Marantaceae, Scitami- neae, and Musaceae ; but their existence in Nepenthes, and, according to Rudolphi, in Heracleum speciosum, renders it probable that future observations will show them to be not uncommon among Dicotyledons also. In Coniferae the spiral vessels have in some cases their spires very remote, and even have glands upon their mem- brane between the spires. (Plate II. fig. 6.) In size, spiral vessels, like other kinds of tissue, are vari- able; they are generally very small in the petals and filaments. Mirbel states them to be sometimes as much as the 288th of an inch in diameter ; Hedwig finds them, in some cases, not exceeding the 3000th ; a very common size is the 1000th. An irritability of a curious kind has been noticed by Mal- pighi in the fibre of a spiral vessel. He says (Anat. p. 3.) that in herbaceous plants and some trees, especially in the winter, a beautiful sight may be observed, by tearing gently asunder a portion of a branch or stem still green, so as to separate the coils of the spires. The fibre will be found to have a peristaltic motion which lasts for a considerable time. An appearance of the same nature has been described by Mr. Don in the bark of Urtica nivea. These observations are, however, not conformable to the experience of others. M. De Candolle is of opinion that the motion seen by Mal- pighi is clue to a hygroscopic quality combined with elasticity; and as spiral vessels do not exist in the bark of Urtica nivea, it seems that there is some inaccuracy in Mr. Don's remark. The situation of spiral vessels is in that part of the axis of the stem surrounding the pith, and called the medullary sheath, and also in every part the tissue of which originates from it ; such as the veins of leaves, and petals ; and of all other c 3 22 ORGANOGRAPHY. BOOK I. modifications of leaves. It has been supposed that they are never found either in the bark, the wood, or the root ; and this appears to be generally true. But there are exceptions to this : Mirbel and Amici have noticed their existence in roots; and Mr. Valentine and Mr. Griffiths have both ex- tracted them from the root of the Hyacinth ; they do not, how- ever, appear to have been hitherto seen in the roots of Dicoty- ledonous plants. I know of no instance of their existence in bark, except in Nepenthes, where they are found in prodigious quantities, not only between the alburnum and the liber, em- bedded in cellular tissue, as was first pointed out to me by Mr. Valentine, but also sparingly both in the bark and wood. They have been described by myself as forming part of the testa of the seed of Collomia, and Mr. Brown has described them as existing abundantly in that of Casuarina. In the former case, the tissue was rather the fibro-ce] hilar, as has been already explained (p. 11.); in the latter, they are appa- rently of an intermediate nature between the cellular -fibrous and the vascular; agreeing with the former in size, situation, and general appearance, but differing in being capable of un- rolling. In the stem of Monocotyledonous plants, spiral vessels occur in the bundles of woody tissue that lie among its cellular substance ; in the leaves of some plants of this de- scription they are found in such abundance, that, according to M. de la Chesnaye, as quoted by De Candolle, they are col- lected in handfuls in some islands of the West Indies for Amadou. The same author informs us, that about a drachm and a half is yielded by every plantain, and that the fibres may be employed either in the manufacture of a sort of down, or may be spun into thread. In Coniferous plants they are few and very small, and in Flowerless plants they are for the most part altogether absent ; the only exceptions being in Ferns and Lycopodiaceae, orders occupying a sort of middle place be- tween flowering and flowerless plants : in these thev no doubt exist. My friend Mr. Griffiths has succeeded in unrolling them in the young shoots of Lycopodium denticulatum. Some have thought that the spiral vessels terminate in those little openings of the cuticle called stomata ; but there does not seem to be any foundation for this opinion* CHAP. I. OF THE ELEMENTARY ORGANS. 23 Ducts {Jig. 8, 9, 10, 11,12.) (Fausses trachees, Fr.; Saft- rohren, Germ ; Tubes corpuscidiferes of Dutrochet, Lymph de- ducts, or Sap>-vessels of Grew and others ; Vaisseaux lympha- tiques of De Candolle, Vaisseaux pneumatiques of others ;) are membranous tubes, with conical or rounded extremities ; their sides being marked with transverse lines, or rings, or bars, or dots arranged spirally, and being incapable of unrolling. In some states these approach so nearly to the spiral vessel, that it is impossible to doubt their being a mere modification of it, as is the case in the annular duct (Plate II. fig. 13.); but in other states, as in the dotted duct, it is impossible to trace the transition from the one form to the other. Some writers confound all the forms under the common name of spiral vessels, but it is more convenient to consider them as distinct, not only because of their peculiar appearances, but because they occupy a station in plants in which true spiral vessels are not found ; and it is therefore probable that their functions are different. All the forms of the duct seem reducible to the following varieties : — 1. The Annular (Jig.ll., and Platell. fig. 1 3.). These are well described by BischofFas being formed of fibrous rings, placed at uncertain intervals ; or, to speak more accurately, they, like spiral vessels, are formed of a spiral thread, but it is broken at every coil, so as to separate into a number of distinct rings. These rings are included within a membranous tube, by which they are held together. When the rings are distant from each other (Plate II. fig. 1. b), the duct has a very peculiar appearance ; when the rings are packed together, so as to touch each other (Plate II. fig. 18.), the external appearance is exactly that of a spiral vessel, from which they are known by being incapable of unrolling. Both these forms are common in the soft parts of plants, particularly in the root, and also in Ferns and Lycopodiaceae among flower- less plants. 2. The Jleticidated (fig. 10. 12., and Plate II. fig. 1 3. a.). In these the spiral fibre, instead of separating into a number of distinct rings, is continuous in some places, anastomoses in others, so as to form a sort of netted appearance, or even c 4 24 ORGANOGRAPHY. BOOK I. breaks into short lengths, which, adhering to the sides of the membrane, give the vessel the appearance of having transverse bars. It is these appearances that have given rise to the notion of cracked, or pierced ducts {fig. 10.) existing in plants; the membrane between the spires, or bars, having been mis- taken for pores ; hence the term vaisseaufendu, used by Mirbel and others. Vessels of this kind are found in the stem of some herbaceous plants ; as, for example, the Impatiens Balsamina, in which they may be found in a great variety of states. 3. The Dotted {fig. 9.). Ducts of this kind are tubes having their sides marked with numerous dots, arranged in a more or less spiral manner, and being divided internally by transverse partitions. Usually, in addition to the dots, there is distinctly visible an oblique or annular transparent line upon the walls of the vessel. (Plate II. fig. 15. 17.) Hence Kieser con- sidered them as spiral vessels, the spires of which, when old, elongate, and become connected by a dotted membrane. Bischoff, on the contrary, considers the dots to be caused by the separation of a spiral fibre into extremely minute portions ; and he gives a figure (Plate II. fig 16.) of the manner in which he considers this change to occur. It is certain, however, that the dotted duct is really an en- tirely distinct kind of vessel, or at least a modification of cellular rather than of vascular tissue, as has been asserted by Du Petit Thouars {Ann. des Sciences, vol. xxi. p. 224.) ; for the following reasons : — If it were such a modification of the spiral vessel as Kieser supposes, it would have none of those internal septa by which it is particularly known. The same remark applies to the theory of Bischoff, which is also im- perfect, in not accounting for the nature of the transverse transparent lines that mark the sides of dotted ducts. Besides, the dotted ducts always terminate abruptly, not in acute cones, as has been seen by myself, and well represented by Mr. Griffiths, in his excellent illustrations of the anatomy of Phytocrene (Plate II. fig. 19. 20.), and they readily separate at the septa ; none of which properties are those of a spiral vessel. That the partitions above alluded to really exist, as has been correctly stated by Dr. Dutrochet, there can be no doubt, notwithstanding the denial of the fact by Link and CHAI\ I. OF THE ELEMENTARY ORGANS. 25 others. They may be seen with the naked eye in the ducts of the Cane, the Bamboo, and many other plants. While, therefore, I conform to the general practice of classing this kind of duct among vascular tissue, I would suggest that it should rather be considered as made up of cylindrical cells, the sides of which are covered with oblong granules, arranged with their principal axis across the tube, and the united ends of which cause the partitions discover- able upon a longitudinal section. It is these partitions that cause externally the appearance of transverse transparent lines. Dotted ducts are the largest of all kinds of tissue. The holes which are so evident to the naked eye, in a transverse section of the oak or the vine, are the mouths of dotted ducts; and the large openings in the ends of the woody bundles of Monocotyledonous stems, as in the Cane, are also almost always caused by the section of a dotted duct. The stem of Arundo Donax, or of any large grass, is an excellent subject for seeking them in ; they can be readily extracted from it when boiled. Vascular tissue always consists of tubes that are unbranched. They have been represented by Mirbel as ramifying in some cases ; but this opinion has undoubtedly arisen from imperfect observation. When forming a series of vessels, the ends of the tubes overly each other, as represented in Plate II. fig. 18. Some anatomists have added to the varieties above enumer- ated, what they call moniliform, or necklace-shaped, or stran- gulated vessels (jfg> 8.) (vaisscaux en chapelet or Strangles, vasa moniliformia, corpuscula vermiformia). These are rightly de- termined by BischofF to be mere accidental forms, caused by their irregular compression, when growing in knots or parts that are subject to an interrupted kind of developement. They may be found figured in Mirbel's Elemens, tab. x. fig. 15.; and in Kieser, fiff. 56. and 57. 26 ORGANOGRAPHY. BOOK I. Sect. IV. Of spurious elementary Organs ; such as Air Cells, Receptacles of Secretion, Glands, fyc. fyc. . The kinds of tissue now enumerated are all that have as yet been discovered in the fabric of a vegetable. There are. however, several other internal parts, which although not elementary, being themselves made up of some one or other of the forms of tissue already described, nevertheless have either been sometimes considered as elementary, or at least are not referable to the appendages of the axis, and can be treated of more conveniently in this place than elsewhere. These are, 1 . Intercellular passages ; 2. Receptacles of secre- tion ; 3. Air cells ; 4. Raphides. 1. Of Intercellular Passages. As the elementary organs are all modifications of either the spherical or cylindrical figure, it must necessarily happen that when they are pressed together, spaces between them will remain, which will be more or less considerable in proportion as the tissue departs in a greater or less degree from the cylindrical or spherical form. When the pressure has been very uniform, as in the case of the tissue of the cuticle, and in many states of cellular substance, no spaces will exist. When they do exist, they are called Intercellular passages [meatus or ductus intercellulares, canaux entrecellidaires). They neces- sarily follow the course of the tissue, being horizontal, ver- tical, or oblique, according to the direction of the angles of the tissue by which they are formed. Their size varies according to the size of the tissue, and the quantity of sap. In plants of a dry character, they are frequently so small as to be scarcely discoverable ; while in succulent plants they are so large as to approach the size of cellules, as in the stem of Tropaeolum majus. (Plate II. fig. 14-.) They are con- tinually filled with fluid, so long as the part of the plant in which they are situated performs its vital functions, and only become dry when it has ceased to live, as in dry pith. CHAP. I. OF THE ELEMENTARY ORGANS. 2/ 2. Of Receptacles of Secretion. But it frequently occurs that the simple intercellular pas- sages are dilated extremely by the secretions they receive, and either increase unusually in size, or rupture the coats of the neighbouring tissue; by which means cavities are formed replete with what is called the proper juice of the plant; that is to say, with the sap altered to the state which is peculiar to the particular species of tree producing it. Cavities of this nature are often called vasa propria ; they are the receptacula sued of Link; the vaisseaux propres of Kieser and De Can- dolle ; and the reservoirs du sue propre of the last author. To this class also are to be referred the turpentine vessels, and the milk vessels of Grew ; the reservoirs accidentels of M. De Candolle ; and also the reservoirs en caecum of the latter, which latter are the clavate vessels filled with oily fluid that are found in the coat of the fruit of Umbelliferae, and which are commonly called vittce. Although the receptacles of secretion have no proper coat, yet they are so surrounded by cellular tissue, that a lining or wall is formed, of perfect regularity and symmetry. The tissue of this lining is generally much smaller than that of the neighbouring parts. In figure the receptacles are extremely variable, most commonly round, as in the leaves of the Orange and of all Myrtacese, where they are called crypta, or glandulce impresses, or reservoirs vesicidaires, or glandes vesiculates, or receptacles of oil. In the Pistacia Te- rebinthus the receptacles are tubular ; in Coniferas they are very irregular in figure, and even position, chiefly forming large hollow cylindrical spaces in the bark. Those in the rind of the orange and lemon are little oblong or spherical cysts; their construction, which is very easily examined, gives an accurate idea of that of all the rest. (Plate II. fig. 21.) 3. Of Air Cells. Besides the common intercellular passages, and the recep- tacles now described, there is another and a very remarkable 28 ORGANOGRAPHY. BOOK I. sort of cavity among the tissue of plants. This is the air cell ; the lacuna of Link, the reservoir d'air and cellule d 'air of Kieser, and the luftbeh'dlter of the Germans. Like the re- ceptacles of secretion, the air cells are built up of tissue, but have no proper membrane of their own ; and this sometimes takes place with a truly wonderful degree of uniformity and beauty. Each cell is often constructed so exactly like its neighbour, that it is impossible to regard them always as mere accidental distensions of the tissue; on the contrary, they are, in those plants to the existence of which they are necessary, evidently formed upon a plan which is uniform in the species, and which has been wisely contrived by Providence in that manner which is most suitable to the purpose for which they are destined. They differ from receptacles of secretion in containing air only, and not the proper juice of the plant ; a peculiarity which is provided for by a curious contrivance of Nature. In receptacles, the orifices of the intercellular passages through which the fluid that is to be deposited drains, are all open ; but, to prevent any discharge of fluid into the air cells, the orifices of all the intercellular passages that would otherwise open into them are closed up. Air cells are very variable in size, figure, and arrangement. In the stems of fistular plants, as Allium, they from a cavity from the base to the summit ; in the stem of the Rush (Juncus articulatus), they consist of a number of tubular ca- vities placed one above the other, and separated by mem- branous partitions composed of a combination of minute cellules; in some aquatic plants they are very small, as in Butomus umbellatus. In form they are either cylindrical, or they assume the figure of the cellules by which they are formed, as in Limnocharis Plumieri (Plate III. fig. 1. and 2.), in which the structure of the air cells and their coats forms one of the most beautiful of microscopical objects. The inner surface of the air cells, when they are essential to the life of a plant, is smooth and uniform ; but in grasses, umbelliferous plants, and others where they are not essential, they seem to be caused by the growth of the stem being CHAP. I. OF THE ELEMENTARY ORGANS. 29 more rapid than the formation of the air cells ; so that the tissue is torn asunder into cavities of an irregular figure and surface. Kieser was the first to observe that in many plants in which the air cells of the stem are regularly separated by partitions, the intercellulary passages of the cellules form- iiirr- the partitions are sometimes left open, so that a free communication is maintained between all the tiers of air cells. (Plate II. fig. 2.) 4. Of Raphides. 14 15 Among the tissue, and particularly in the intercellular passages° of Monocotyledonous plants, are found certain needle-shaped transparent bodies, lying either singly or in bundles, and called raphides. They were first discovered by Rafa, who found them in the milky juice of Euphorbias ; afterwards they were met with by M. Jurine, in the leaves of Leucojum vernum, and elsewhere; and they are now well known to all vegetable anatomists. It is probable that the first discoverers considered them a kind of special organ ; but they have subsequently been recognised to be crystals of extreme minuteness, and, according to M. Raspail, of oxalate of lime. If a common Hyacinth is wounded, a considerable discharge of fluid takes place, and in this myriads of raphides are found floating ; or if the cuticle of the leaf of Mirabilis Jalapa is lifted up, little whitish spots are observable, which are composed of them; all these are acicular in form, whence their name. But in the Cactus peruvianus they are, according to M. Turpin, found in the inside of the vesicles of cellular tissue, and, instead of being needle-shaped, have the form of 30 ORGANOGRAPHY. BOOK I. extremely minute conglomerated crystals, which are rectangular prisms with tetraedral summits, some with a square, others with an oblong base. Crystals of a similar figure have been remarked by the same observer in Rheum palmatum ; and their presence, according to him, is sufficient to distinguish samples really from China and Turkey, from those produced in Europe. The former abound in these crystals, the latter have hardly any. In the above figure, 15 represents the raphides of Aloe verrucosa (from Kieser); 14, those of Cactus peruvianus; 13, those of Rheum palmatum : the two latter from Turpin. 31 CHAPTER II. OF THE COMPOUND ORGANS IN FLOWERING PLANTS. Having now explained the more important circumstances connected with modifications in the elementary organs of vegetation, the next subject of enquiry will be the manner in which they are combined into those masses which constitute the external or compound organs, or in other words the parts that present themselves to us under the form of roots, stems, leaves, flowers, and fruit, and that constitute the apparatus through which all the actions of vegetable life are performed. In doing this, I shall limit myself in the first place to Flower- ing Plants {Introduction to the Natural System, p. 1.); reserv- ing for the subject of a separate chapter the explanation of some of the compound organs of Flowerless plants [Ibid. p. 307. ), which differ so much in structure from all others, as to require in most cases a special and distinct notice. Sect. I. Of the Cuticle and its Appendages. 1. Of the Cuticle. Vegetables, like animals, are covered externally by a thin membrane or cuticle, which usually adheres firmly to the cellular substance beneath it. To the naked eye it appears like a transparent homogeneous pellicle, but under the mi- croscope it is found to be traversed in various directions by lines, which, by constantly anastomosing, give it a reticulated character. In some of the lower tribes of plants, consisting entirely of cellular tissue, it is not distinguishable, but in all others it is to be found upon every part, except the stigma and the spongioles of the roots. Its usual character is that of a delicate membrane, but in some plants it is so hard as almost to resist the blade of a knife, as in the pseudo-bulbs of 32 ORGANOGRAPHY. BOOK I. certain Orchideous plants. The most usual form of the reticulations is the hexagonal (Plate III. fig. 11.) : sometimes they are exceedingly irregular in figure; often prismatical ; and not unfrequently bounded by sinuous lines, so irregular in their direction as to give the meshes no determinate figure (fig. 5.). Botanists have not agreed as to the exact nature of the cuticle ; while the greater number incline to the opinion that it is an external layer of cellular tissue in a dry and compressed state; others, among whom are included both Kieser and Amici, consider it a membrane of a peculiar nature, transversed by veins, or vasa lymphatica. By the latter it is contended, that the sinuous direction of the lines in many cuticles is incompatible with the idea of any thing formed by the adhesion of cellular tissue; that when it is once removed, the subjacent tissue dies, and does not become cuticle in its turn, and that it may often be torn up readily without laceration. On the other hand, it is contended, that the reticulations of the cuticle are mostly of some figure analogous to that of cellular tissue, and that the sinuous meshes themselves are not so different as to be incompatible with the idea of a mem- brane formed of adhering cellules. We are accustomed to see so much variety in the mere form of all parts of plants, that an anomalous configuration in cellular tissue should not surprise us. The lines, or supposed vasa lymphatica, are nothing more than the united sides of the cellules, and are altogether the same as are presented to the eye by any section of a mass of cellular substance. It is certain that the cuticle cannot be removed without lacerating the subjacent tissue, with however much facility it may be sometimes separable: on the under surface of the leaf of the Box, for instance, there has plainly been some tearing of the tissue, before the cuticle acquired the loose state in which it is finally found. If the subjacent epidermis never becomes cuticle when the latter is removed, this is no reason why the cuticle itself should not be composed of cellular tissue ; for it is an axiom in vegetable physiology, that a part once fully formed is incapable of any subsequent change. Thus, pith never alters its dimensions, CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 33 after the medullary sheath that encloses it has been once completed, and a zone of wood never contracts or expands after it has been deposited : new matter may be added to any part, but the arrangement of the tissue, once fixed, re- mains unchangeable. The principal argument, however, in favour of cuticle being compressed cellular tissue, is, that in the cuticle of many plants the cellular state is distinctly visible upon a section (Plate I. fig. 2. a) ; that it even consists occasionally of several layers of cellules, as in many epiphytes of the Orchis tribe; and that, as there is no reason to doubt that Nature is as uni- form in the plan upon which cuticle is constructed as in all her other works, in those cases in which the cellular structure is less distinctly visible, we are nevertheless ' istified by sound philosophy in recognising it; while, on the other hand, it would be highly unphilosophical to suppose that the cuticle is formed in some plants upon one plan, and in others upon a totally different one. It may be farther remarked, that separable cuticle may often be traced into that which, being younger, is both inseparable and undistinguishable from the other cellular substance with which it is in contact, and from which it possesses no organic difference. There is some reason to suppose that there is occasionally present, on the outside of the cuticle, a transparent, very deli- cate membrane, having no organic structure, as far as can be discovered with the most powerful microscopes. Something of this kind has been noticed by M. Adolphe Brongniart in the Cabbage leaf, and an analogous structure has been re- marked by Professor Henslow in the Digitalis. 2. Of Stomata. In many plants the cuticle has certain openings of a very peculiar character, which appear connected with respiration, and which are called Stomata. (Plate III. passim.) Stomata [Pores of the epidermis ; Pores corticaux, allonge's, evajioratoires, or grands pores •, Glands corticales, miliaires, or epidermoi dales ; Gland/da? cutanea? ,■ Oejfnungen ,■ Stomal ia ,•) are passages through the cuticle, having the appearance of D 34. ORGANOGRAPHY. BOOK I. areolae, in the centre of which is a slit that opens or closes according to circumstances, and lies over a cavity in the sub- jacent tissue. There is, perhaps, nothing in the structure of plants upon which it is more difficult to form any satisfactory opinion than these stomata. Malpighi, and Grew, who seems first to have figured them {see his plate xlviii. fig. 4.), call them openings or apertures, but had no exact idea of their struc- ture. Mirbel also, for a long time, considered them pores, and figured them as such ; admitting, however, that he suspected the openings to be an optical deception. M. De Candolle entertains no doubt of their being passages through the cu- ticle. He says their edge has the appearance of a kind of oval sphincter, capable of opening and shutting. The mem- brane that surrounds this sphincter is always continuous with those which constitute the network of the cuticle : under the latter, and in the interval between the pore and the edge of the sphincter, are often found molecules of adhesive green matter {Organogr.'i. 80.); and recently M. Adolphe Brong- niart, in his beautiful figures of the anatomy of leaves, would seem to have settled the question beyond all dispute. {Annales des Scie?ices, vol. xxi.) Nevertheless, there are anatomists of high reputation who entertain a directly opposite opinion ; denying the existence of passages, and considering the sto- mata rather in the light of glands. Nees von Esenbeck and Link deny the existence of any perforation in the sto- mata, and consider that the supposed opening is a space more pellucid than the surrounding tissue, and that what seems a closed up slit is the thickened border of the space. Link further adds, that the obscuration of the centre of the stomata is caused by a peculiar secretion of matter, as is plainly visible in Baryosma serratum. {Elementa, p. 225.) To the views of these writers is to be added the testimony of Mr. Brown {Suppl. prim. Prodr. p. 1.), who describes the stomata as glands which are really almost always imperforate, with a disk formed by a membrane of greater or less opaqueness, and even occasionally coloured ; at the same time he speaks of this disk being, perhaps, sometimes perforated. In the midst of such conflicting testimony, an observer CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 35 necessarily finds much difficulty in fixing his opinion. I shall take the liberty of stating what I myself have seen, without, however, supposing that my humble authority can in any degree contribute to the determination of the question. In no plants are stomata larger than in some Monocotyledons ; they are, therefore, the best subjects for examination for general purposes. In Crinum amabile they evidently consist of two kidney-shaped bodies filled with green matter, lying upon an area of the cuticle smaller than those that surround it, and having their incurved sides next each other. In some, at the part where the kidney-shaped bodies come in contact, there is an elevated ridge, dark, as if filled with air, and having its principal diameter distinctly divided by a line. (Plate III. fig. 11.) In this state the stoma is at rest: but in others the kidney-shaped bodies are much more curved ; their sides are more separated from each other ; and there is no elevated ridge : at their former line of contact there is an opening so distinct and wide as to be equal to half the diameter of one of the kidney -shaped bodies; this, I presume, is the stoma open. That what is described to be an opening, is really so, seems to be demonstrated by the following tests : — 1. It is more transparent than any part of the most trans- parent portion of the cuticle ; 2. It admits transmitted light without interruption ; as is seen by gradually augmenting the magnifying power by which it is viewed, when the opening continues transparent, notwithstanding the great loss of light that attends the use of very high powers in compound mi- croscopes; and, 3. None of those arts which the microscopic observer knows so well how to employ, such as shifting, augmenting or decreasing the light, interposing moveable shadows between the mirror and the object, and the like, give the least indication of the presence of any membrane across the orifice of the stoma. I therefore conclude, that, in the Crinum amabile, the stomata are formed by two elastic reni- form cellules, lying over an opening in the centre of a con- tracted area of cuticle ; that these cellules, when expanded, meet, and press powerfully against each other, like two opposing springs ; thus causing the elevated ridge-like ap- pearance visible in the axis of the stoma in the figure above d 2 36 ORGANOGRAPHY. BOOK I. referred to ; and that, when contracted, they curve in an oppo- site direction, separating from each other, and ceasing to close up the aperture over which they lie. If it were possible to be absolutely certain of the accuracy of this description, the structure of the stoma in Crinum amabile might be safely taken as the type of all others ; for, no doubt, they are all constructed upon a similar plan. Without actually asserting so much as this, I may venture to state, that, of many hun- dreds of observations I have made upon this subject, I have not met with any thing that has led me to doubt the uni- formity of their nature, or their general accordance with what is found in Crinum amabile, whatever that may be. Or at least, the only difference is this, that while the two cellules that form the edges of the aperture are distinctly separated at their extremities in this plant, they are often confluent in others, as in Caladium esculentum. (Plate III. fig. 9.) Several varieties are represented at Plate III. ; besides which, they have been noticed by Link to be occasionally quadrangular, as in Yucca gloriosa (Plate III. fig. 10.), and Agave ameri- cana, and by Mr. Brown to be very rarely angular, of which, however, no instance is cited by that botanist. The former case is one in which the quadrangular figure is caused by the cellules being straight ; I am not aware if Mr. Brown means the same thing. I have never been so fortunate as to discover the mem- brane which Mr. Brown describes as generally overlying the apertures ; nor do I know of any other botanist having con- firmed that observation. Stomata are not found in Mosses, Hepaticae, Fungi, Algas, or Lichens (see Introduction to the Natural System)', in no submersed plants, or submersed parts of amphibious plants ; it is also said, not in Monotropa hypopithys, Neottia nidus avis, and Cuscuta europaea. They are not formed in the cuticle of plants growing in darkness ; are very small in trees and shrubs, particularly evergreens, and more espe- cially in such as have coriaceous leaves, and acrid or aromatic juices. {Iiudolphi.) They are not present upon roots, or the ribs of leaves. It also frequently happens that they are found upon one surface of a leaf, but not on another, and CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 37 generally in most abundance on the under side. In suc- culent plants, or in the succulent parts of other plants, they are either rare, or wholly wanting. They may be generally seen upon the calyx ; often on the corolla ; and rarely, but sometimes, upon the filaments, anthers, and styles. In fruit, they have only been noticed upon such as are membranous, and never upon the coat of the seed ; they exist, however, upon the surface of cotyledons. Mr. Brown thinks that the uniformity of the stomata, in figure, position, and size, with respect to the meshes of the cuticle, is often such as to indicate the limits, and sometimes the affinities, of genera, and of their natural sections. He has shown, with his usual skill, that this is the case in Pro- teaceae. He also remarks, that on the microscopic character of the equal existence of stomata on both surfaces of the leaf depends that want of lustre which is so remarkable in the forests of New Holland. {Journal of the Royal Geogr. Society, i. 21.) The same botanist is of opinion, that the two glands, or cel- lules, of which a stoma is composed, are each analogous to the single cellules found occupying the inner face of the meshes of the cuticle. (Plate iii. fig. 9.) See the Memoir on the Impregnation of Orchidece.) The surface of the cuticle is either perfectly smooth, or furnished with numerous processes, consisting of cellular tissue in different states of combination, which may be arranged under the heads of hairs, scales, glands, and jnnckles. All these oi-iginate either directly from the cuticle, or from the cellular substance beneath it ; never having any communication with the vascular or ligneous system. In Nepenthes the cuticle in the inside of the pitchers is pierced by a great number of holes, each of which is closed up by a firm thick disk of small cellular tissue, deep brown in colour, and connected with the cavernous parenchyma of the pitcher. Besides these, Nepenthes has also stomata. Nerium oleander and some other plants have, in lieu of stomata, cavities in the cuticle, curiously filled up or protected by hairs. (See Annales des Sciences, xxi. 438.) D 3 38 ORGANOGRAPHY. BOOK I. 3. Of Hairs. These (Jig.16.) are minute, transparent, filiform, acute processes, composed of cel- lular tissue more or less elon- gated, and arranged in a single row. They are found occasion- ally upon every part of a plant, even in the cavities of the and stem, as in Nymph ae a and other aquatic plants. In the Cotton Plant (Gossypium herbaceum, &c.) they form the substance which envelopes the seeds, and is wrought into linen; in the Cowhage (Mucuna urens and pruriens), it is they that produce the itching; and in the Palm tribe they are the long, entangled, soft, strangulated filaments that are used for Ama- dou. They vary extremely in length, density, rigidity, and other particulars ; on which account they have been distin- guished by the following names : — Down or Pubescence (pubes, adj. pubescens), when they form a short soft stratum, which only partially covers the cuticle, as in Geranium molle. Hairiness (hirsuties, adj. hirsutus), when they are rather longer and more rigid, as in Galeopsis Tetrahit. Pili (pilus, adj. pilosus), when they are long, soft, and erect, as in Daucus Carota. Villus (adj. villosus), when they are very long, very soft, erect, and straight, as in Epilobium hirsutum. Crini (adj. crinitus) are this variety in excess. Velvet (velumen, adj. vehrfinus), when they are short, very dense and soft, but rather rigid, and forming a surface like velvet, as in many Lasiandras. Tomentum (adj. totnentosus), when they ai'e entangled, and close pressed to the stem, as in Geranium rotundifolium. Cilice (adj. ciliatus), when long, and forming a fringe to a margin, like an eyelash, as in Sempervivum tectorum. Bristles (setce, adj. setosus), when short and stiff, as on the stems of Echium. Stings (stimuli, adj. stimulans; pili subulati of De Candolle), CHAP. II. COMPOUND ORGANS IN FLOWERING TLANTS. 39 when stiff and pungent, giving out an acrid juice if touched, as in the Nettle. Glandular hairs (pili capitati), when they are tipped with a glandular exudation, as in Primula sinensis. These must not be confounded with stalked elands. Hooks (hami, unci, roslella), when curved back at the point, as in the nuts of Myosotis Lappula. Barbs (glochis, adj. glochidatus), if forked at the apex, both divisions of the fork being hooked, as in the nuts of the same plant. Hairs also give the following names to the surface of any thing : — 1. Silhj (scriceus), when they are long, very fine, and pressed closely to the surface, so as to present a sublucid silky appearance ; ex. Protea argentea. 2. Arachnoid, when very long, and loosely entangled, so as to resemble cobweb : ex. Calceolaria arachnoidea. 3. Manicate, when interwoven into a mass that can be easily separated from the surface : ex. Cacalia canescens, Bupleurum giganteum. 4. Bearded {barbatus), when the hairs are long, and placed in tufts : ex. the lip of Chelone barbata. 5. Bough (asper), when the surface is clothed with hairs, the lower joint of which resembles a little bulb, and the upper a short rigid bristle : ex. Borago officinalis. Hairs are either formed of a single cell of cellular tissue (Plate I. fig. 8. b), or of several placed end to end in a single series (Plate I. fig. a, b.), whence, if viewed externally, they have the appearance of being divided internally by transverse partitions. They are sometimes divided into two or three forks at the extremity, as in Alyssum, some species of Apargia, &c. Occasionally they emit little branches along their whole length : when such branches are very short, the hairs are said to be toothed or toothleted, as in the fruit of Torilis Anthris- cus ; when they are something longer, the hairs are called branched, as in the petioles of the gooseberry ; if longer and finer still, the term is pinnate, as in Hieracium Pilosella; if the branches are themselves pinnate, as in Hieracium undulatum, d 4 40 ORGANOGRAPHY. BOOK I. the hairs are then said to be plumose. It sometimes happens that little branchlets are produced on one side only of a hair, as on the leaves of Siegesbeckia orientalis, in which case the hair is called one-sided {secundatus) ; very rarely they appear upon the articulations of the hair, which in that case is called ganglioneous. (Plate I. fig. 9. Verbascum Lychnitis) : \\~\e poils en goupillon of De Candolle are referable to this form. Be- sides these, there are many other modifications. Hairs are conical, cylindrical, or moniliform, thickened slightly at the articulations (torulose), as in Lamium album, or much en- larged at the same point (nodulose), as in the calyx of Achy- ranthes lappacea. Hairs are sometimes said to be Jixed by their middle (Plate I. fig. 10. c) ; a remarkable structure, common to many different genera; as Capsella, Malpighia, Indigofera, &c. This expression, however, like many others commonly used in botany, conveys a false idea of the real structure of such hairs. They are in reality formed by an elevation of one cel- lule of the cuticle above the level of the rest, and by the developement of a simple hair from its two opposite sides. Such would be more correctly named divaricating hairs. When the central cellule has an unusual size, as in Malpighia, these hairs are called poils en navette (pili Malpighiacei) by M. De Candolle ; and when the central cellule is not very apparent, poils en fausse navette (pili pseudo-Malpighiacei, biacuminati), as in Indigofera, Astragalus, Asper, &c. In many plants the hairs grow in clusters, as in Malvaceae, and are occasionally united at their base : such are called stellate, and are frequently peculiar to certain natural orders. (Plate I. fig. 10. a.) All these varieties belong to one or other of the two prin- cipal kinds of hairs ; viz. the Lymphatic and the Secreting. Of these, lymphatic hairs consist of tissue tapering gradually from the base to the apex ; and secreting, of cellules visibly distended either at the apex or base into receptacles of fluid. Malpighiaceous and glandular hairs, stings, and those which cause asperity on the surface of any thing, belong to the latter; almost all the other varieties to the former. When hairs arise from one surface only of any of the CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 41 appendages of the axis, it is almost always from the under surface ; but the seed leaves of the nettle, and the common leaves of Passerina hirsuta, are mentioned by M. De Candolle as exceptions to this rule : certain states of Rosa caniua might also be mentioned as exhibiting a similar phenomenon. When a portion only of the surface of any thing is covered by hairs, that portion is uniformly the ribs or veins. According to M. De Candolle, hairs are not found either upon true roots, except at the moment of germination, nor upon any part of the stem that is formed under ground, nor upon any parts that grow under water. 4. Of Scales. Scales are thin flat membranous processes, formed of cellular tissue springing from the cuticle. They may be con- sidered as hairs of a higher order, — as organs of the same nature, but more developed ; for they differ from hairs only in their degree of composition. They are of two kinds, Scales properly so called, and Momenta. Care must be taken not to confound scales of this description with scales of the stem, to be described hereafter : those now under consideration being mere processes of the cuticle ; those to be noticed hereafter being peculiar modifications of leaves. Scales, properly so called, are the small, roundish, flattened particles which give a leprous appearance to the surface of certain plants, as the Elaeagnus and the Ananassa. (Plate I. fig. 10. b.) They consist of a thin transparent membrane, attached by its middle, and, owing to the imperfect union towards its circumference, of the cellular tissue of which it is composed, having a lacerated irregular margin. A scale of this nature is called in Latin composition lepis, and a surface covered by such scales lepidotus, and not squamosus, which is only applied to a surface covered with the rudiments of leaves. Scales are the jjoils en ecusson {pill scutati) of De Candolle. Ramenta (Vaglnella:) are thin, brown, foliaceous scales, appearing sometimes in great abundance upon young shoots. They are particularly numerous, and highly developed, upon the petioles and the backs of the leaves of Ferns. They consist of cellular tissue alone, without any vascular 42 ORGANOGRAPHY. BOOK I. bundles, and are known from leaves not only by their anato- mical structure, but also by their irregular position, and by the absence of buds from their axillae. The student must particu- larly remark this, or he will confound with them leaves having a ramentaceous appearance, such as are produced upon the young shoots of Pinus. Link remarks, that they are very similar in structure to the leaves of mosses. The term striga has occasionally been applied to them (Dec. Theor. Elan. ed. 2. 376. Link, Elcm, 240.) ; but that word was employed by Linnaeus to designate any stiff bristle-like process, as the spiiies of the Cactus, the divaricating hairs of Malpighia, and the stiff stellated hairs of Hibiscus. So vague an application of the term is very properly avoided at the present day, and the substantive is rejected from modern glossology ; the ad- jective term strigose is, however, occasionally still employed to express a surface covered with stiff hairs. 5. Of Glands. Glands are elevated spaces in the stratum of parenchyma lying immediately below the cuticle, in which they cause pro- jections. They are of several kinds. Stalked glands are elevated on a stalk which is either sim- ple or branched : they secrete some peculiar matter at their extremities, and are often confounded with the glandular hairs above described, from which they have been well dis- tinguished by Link. According to that botanist, they are either simple or compound ; the former consisting of a single cell, and placed upon a hair acting as a simple conduit, oc- casionally interrupted by divisions ; the latter consisting of several cells, and seated upon a stalk containing several con- duits, formed by rows of cellular tissue. They are common upon the rose and the bramble, in which they become very rigid, and assume the nature of aculei. For the sake of dis- tinguishing them from the latter, they have been called sctce by Woods and myself, but improperly ; they are also the aigxdllons of the French. In Hypericum they abound on the calyx and corolla of some species, but do not give out any exudation ; they contain, however, a deep red juice within CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 43 their cells. In some Jatrophas they are much branched ; in many Diosmeee they form a curious humid appendage at the apex of the stamens. Sessile glands are produced upon various parts, and are extremely variable in figure. In Cassias, they are seated upon the upper edge of the petiole, and are usually cylin- drical or conical; in Cruciferous plants they are little roundish shining bodies, arising from just below the base of the ova- rium ; in the leafless Acacias, they are a little depressed, with a thickened rim, and placed on the upper edge of the phyllo- dium ; they are little kidney-shaped bodies upon the petiole of the Peach and other drupaceous plants ; and they assume many more appearances. Verruca, or warts, are roundish excrescences, formed of cellular tissue filled with opaque matter, and are situated upon various parts. They are common upon the surface of the leaves of the Aloe, where they are very large ; upon the stem, as in Euonymus verrucosus ; upon the petiole, as in Passiflora; they are also found upon the calyx, as in some species of Campanula, and at the serratures of the leaves, when they are considered by M. Roper (De Floribus Bal- saminearum, p. 15.) to be abortive ovula. They also appear upon the pericarpium and the testa of the seed; in the latter case they are called spongioid: seminales by De Candolle. They are round, oblong, or reniform, and occasionally cupu- late, when they receive the name of glandes a godet (glan- didte urceolares) from some French writers. Verrucas are the glandes ccllulaiTes of Mirbel ; but they must not be confounded with the glandes vasculaires of the same writer, which are not mere excrescences of the epidermis, but modifications of well known organs. (See Discus, further on.) The presence of minute verrucas upon the surface of a leaf gives rise to a peculiar kind of roughness which is called scabrities, and such a surface is then said to be scabrous (scaber) : this must not be confounded with asperity. Papilla; {Glandidce idriculaires of Guettard) are minute transparent elevated points of the cuticle, filled with fluid, and covering closely the whole surface upon which they appear. In other words, they are elevated, distended cellules of the 44< ORGANOGRAPHY. BOOK I. cuticle. The presence of papillae upon the leaves of the ice plant gives rise to the peculiar crystalline nature of its surface; they also cause the satiny appearance of the petals, upon which they almost always exist in great quantities. Link remarks, that the petals of Plantago, which are destitute of papillae, are also without the usual satiny lustre of those organs. When the papillae are much elongated beyond the surface, as in many stigmas, of which they form the collecting fringes, they receive sometimes the name of papulce. It should be observed, that in M. De Candolle's Theorie Elemen- taire, these two terms are transposed, each having received the definition belonging to the other. Lenticular glands (Lenticelles of De Candolle ; Glandes lenticulaires of Guettard ;) are brown oval spots found upon the bark of many plants, especially willows : they indicate the points from which roots will appear if the branch be placed in circumstances favourable to their production. They are considered by M. De Candolle to bear the same relation to the roots that buds bear to young branches. (De Candolle^ Pre- mier Mem. sur les Lentic., in the Ann. des Sciences Nuturellcs.) 6. Of Prickles. Prickles (aculei) are rigid, opaque, conical processes, formed of masses of cellular tissue, and terminating in an acute point. They may be, not improperly, considered as very compound indurated hairs. They have no connection with the woody fibre, by which character they are obviously distin- guished from spines, of which mention will be made under the head of branches, of which spines are an abortion. Prickles are found upon all parts of a plant, except the sti- pulae and stamens. They are very rarely found upon the corolla, as in Solanum Hystrix ; their most usual place is upon the stem, as in Rosa, Rubus, &c. Sect. II. Of the Stem or Ascending Axis. When a plant first begins to grow from the seed, it is a little body called an embryo, with two opposite extremities, of CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 45 which the one elongates in the direction of the earth's centre, and the other, taking a direction exactly the contrary, extends upwards into the air. This disposition to develope in two diametrically opposite directions is found in all seeds, pro- perly so called, there being no known exception to it; and the tendency is moreover so powerful, that, as we shall hereafter see (Book II.), no external influence is sufficient to overcome it. The result of this developement is the axis, or centre, round which the leaves and other appendages are arranged. That part of the axis which forces its way downwards, con- stantly avoiding light, and withdrawing from the influence of the air, is the descending axis, or the root ; and that which seeks the light, always striving to expose itself to the air, and expanding itself, to the utmost extent of its nature, to the solar rays, is the ascending axis, or the stem. As the double elonga- tion just mentioned exists in all plants, it follows that all plants must necessarily have, at an early period of their existence at least, both stem and root ; and that, consequently, when plants are said to be rootless, or stemless, such expressions are not to be considered physiologically correct. The Stem has received many names ; such as candex asce?ide?is, caudex intermedins, cidnms, stipes, truncus, and truncus ascendens. It always consists of bundles of vascular and woody tissue, embedded in cellular substance in various ways, and the whole enclosed within a cuticle. The manner in which these parts are arranged with respect to each other will be explained hereafter. The more immediate subject of consideration must be the parts that are common to all stems. 1. Of its Parts. Where the stem and root, or the ascending and descend- ing axes, diverge, there commences in many plants a differ- ence of anatomical structure, and in all a very essential physiological dissimilarity ; as will be hereafter seen. This portion of the axis is called the neck or collum, (coarclure of Grew, nceud vital of Lamarck, limes cojjummis, or fundus planter, of Jungius,) and has been thought by some to be the 46 ORGANOGRAPHY. BOOK X. seat of vegetable vitality ; an erroneous idea, of which more will be said in the next book. At first it is a space that we have no difficulty in distinguishing, so long as the embryo, or young plant, has not undergone any considerable change ; but in process of time it is externally obliterated ; so that in trees of a few years' growth its existence becomes a matter of theory, instead of being actually evident to our senses. Immediately consequent upon the growth of a plant is the formation of leaves. The point of the stem whence these arise is called the nodus (geniculum, Jungius; nceud, Fr. ;) and the space between two nodi is called an internodium (en- trenceud Fr.; merithallus, Du Petit Thouars). In internodia the arrangement of the vascular and fibrous tissue, of whatever nature it may be, of which they are composed, is nearly parallel, or, at least, experiences no horizontal interruption. At the nodi, on the contrary, vessels are sent off horizontally into the leaf; the general developement of the axis is mo- mentarily arrested while this horizontal communication is effecting, and all the tissue is more or less contracted. In many plants this contraction, although it always exists, is scarcely appreciable ; but in others it takes place in so re- markable a degree as to give their stems a peculiar character; as, for instance, in the Bamboo, in which it causes diaphragms that continue to grow and harden, notwithstanding the pow- erfully rapid horizontal distension to which the stems of that plant are subject. In all cases, without exception, a leaf-bud or buds is formed at a nodus immediately above the base of the leaf; generally such a bud is either sufficiently apparent to be readily recognised by the naked eye, or, at least, it becomes apparent at some time or other : but in certain plants, as Heaths, the buds are often never discoverable ; nevertheless, they always exist, in however rudimentary a state, as is proved by their occasional developement under favourable or uncom- mon circumstances. By some writers nodi, upon which buds are obviously formed, are called compound, or artiphyllous • and those in which no apparent buds are discoverable, are named simple, or plci plujilous : they are also said to be divided, when they do not surround the stem, as in the apple and other alternate-leaved genera; or entire, when they do surround CIlAr. II. COMPOUND ORGANS IN FLOWERING PLANTS. 47 it, as in grapes and umbelliferous plants : they are further said to he pervious, when the pith passes through them without interruption, or closed, when the canal of the pith is inter- rupted, as if by a partition. Pervious and divided, and closed and entire nodi usually accompany each other. For other remarks upon this subject, see Link's Elementa. All the divisions of a stem are in general terms called branches {rami) ; but it is occasionally found convenient to ex- press particular kinds of branches by special names. Thus, the twigs, or youngest shoots, are called ramuli or branchlets {brindilles or ramilles, Fr.), and by the older botanists Jlagclla : the assemblage of branches which forms the head of a forest tree is called the coma : cyma is sometimes used to express the same thing, but improperly. Shoots which have not completed their growth have re- ceived the name of innovations, a term usually applied to mosses. When such a shoot is covered with scales upon its first appearance, as the Asparagus, it is called turio : by the old botanists all such shoots were named asparagi. When a shoot is Ions and flexible, it receives the name of vimcn. This word, however, is seldom used ; its adjective being employed instead : thus, we say, rami viminei, or caulis vimineus ; and not vimen. From this kind of branch, that called a virgate stem, caulis virgatus, differs only in being less flexible and more riend. A vounsr slender branch of a tree or shrub is some- © Jo times named virgultum. When the branches diverge nearly at right angles from the stem, they are said to be brachiate. Small stems, which proceed from buds formed at the neck of a plant without the previous production of a leaf, are called cauliculi. Besides these terms, Du Petit Thouars employed cer- tain French words in a way peculiar to himself. The first young shoot produced during the year by a tree, he named scion ; any subsequent shoots formed by the scion, he termed ramilles ; the shoot that supports the scion was a ramcau; that which supports the rameau a branche ; and the trunk which bears the whole the tronc. Professor Link calls a stem which proceeds straight from the earth to the summit, bearing its branches on its sides, as Pinus, a caidis excurrens, and a 48 ORGANOGRAPHY. BOOK I. stem which at a certain distance above the earth breaks out into irregular ramifications, a caidis deliquescens. From the constitution and ramifications of their branches, plants are divided into trees, shrubs, and herbs. When the branches are perennial, and supported upon a trunk, a tree (arboy-) is said to be formed ; for a small tree, the term arbus- culus is sometimes employed. When the branches are peren- nial, proceeding directly from the surface of the earth without any supporting trunk, we have a shrub (frutex or arbustum, Lat., and arbrisseau, Fr.), which occasionally, when very small, receives the diminutive name of fruticulus. If a shrub is low, and very much branched, it is often called dumosus (subst. dumas) : this kind of shrub is what the French understand by their word buisson. The suffrutex, undcr-shrub, or sous-arbris- sean, differs from the shrub, in perishing annually, either wholly or in part ; and from the herb, in having branches of a woody texture, which frequently exist more than one year : such is the Mignonette (Reseda odorata) in its native country, or in the state in which it is known in gardens as the Tree Mignonette. The under-shrub is exactly intermediate between the shrub and the herb. All plants producing shoots of annual duration from the surface of the earth are called herbs. Some botanists distinguish two sorts of stems, the characters of which are derived from their mode of growth. When a stem is never terminated by a flower-bud, nor has its growth stopped by any other organic cause, as in Veronica arvensis, and all perennial and arborescent plants, it is said to be indeterminatus ; but when a stem has its growth uniformly stopped at a particular period of its existence by the production of a terminal bud, or by some such cause, it is called determi- natiis. The capitate and verticillate species of Mint owe their differences to causes of this nature; the stem of the former being determinate, the latter indeterminate. Some branches are imperfectly formed, lose their power of extension, become unusually hard, and acquire a sharp point. They are then called spines (spina), and must not be confounded with prickles, already described, from which they are distinguishable by their woody vascular centre. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 49 Occasionally, as in the Whitethorn, they bear leaves. In do- mesticated plants they often entirely disappear, as in the Apple and Pear, the wild varieties of which are spiny, and the culti- vated ones spineless. The point whence two branches diverge is called the axilla, or, in old botanical language, the ala. Leaf-buds {Gemma, Linn. Bourgeon, Fr.), being the rudi- ments of young branches, are of great importance in regard to the general structure of a plant. They consist of scales 17 18 imbricated over each other, the outermost being the hardest and thickest, and surrounding a minute axis which is in direct communication with the woody and cellular tissue of the stem. Linnaeus called this bud Hybemacidum, because it serves for the winter protection of the young and tender parts ; and dis- tinguished it into the Gemma, or leaf-bud of the stem, and the Bulb, or leaf-bud of the root. The leaf-bud has been compared by Du Petit Thouars and some other botanists to the embryo, and has even been de- nominated a Jixed embryo. This comparison must not, how- ever, be understood to indicate any positive identity between these two parts in structure, but merely an analogous func- tion, both being formed for the purpose of reproduction ; both in origin and structure they are entirely different. The leaf-bud consists of both vascular and cellular tissue, the embryo of cellular tissue only: the leaf-bud is produced without sexual intercourse, to the embryo this is essential : 50 ORGANOGRAPHY. BOOK I. finally, the leaf-bud perpetuates the individual, the embryo continues the species. The usual, or normal, situation of'leaf-buds is in the axillae of leaves; and all departure from this position is either irre- gular or accidental. Botanists give them the name of regular when they are placed in their normal station, and they call all others latent or adventitious. The latter have been found in almost every part of plants; the roots, the internodia, the petiole, the leaf itself, have all been remarked producing them. On the leaf they usually proceed from the margin, as in Malaxis paludosa, where they form minute granulations, first deter- mined to be buds by Professor Henslovv, or as in Bryophyl- lum calycinum and Tellima grandiflora ; but they have been seen by M. Turpin proceeding from the surface of the leaf of Ornithogalum. (fig. 19.) We are wholly unacquainted with the cause of the form- ation of leaf-buds ; all we know is, that they appear to pro- ceed from woody or vascular, and not from cellular tissue. There is, indeed, an opinion, which I believe is that of Mr. Knight, that the sap itself can at any time generate buds without any previously formed rudiment ; and that they de- pend, not upon a specific alteration of the arrangement of the vascular system, called into action by particular circum- stances, but upon a state of- the sap favourable to their creation. In proof of this it has been said, that if a bud of the Prunus Pseudo-cerasus, or Chinese Cherry, be inserted upon a cherry stock it will grow freely, and after a time will emit small roots from just above its union with the stock ; CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 51 at the time when these little roots are formed, let the shoot be cut back to within a short distance of the stock, and the little roots will then, in consequence of the great impulsion of sap into them, become branches emitting leaves. The leaf-buds of the deciduous trees of cold climates are covered by scales, which are also called tegmenta ; these afford protection against cold and external accidents, and vary much in texture, thickness, and other characters. Thus, in the Beech, the tegmenta are thin, smooth, and dry ; in many Wil- lows they are covered with a thick down ; in Popidus balsa- mifera they exude a tenacious viscid juice. In herbaceous plants and trees of climates in which vegetation is not exposed to severe cold, the leaf-buds have no tegmenta ; which is also, but very rarely, the case in some northern shrubs, as Rham- nus Frangula. The scales of the bud, however dissimilar they may be to leaves in their ordinary appearance, are nevertheless, in reality, leaves in an imperfectly formed state. They are the last leaves of the season, developed at a period when the current of vegetation is stopping, and when the vital powers have become almost torpid. That such is really their nature is apparent from the gradual transition from scales to perfect leaves, that occurs in such plants as Virburnum prunifolium, Magnolia acuminata, Liriodendron tulipifera, and iEsculus Pavia ; in the latter the transition is, perhaps, most satisfac- torily manifested. In this plant the scales on the outside are short, hard, dry, and brown ; those next them are longer, and greenish, and delicate ; within these they become dilated, are slightly coloured pink, and occasionally bear a few imperfect leaflets at their apex ; next to them are developed leaves of the ordinary character, except that their petiole is dilated and membranous like the inner scales of the bud ; and, finally, perfectly formed leaves complete the series of transitions. Among the varieties of root is sometimes classed what botanists call a bulb ,• a scaly body, formed at or beneath the surface of the ground, emitting roots from its base, and pro- ducing a stem from its centre. Linnaeus considered it the leaf-bud of a root; but in this he was partly mistaken, roots being essentially characterised by the absence of buds. He f. 2 52 ORGANOGRAPHY. BOOK I. was, however, perfectly correct in identifying it with a leaf- bud. A bulb has the power of propagating itself by deve- loping in the axillae of its scales new bulbs, or what gardeners call cloves, ( Cayeu, French ; Nucleus and Adnascens of the older botanists ; Adnatum of Richard ;) which grow at the ex- pense of their parent bulb, and eventually destroy it. Every true bulb is, therefore, necessarily formed of imbricated scales, and a solid bulb has no existence. The bulbi solidi, as they have been called, of the Crocus, the Colchicum, and others, are, as we shall hereafter see (see Cormus), a kind of ubterranean stem : they are distinct from the bulb in being not an imbricated scaly bud, but a solid fleshy stem, itself emitting buds. It has been supposed that they were buds, the scales of which had become consolidated; but this hypo- thesis leads to this very inadmissible conclusion, — that as the cormus or bulbus solidus of a Crocus is essentially the same, except in size and situation, as the stem of a Palm, the stem of a Palm must be a bulbus solidus also, which is absurd. In truth, the bulb is analogous to the bud that is seated upon the cormus, and not to the cormus itself; a bulb being an enlarged subterranean bud without a stem, the cormus a subterranean stem without one enlarged bud. Of the bulb, properly so called, there are two kinds. 1. The tunicated bulb (fig. 21.), of which the outer scales are thin and membranous, and cohere in the form of a distinct covering, as in the onion ; and, 2. the naked bulb [Bulbus CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 53 squamosus) (fig. 23. 22.), in which the outer scales are not mem- branous and coherent, but distinct and fleshy like the inner scales, as in Lilium. The outer covering of a bulb of the first kind is called the tunic. Besides the bulbs properly so called, there are certain leaf-buds, developed upon stems in the air, and separating spontaneously from the part that bears them, which are alto- gether of the nature of bulbs. Such are found in Lilium tigrinum, some Alliums, &c. They have been called bulbilli, propagines, sautilles, bacilli, &c. Care must be taken not to follow some botanists, in confounding with them the seeds of certain Amaryllideae, which have a fleshy testa; but which, with a vague external resemblance to bulbs, have in every in- spect the structure of genuine seeds. The tegmenta, or scales of the bud, have received the fol- lowing names, according to the part of the leaf of which they appear to be a transformation ; such terms are, however, but seldom employed : — 1. Foliacea, when they are abortive leaves, as in Daphne Mezereum. 2. Petiolacea, when they are formed by the persistent base of the petiole, as in Juglans regia. 3. Stipulacea, when they arise from the union of stipulae, which roll together and envelope the young shoot, as in Car- pinus, Ostrya, Magnolia, &c. 4. Fulcracea, when they are formed of petioles and stipules combined, as in Prunus domestica, &c. — [Rich. Nouv. Elem. 134. ed. 3.) The manner in which the nascent leaves are arranged with- in the leaf-bud is called foliation or vernation. The names applied to the various modifications of this will be explained in Glossology. They are of great practical importance both for distinguishing species, genera, and even natural orders; but have, nevertheless, received very little general attention. The vernation of Prunus Cerasus is conduplicate ; of Prunus domestica, convolute; of Filicesand Cycadeae, circinate. and so on. e 3 54- ORGANOCiltAPHY. BOOK I. 2. Of its External Modifications. It has already been stated, that the first direction taken by the stem immediately upon its developement is upward into the air. While this ascending tendency is by many plants maintained during the whole period of their existence, by others it is departed from at an early age, and a horizontal course is taken instead ; while also free communication with light and air is essential to most stems, others remain during all their lives buried under ground, and shun rather than seek the light. From these and other causes, the stems of plants assume a number of different states, to which botanists attach particular terms. It will be most convenient to divide the subject into the consideration of the varieties of — 1 . The subterranean stem ; and, 2. The aerial stem. The subterranean stem, often called soucheby the French, was confounded by all the older botanists, as it still is by the vulgar, with the root, to which it bears an external resem- blance, but from which it is positively distinguished both by its ascending direction, and by its anatomical structure. (See Root.) 24 25 The following are the varieties which have been distin- guished : — The Cormns, fig. 24. (Lecus of Du Petit Thouars, Plateau of De Candolle), is the dilated base of the stem of Monocoty- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 55 ledonous plants, intervening between the roots and the first buds ; and forming the reproductive portion of the stem of such plants when they are not caulescent. It is composed of cellular tissue, traversed by bundles of vessels and woody fibre, and has the form of a flattened disk. The fleshy root of the Arum, that of the Crocus and the Colchicum, are all differ- ent forms of the Cormus. It has been called bulbo-tuber by Mr. Ker, and bulbus solidus by many others ; the last is a contradiction in terms. (See Bulb.) The stems of Palms have by some writers been considered as an extended cormus, and not a true stem, but this seems an extravagant application of the term ; or rather an application which reduces the signification of the term to nothing. A cormus is a depressed subterranean stem of a particular kind; the trunk of a Palm is, as far as its external character is con- cerned, as much a stem as that of an oak. M. De Candolle applies the name cormus only to the stems of Cryptogamous plants, and refers to it the Anabices of Necker. The Tuber, fig. 25. (Tuberculum if very small), is an annual thickened subterranean stem, provided at the sides with latent buds, from which new plants are produced the succeeding year, as in the Potato and Arrow root. A tuber is, in reality, a part of a subterranean stem, excessively en- larged by the developement to an unusual degree of cellular tissue. The consequences frequently attendant upon a state of anamorphosis, as this is called, take place; the regular and symmetrical arrangement of the buds is disturbed, the buds themselves are sunk beneath the surface, or half obliterated, and the whole becomes a shapeless mass. Such is not, however, always the case ; the enlargement sometimes occurs without being accompanied by much distortion. In most, perhaps all, tubers, a great quantity of amylaceous matter is deposited, on which account they are frequently found to possess highly nutritive properties. The Creeping stern, fig. 26. (soboles), is a slender stem, which creeps along horizontally below the surface of the earth, emit- ting roots and new plants at intervals, as in the Triticum repens. e 4 56 ORGANOGRAPHY. BOOK I. This is what many botanists call a creeping root. It is one of those provisions of nature, by which the barren sands that bound the sea are confined within their limits ; most of the plants which cover such soils being provided with subterranean stems of this kind. Jt is also extremely tenacious of life, the buds at every nodus being capable of renewing the existence of the individual ; hence the almost indestructible properties of the Couch grass, Triticum repens, by the ordinary opera- tions of husbandry ; divisions of its creeping stem by cut- ting and tearing, producing no other effect than that of calling new individuals into existence as fast as others are destroyed. The term soboles is applied by Link and De Candolle to the sucker of trees and shrubs. (See Surculus.) Of the aerial stem, the most remarkable forms are the following : — The term stem (caulis) is generally applied to the ascending caudex of herbaceous plants or shrubs, and not to trees, in which the word truncus is employed to indicate their main stem ; sometimes, however, this is called caulis arboreus. From the caulis, Linnaeus, following the older botanists, distinguished the culmus or strata (Chawne, Fr.), which is the stem of Grasses; and M. De Candolle has further adopted the name Calamus (Chalumeau, Fr.) for all fistulous simple stems without articulations, as those of Rushes ; but neither of these differ in any material degree from common stems, and the employment of either term is superfluous. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 57 This has been already remarked with respect to Culmus by Link, who very justly enquires (Linnaea, ii. 235.) "cur Gra- minibus caulem denegares et culmum diceres?" The Runner, fig. 27. (sarmentum of Fuchsius and Linnaeus, coulant of the French,) is a prostrate filiform stem, forming at its extremity roots and a young plant, which itself gives birth to new runners, as in the Strawberry. Rightly considered it is a prostrate viviparous scape ; that is to say, a scape which pro- duces roots and leaves instead of flowers. It has been called flagellum by some modern botanists, but that term properly applies to the trailing shoots of the vine. The Sucker, fig. 29. (surculus), called by the French Dragon or Surgeon, is a branch which proceeds from the neck of a plant beneath the surface, and becomes erect as soon as it emerges from the earth, immediately producing leaves and branches, and subsequently roots from its base, as in Rosa spinosissima, and many other plants. Link applies the term soboles to this form of stem. From this has been distinguished by some botanists the Stole, {Stolo, Lat. ; and Jet, French ;) which may be considered the reverse of the sucker, from which it differs in proceeding from the stem above the surface of the earth, into which it afterwards descends and takes root, as in Aster junceus ; but there does not appear to be any ma- terial distinction between them. Willdenow confines the term surculus to the creeping stems of Mosses. By the older bota- nists a sucker was always understood by the word Stolo, and Surculus indicated a vigorous young shoot without branches. The shoots thrown up from the buried stems of Monocoty- ledonous plants, as the Pineapple for example, (the Adnata, Adnascentia, or Appendices of Fuchsius,) are of the nature of suckers. It may be here remarked, that Stolo. has given rise to the name Stool, which is applied to the parent plant, whence young individuals are propagated by the process of laying, as it is technically called by gardeners. The branch laid down was termed Propago by the older botanists, and the layer was called Malleolus, which literally signifies a hammer, and which was thus applied, because when the layer is separated from its 58 ORGANOGRAPHY. BOOK I. parent, its lower end resembles a hammer head, of which the new plant represents the handle. The Offset, fig. 30. {propaadum, Link), is a short lateral branch in some herbaceous plants, terminated by a cluster of leaves, and capable of taking root when separated from the mother plant, as in Sempervivum. It differs very little from the runner. The Rootstock, fig. 28. (rhizoma), is a prostrate thickened rooting stem, which yearly produces young branches or plants. It is chiefly found in Irideae and epiphytous Orchideae, and is often called Caudex repens. Link considers it scarcely dis- tinct from the sucker or stole. The old botanists called it Cervix, — a name now forgotten. The Vine, fig. 31. (viticula, Fuchs.), is a stem which trails along the ground without rooting, or entangles itself with other plants, to which it adheres by means of its tendrils, as the Cucumber and the Vine. This term is now rarely em- ployed. M. De Candolle refers it to the runner or sarmentum ; but it is essentially distinct from that form of stem. If a plant is apparently destitute of an aerial stem, it is technically called stemless [acaulis) ; a term which must not however be understood to be exact, because it is, from the nature of things, impossible that any plant can exist without a stem in a greater or less degree of developement. All that the term acaulis really means, is that the stem is very short. The Pseudobulb is an enlarged aerial stem, resembling a tuber, from which it scarcely differs, except in its being formed above ground, in having a dliticle that is often extremely hard, and in retaining upon its surface the scars of leaves that it once bore. This is only known in Orchideous plants, in which it is very common: the tuber of Arrow root is inter- mediate between the Pseudobulb and the genuine tuber. 3. Of its Internal Modifications. The internal structure of the stems of Flowering plants, is subject to two principal and to several subordinate modifica- tions. The former are well illustrated by such plants as the Oak CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 59 and the Cane, specimens of which can be easily obtained for comparison. A transverse slice of the former exhibits a cen- tral cellular substance or pith, an external cellular and fibrous ring or bark, an intermediate woody mass, and certain fine lines radiating from the pith to the bark, through the wood, and called medullary rays ; this is called Exogenous structure. In the Cane, on the contrary, neither bark, nor pith, nor wood, nor medullary rays, are distinguishable; the transverse section exhibits, on the contrary, a large number of hard spots caused by the section of bundles of woody tissue, and a mass of cellular substance in which they lie imbedded. This kind of structure is named Endogenous. In both cases there is a cellular and vascular system distinct from each other ; by a diversity in the respective arrangement of which the differences above described are caused. In de- scribing in detail the peculiar structure of Exogenous and Endogenous stems, it will be more convenient to consider them with reference to those two systems, than to follow the usual method of leaving the fact of there being two distinct systems out of consideration. § 1. Of the Exogenous Structure. The Cellular system in an Exogenous stem chiefly occupies the centre and the circumference, which are connected by thin 60 ORGANOGRAPHY. BOOK I. vertical plates of the same nature as themselves. The central part (a fig. 33.) is the pith, that of the circumference (b) is the bark, and the connecting vertical plates (c) are medullary rays. The pith is a cylindrical or an- gular column of cellular tissue, arising at the neck of the stem and terminating at the leaf-buds, with all of which, whether they are lateral or terminal, it is in direct communication. Its tissue, when cut through, almost always exhibits an hexagonal character, and is frequently larger than in any other part. When newly formed, it is green, and filled with fluid ; but its colour gradually disappears as it dries up, and it finally becomes colourless. After this it undergoes no further change, un- less by the deposition in it, in course of time, of some of the peculiar secretions of the species to which it belongs. It has been contended, indeed, by some physiologists, that it is gradually pressed upon by the surrounding part of the vas- cular system, until it is either much reduced in diameter or wholly disappears ; and in proof of this assertion, the Elder has been referred to, in which the pith is very large in the young shoots, and very small in the old trunks. Those, however, who entertain this opinion, seem not to consider that the diameter of the pith of all trees is different in different shoots, according to the age of those shoots; — that in the first that arises after germination, the pith is a mere thread, or at least of very small dimensions — that in the shoots of the succeeding year it becomes larger — and that its dimensions increase in proportion to the general rapidity of developement of the vegetable system : the pith, therefore, in the first-formed shoots, in which it is so small compared with that in the branches of subsequent years, is not so because of the pressure of surrounding parts ; it never was any bigger. The pith is always, when first forming, a uniform compact mass, connected without interruption in any part; but the vascular system sometimes developing more rapidly than CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 61 itself, it occasionally happens that it is either torn or divided into irregular cavities, as in the Horse Chestnut, the Rice-paper plant, and many others ; or that it is so much lacerated as to lose all resemblance to its original state, and to remain in the shape of ragged fragments adhering to the inside of the vas- cular system : this is what happens in Umbelliferous and other fistular-stemmed plants. Sometimes the pith is much more compact at the nodi than in the internodia, as in the Ash ; whence an idea has arisen that it is actually interrupted at those places : this is, however, an obvious mistake; there is no interruption of continuity, but a mere alteration in compactness. It very seldom happens that any part of the vascular system intermixes with the pith, which is almost always composed of cellular tissue exclusively ; but in Ferula and the Marvel of Peru, it has been proved by Messrs. Mirbel and De Candolle, that bundles of woody fibre are intermixed ; and in the Ne- penthes there is a considerable quantity of spiral vessels scattered among the cellular tissue of the pith. The Bark is the external coating of the stem, lying imme- diately over the wood, to which it forms a sort of sheath, and from which it is always distinctly separable. When but one year old, it consists of an exterior coating of cellular substance, called the cellular integument or the epidermis, and of an in- terior lining of woody fibre, called the liber or inner bark : if more than one year old, then it is composed of as many layers of cellular integument and woody fibre as it is years old, the former being invariably external, and the latter internal, in each layer ; and every layer being formed beneath the previous one, and therefore next the wood. In consequence of the new bark being continually generated within that of the pre- vious year, it is necessary that the latter, which is pushed outwards, should be extensible ; and in many plants this ex- tensibility takes place to a considerable degree. In the Apple, several successive zones of bark are formed without any appearance of a dislocation or disruption of the tissue of the outside; and in the Daphne Lagetto, the fibres of the liber are so tenacious that, instead of being ruptured by 62 ORGANOGRAPHY. BOOK I. the force of the inward growth, they are separated into lozenge-shaped meshes, arranged in such beautiful order, as to have acquired for the plant itself the name of the Lace Bark Tree. There exists, however, in all cases, a limit to the extensibility of the old layers of bark ; and when this ceases, the outer bark either splits into deep fissures, as in the Oak, the Elm, the Cork, and most of our European trees, or it falls away in broad plates, as in the Plane, or it peels off in long thin ribands, as in the Birch. As there is a double layer of cellular integument and woody fibre formed every year, it follows that the age of a tree ought to be indicated by the number of such deposits contained in its bark. But the arrangement of the zones is so very soon disturbed, and the distinction between them becomes so im- perfect, that even when the outermost coating is still entire, it is scarcely practicable to count the zones; and as soon as the outside begins to split or peel off, all traces of their full number necessarily disappear. That the bark really increases by constant deposits of new matter between it and the wood, is demonstrated by intro- ducing a piece of metal into the liber of a tree, and watching it subsequently : in process of time it will be protruded to the outside, and will finally fall away. Notwithstanding the fibrous character of a certain portion of the bark, it is generally so brittle as to be capable of breaking in all directions with a clean fracture, as soon as it becomes dry and ceases to live ; but in many plants, when young, it is so tough as to be applied to different economical purposes. For instance, the Russia mats of commerce are prepared from the liber of two or three species of Tilia, that of many Malvaceae is manufactured into cordage, and similar properties are found in that of many other plants. When stems are old, the bark usually bears but a small proportion in thickness to the wood ; yet in some plants its dimensions are of a magnitude that is very remarkable. For instance, Pinus Douglasii specimens have been brought to Europe twelve inches thick, and these are said not to be of the largest size. Lacunae and Vasa propria are exceedingly common in the CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 63 bark, but there is no well authenticated instance of any spiral vessels having been found in it; except in Nepenthes, in which they are found in almost every part, and exist in no inconsiderable numbers in the bark. Mr. Don states that spiral vessels abound in the bark of Urtica nivea, but I have not succeeded in discovering them there. Beneath the bark and above the wood is interposed in the spring a mucous viscid layer, which, when highly magnified, is found to consist of numerous minute transparent granules, and to exhibit faint traces of a delicate cellular organization. This secretion is named the Cambium, and appears to be exuded both by the bark and wood, certainly by the latter. The cellular system of the pith and that of the bark are, in the embryo, and youngest shoots, in contact ; but the vascular system, as it forms, gradually interposes between them, till after a few weeks they are distinctly separated, and in very awed trunks are sometimes divided by a space of several feet ; that is to say, by half the diameter of the wood. But whatever may be the distance between them, a horizontal communi- cation of the most perfect kind continues to be maintained. When the vascular system is first insinuated into the cellular system, dividing the pith and bark, it does not completely separate them, but pushes aside a quantity of cellular tissue, pressing it tightly into thin vertical radiating plates ; as the vascular system extends, these plates in- crease outwardly, continuing to maintain the connection between the centre and the circumference. Botanists call them medullary rays (or plates) ; and carpenters, the silver grain. They are composed of muriform cellular tissue (Plate I. fig. 7.), often not consisting of more than a single layer of cellules ; but sometimes, as in Aristolochias, the number of layers is very considerable (Plate II. fig. 12. a). In horizontal sections of an Exogenous stem, they are seen as fine lines radiating from the centre to the circumference ; in longitudinal sections they give that glancing satiny lustre which is in all discoverable, and which gives to some, such as the Plane and the Sycamore, a character of remarkable beauty. No vascular tissue is ever found in the medullary rays, 64 ORGANOGRAPHY. BOOK I. unless those curious plates described by Mr. Griffith in the wood of Phytocrene gigantea, in which vessels exist, should prove to belong to the medullary system. The vascular system in an Exogenous stem is confined to the space between the pith and the bark, where it chiefly consists of ducts and woody fibre collected into compact wedge-shaped vertical plates (fig. 33. d), the edges of which rest on the pith and bai'k, and the sides of which are in contact with the medullary rays. That portion of the vascular system which is first generated is in immediate contact with the pith, to which it forms a complete sheath, interrupted only by the passage of the medullary rays through it. It consists of spiral vessels and woody fibre intermixed, and forms an exceedingly thin layer, called the medullary sheath. This is the only part of the vascular system of the stem in which spiral vessels are ordinarily found ; the whole of the vessels subsequently depo- sited over the medullary sheath being ducts, and mostly dotted ones, with a few exceptions. The medullary sheath establishes a connection between the axis and all its append- ages, the veins of leaves, flowers, and fruits, being in all cases prolongations of it. It has been remarked by Senebier, and since by M. De Candolle, that it preserves a green colour even in old trunks, which proves that it still continues to retain its vitality when that of the surrounding parts has ceased. The vascular system of a stem one year old consists of a zone of wood lying between the pith and the bark, lined in the inside by the medullary sheath, and separated into wedge- shaped vertical plates by the medullary rays that pass through it. All that part of the first zone which is on the outside of the medullary sheath is composed of woody fibre and ducts intermixed in no apparent order ; but the ducts are generally either in greater abundance next the medullary sheath, or confined to that side of the zone, and the woody fibre alone forms a compact mass on the outside. The second year another zone is formed on the outside of the first, with which it agrees exactly in structure, except that there is no medul- lary sheath ; the third year a third zone is formed on the CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 65 outside the second, in all respects like it; and so on, one zone being deposited every year as long as the plant con- tinues to live. As each new zone is formed over that of the previous year, the latter undergoes no alteration of struc- ture when once formed : wood is not subject to distension by a force beneath it, as the bark is , but, whatever the first arrangement or direction of its tissue may be, such they re- main to the end of its life. The formation of the wood is, therefore, the reverse of that of the bark ; the latter increas- ing by addition to its inside, the former by successive de- posits upon its outside. It is for this reason that stems of this kind are called Exogenous (from two Greek words, sig- nifying to grow outwardly). According to M. Dutrochet, each zone of wood is in these plants separated from its neigh- bour by a layer of cellular tissue, forming part of the system of the pith and bark. After wood has arrived at the age of a few years, or some- times even sooner, it acquires a colour different from that which it possessed when first deposited, becoming what is called heart-wood, or duramen. For instance, in the beech it becomes light brown, in the oak deep brown, in Brazil wood and Guaiacum green, and in ebony black. In all these it was originally colourless, and owes its different tints to matter deposited at first in the ducts, and subsequently in all parts of the tissue ; as may be easily proved by throwing a piece of heart-wood into nitric acid, or some other solvent, when the colouring matter is discharged, and the tissue recovers its original colourless character. That part of the wood in which no colouring matter is yet deposited, and consequently that which, being last formed, is interposed between the bark and duramen, is called alburnum. The distinction between these is physiologically important, as will hereafter be explained. Each zone of the vascular system of an Exogenous stem being the result of a single year's growth, it should follow that to count the zones apparent in a transverse section is sufficient to determine the age of the individual under ex- amination ; and further, that, as there is not much difference in the average depth of the zones in very old trees, a certain rate of growth being ascertained to be peculiar to particular F 66 ORGANOGRAPHY. BOOK I. species, the examination of a mere fragment of a tree, the diameter of which is known, should suffice to enable the botanist to judge with considerable accuracy of the age of the indivi- dual to which it belonged. It is true, indeed, that the zones become less and less deep as a tree advances in age ; that in cold seasons, or after transplantation, or in consequence of any causes that may have impeded its growth, the formation of wood is so imperfect as scarcely to form a perceptible zone : yet the learned M. De Candolle has endeavoured to show, in a very able paper Sur la Longevite des Arbres, that the general accuracy of calculations is not much affected by such accidents ; occasional interruptions to growth being scarcely appreciable in the average of many years. This is possibly true in European trees, and in those of other cold or temperate regions in which the seasons are distinctly marked ; in such the zones are not only separated with tolerable distinctness, but do not vary much in annual dimensions. But in many hot countries the difference between the growing season and that of rest, if any occur, is so small, that the zones are as it were confounded, and the observer finds himself incapable of distinguishing with exactness the formation of one year from that of another. In the wood of Guaiacum, Phlomis fruticosa, Metrosideros polymorpha, and many other Myrtaceae, for instance, the zones are extremely indistinct ; in some Bauhi- nias they are formed with great irregularity ; and in Holbollia latifolia, some kinds of Ficus, certain species of Aristolochia, as A. labiosa, and many other plants, they are so confounded, that there is not the slightest trace of annual separation. With regard to judging of the age of a tree by the inspection of a fragment, the diameter of the stem being known, a little reflection will show that this is to be done with great caution, and that it is liable to excessive error. If, indeed, the zones upon both sides of a tree were always of the same, or nearly the same, thickness, much error would, perhaps, not attend such an investigation ; but it happens that, from various causes, there is often a great difference between the growth of the two sides, and consequently, that a fragment taken from either side must necessarily lead to the falsest inferences. For example, I have now before me four specimens of wood, taken almost at CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 67 hazard from among a fine collection, for which I am indebted to the munificence of the East India Company. The mea- surements of either side, and their age, as indicated by the number of zones they comprehend, are as follows : — Cornus capitata Pyrus foliolosa Magnolia insignis - Alnus napalensis - Diameter of Side A. Side B. 9 lines. 8 lines. 1 1 lines. 1 1 lines. 36 lines. 22 lines. 20 lines. 23 lines. Total. 45 lines. 30 lines. 31 lines. 34 lines. Real Age, or No. of Zones. 40 36 17 Now, in the first of these cases, suppose that a portion of the side A. were examined, and the observer were told that the diameter of the whole stem was 45 lines, he would find that each zone was 0.45 of a line deep, and that, consequently, the tree had been twenty years forming a radius of 9 lines, or in- creasing 18 lines in diameter: the total diameter being 45 lines, he would necessarily infer that the age of the tree was one hundred years ; its real age being only forty, as indicated by its zones. And so of the rest. When we hear of the Baobab trees of Senegal being 5150 years old, as computed by Adanson, and the Taxodium dis- tichum still more aged, according to the calculations of the ingenious M. Alphonse De Candolle, it is impossible to avoid suspecting that some such error as that just explained has vitiated their conclusions. To the characters above assigned to the stem of Exogenous plants there are several remarkable exceptions, some of which have been described by botanists ; others are mentioned now for the first time. M. Mirbel has noticed the unusual structure of Calycanthus, (Annates des Sciences, vol. xiv.) in the bark of which, at equal distances, are found four minute extremely eccentrical woody axes, the principal diameter of which is inwards ; that is to say, next the wood. The existence of this structure, noticed by the discoverer only in C. floridus, I have since ascertained in all the other species, and also in Chimonanthus. f 'J 68 ORGANOGRAPHY BOOK I. In Coniferous wood {fig. 34?.) there is scarcely any mixture of ducts among woody fibre, as in other exogenous plants ; in consequence of which a cross section exhibits none of those open mouths which are caused by the division of ducts, and which give what is vulgarly called po- rosity to wood. Instead of this, the vascular system generally consists exclu- sively of that kind of woody fibre which has been described at p. 15., under the name of glandular, with the exception of the medullary sheath, in which spiral vessels are present in small numbers. The Yew is the principal exception : in this plant the woody fibre is the same as that of other Coniferae ; but many tubes have a great quantity of little fibres lying obliquely across them at nearly equal distances, sometimes arranged with considerable regularity, — sometimes disturbed as it were, so that the transverse fibres, although they retain their obliquity, are not parallel, — and sometimes, but more rarely, so regular as to give to the tubes of woody fibre the appearance of spiral vessels, the coils of which are separated by considerable intervals. The latter only is represented by Kieser, at his tab. xxi. fig. 103, 104?.; but the former is by far the most common appearance. In Cycadeae the vascular system is destitute of ducts, as in Coniferae ; their place being supplied by such woody fibre as has been already described at p. 14. But the zones of wood are separated by a layer of cellular substance resembling that of the pith, and often as thick as the zones themselves. This structure is represented by M. Adolphe Brongniart, in the 16th volume of the Annales des Sciences. My friend Mr. Griffith has beautifully illustrated the struc- ture of a plant called Phytocrene {Jig. 35.), in Dr. Wallich's Plantse Asiatics?, vol. iii. t. 216. In this curious produc- tion, the wood consists of vessels encompassed by woody fibre ; and in the place of medullary rays are thick plates, con- nected neither with the medulla nor with the bark, nor even with each other in different zones. When the wood is dry, CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 69 these plates separate from the wood, in which they finally lie loose ; and, what is very remarkable, they contain vessels. 35 In Nepenthes distillatoria the pith contains a great quantity of spiral vessels; the place of the medullary sheath is occu- pied by a deep and very dense layer of woody fibre, in which no vessels, or scarcely any, are discoverable ; there are no medullary rays; the wood has no concentric zones; between the bark and the wood is interposed a thick layer of cellular tissue, in which an immense quantity of very large spiral ves- sels is formed ; on the outside of this layer is a thinner coating of woody fibre, containing some very minute spiral vessels ; and, finally, the whole is enclosed in a cellular inte- gument, also containing spiral vessels of small size. In this singular plant the outer layers are, it is to be presumed, liber and epidermis; and the cellular deposit between the former and the wood is analogous to cambium in an organised state, belonging equally to the wood and the bark. What is so exceedingly remarkable is the complete intermixture of the vascular and cellular systems, so that limits no longer exist between the two. I have a specimen of the twisted compressed stem of a Bauhinia from Colombia (Jig. 36.), in which there are no con- centric circles, properly so called ; but in which there are cer- tain irregular flexuous zones, consisting of a layer of cellular tissue coated by a stratum of woody fibre, enclosing, at irre- w 3 70 ORGANOGRAPHY. BOOK I. gular distances from the centre, very un- equal portions of the vascular system. The pith is exceedingly eccentrical ; and the medullary rays, which are very im- perfectly formed, do not all radiate from the pith, but on the thickest side form curves passing from one side of the stem to the other, their convexities turned to- wards the pith. In the stem of an unknown climber in my possession from Colombia {Jig. 37.), the vascular system is divided into four 37 In Hollbollia latifolia (Jig. 38.), which has atwin- ing stem, there are no con- centric circles, and the me- dullary rays are curved, part from right to left, and part from left to right, di- verging at one point and nearly equal parts, by four short thick plates radiating from the pith, and consisting of woody fibre with a very few ducts. These plates are not more than one third the depth of the wood; so that between their back and the bark there is a considerable vacancy, by which the four divisions of the vascular system are separated. This vacancy is nearly filled with bark, which projects into the cavity. 38 converging at another bark is tensive fo rations. pier ced longitudinal the with ex- per- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 71 In Euonymus tingens (fig. 39.) the vessels near the centre of the stem are arranged in con- centric interrupted circles, but towards the bark there is no trace of such circles; and the vessels are all confounded in an uniform mass. InMenispermum laurifolium (fig. 40.) the concentric lines evidently belong to the medullary sys- tem; they are extremely interrupted and unequal, often only half encir- cling the stem, or even less, and they anastomose in various ways ; the me- dullary rays are unusually large, and lie across the wood like parallel bars ; and, finally, the plates of which the wood consists each contain but one vessel, which is situated at the ex- ternal edge of the plate. None of the anomalous forms of 41 Exogenous stems are, however, so remarkable as an unknown Burmese p tree ( fig. 41.), for a specimen of which I * am indebted to my friend Dr. Wallich. In a section of this the general ap- pearance is so much that of an Endo- genous stem, that without an attentive examination it might be actually mis- taken for one. The diameter of this stem is two inches seven lines ; it is nearly perfectly circular, and has a very thin but distinct bark, with a central pith surrounded by very compact woody fibre. There are neither zones nor medullary rays ; but the vascular system consists of an uniform mass of ducts and woody fibre, disposed with great symmetry, and of the same degree of compactness at the circumference as well as in the centre. Amongst his wood are interspersed, at f 4 ORGANOGRAPHY. BOOK I. the distance of about half a line, with very great regularity, passages containing loose cellular tissue. These passages are convex at the back and rather concave in front, run parallel with the ducts, and do not seem to have any kind of com- munication with each other. They, no doubt, represent the medullary rays of the cellular system of this highly curious plant. It must be remarked, that the resemblance borne by this stem to that of an Endogenous plant is more apparent than real ; for whilst, in the latter, the vascular system is separated into bundles surrounded by the cellular system, in this, on the contrary, the cellular system consists of tubular passages surrounded by masses of the vascular system. These examples of anomalous structure will show the stu- dent that it is neither medullary rays nor concentric zones in the wood that are the certain indications of Exogenous growth ; both the one and the other being sometimes absent ; but that the presence of a central pith, and a greater degree of hardness in the centre than in the circumference, are the signs from which alone any absolute evidence can be derived. § 2. Of the Endogenous Structure. 42 Plants of an arborescent habit having this structure being almost exclusively extra-European, and most of them being CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 73 natives only of the tropics, botanists have had much fewer occasions of examining them, and, consequently, their know- ledge concerning them is far more limited. It is, therefore, probable that in this department of the subject there will be hereafter much to add and much to correct. In the mean while it must suffice, that what has been published concerning Endogenous plants is given with such corrections and additions as my own experience may have suggested. In Endogenous plants the vascular and cellular systems are as distinct as in Exogenous, but they are differently arranged. The cellular system, instead of being distinguishable into pith, bark, and medullary rays, is a uniform mass, in which the vascular system lies im- bedded in the form of thick fibres; and the vascular system itself has no tendency to collect into zones or wedges resembling wood, but in all cases retains the form of bundles resembling fibres. These bundles consist of woody fibre, enclosing spiral vessels or ducts ; most commonly the latter. The diameter of an Endogenous stem is increased by the constant addition of fibrous bundles to the centre, whence the name. Those bundles displace such as are previously formed, pushing them outwards; so that the centre, being always most newly formed, is the softest; and the outside, being older, and being gradually rendered more and more compact by the pressure exercised upon the bundles lying next it by those forming in the centre, is the hardest. In Endogenous plants that attain a considerable age, such as many Palms, this operation goes on till the outside becomes sometimes hard enough to resist the blow of a hatchet. It does not, however, appear that each successive bundle of fibres passes exactly down the centre, or that there is even much regularity in the manner in which they are arranged in that part: it is only certain that it is about the centre that they descend, and that on the outside no new formation takes place. This appears from the manner in which the bundles cross and interlace one another, as is shown in the figure of 74 ORGANOGRAPHY. BOOK I. Pandanus odoratissimus given by M. De Candolle in his Organographie (tab. vi.), or still more clearly in the lax tissue of the inside of the stems of Dracaena Draco. The epidermis of an Endogenous stem seems capable of very little distension. In many plants of this kind the diameter of the stem is the same, or not very widely different, at the period when it is first formed, and when it has arrived at its greatest age : Palms are, in particular, an instance of this ; whence the cylindrical form that is so common in them. That the increase in their diameter is really inconsiderable, is proved in a curious, and at the same time very conclusive, manner by the circumstance of gigantic woody climbing plants sometimes coiling round such stems, and retaining them in their embrace for many years, without the stem thus tightly wound round indicating in the slightest manner, by swelling or otherwise, that such ligatures inconvenience it. A specimen illustrative of this is preserved in the Museum of Natural History at Paris, and has been figured both by M. Mirbel in his Elemens (tab. xix.), and M. De Candolle in his Organographie (tab. iv.). We know, from the effect of the common Bindweed upon the Exogenous plants of our hedges, that the embrace of a twining plant is, in a single year, destructive of the life of every thing that increases in diameter; or at least produces, above the strangled part, extensive swellings that always end in death. It is, however, certain that other Exogenous plants do increase extensively in diameter up to a certain point; but this is effected with great rapidity ; and the horizontal growth once stopped appears never to be renewed. Thus, in the Bamboo, stems are sometimes found as much as two feet in circum- ference, which were originally not more than half an inch in diameter. Others would seem to have an unlimited power of distension. In the Dracaenas, called in French colonies in Africa Bois-chandelles, the first shoot from the ground is a Turio (sucker), an inch in diameter, and perhaps fifteen feet high ; but in time it distends so much that sometimes two men can scarcely embrace it in their extended arms. ( TJwu- ars, Essais, p. 3.) As Endogenous stems contain no concentric zones, there is CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 75 nothing in their internal structure to indicate age ; but, in the opinion of some botanists, there are sometimes external cha- racters that will afford sufficient evidence. It is said that the number of external rings that indicate the fall of leaves from the trunk of the Palm tribe coincide with the number of years that the individual has lived. There is, however, nothing like pi'oof of this at present before the public ; such statements must therefore be received with great caution. It may further be remarked, with reference to this subject, that in many Palms these rings disappear after a certain number of years. In arborescent Endogenous plants, it usually happens that only one terminal leaf-bud developes ; and in such cases the stem is cylindrical, or very nearly so, as in Palms. If two terminal leaf-buds constantly develope, the stem becomes dichotomous, but the branches are all cylindrical, as in Pan- danus and the Doom Palms of Egypt; but if axillary leaf- buds are regularly developed, as in the Asparagus, Dracaena Draco, or in arborescent grasses, then the conical form that prevails in Exogenous plants uniformly exists in Endogenous ones also. Besides the difference now mentioned, there is one other form of the Endogenous stem that it is necessary to describe ; viz. that of Grasses. In those plants the stem is hollow ex- cept at the nodi, where transverse partitions intercept the cavity, dividing it into many cells. In the Bamboo these cells and partitions are so large that, as is well known, lengths of that plant are used as cases to contain papers. In consequence of this great apparent deviation from the usual structure, a celebrated Swedish botanist has remarked, that Grasses are the least Endogenous of all Endogenous plants. But if the gradual developement of a grass be attentively observed, it will be found that the stem is originally solid ; that it then becomes hollow in consequence of its increasing in diameter more rapidly than new tissue can be formed ; and that, finally, in old arborescent stems, it again becomes solid by the constant addition of matter to its inside; so that its deviation from the ordinary characters of Endogenous structure i> much less considerable than it seems to be at first sight. 76 ORGANOGRAPHY. BOOK I. Sect. III. Of the Root, or descending Axis. At or about the same time that the ascending axis seeks the light and becomes a stem, does the opposite extremity of the seed or bud bury itself in the earth and become a root, with a tendency downwards so powerful, that no known force is sufficient to overcome it. Correctly speaking, nothing can be considered a root except what has such an origin ; for those roots which are emitted by the stems of plants, are in reality the roots of the buds above them, as will be hereafter explained. Nevertheless, nothing is more common than even forbotanists to confound subterranean stems or buds with roots, as has been already seen. (See Bulb, Tuber, Soboles, &c. &c.) Independently of its origin, the root is to be distinguished from the stem by many absolute characters. In the first place, its ramifications occur irregularly, without any sym- metrical arrangement: they do not, like branches, proceed from certain fixed points (buds), but are produced from all and any points of the root. Secondly, a root has no leaf-buds, unless indeed, as is sometimes the case, it has the power of forming adventitious ones ; but, in such a case, the irregular manner in which such are produced is sufficient evidence of their nature. Thirdly, roots have no scales, leaves, or other appendages ; neither do they ever indicate upon their surface, by means of scars, any trace of such : all underground bodies upon which scales have been found are stems, whatever they may have been called ; the only appendages they ever have are such things as the little hollow floating bladders found in Utricularia. A fourth distinction between roots and stems is, that the former have never any stomata upon their cuticle ; and, finally, in Exogenous plants, the root has never any pith. It has been also said that roots are always colourless, while stems are always coloured ; but aerial roots are often green, and all underground stems are colourless. The body of the root is sometimes called the caudex ; the minute subdivisions have been sometimes called radicular — a term that should be confined to the root in the embryo; others name them Jibrillcc, a term more generally adopted ; while the terms rhizina and rhizula have been given by Pro- fessor Link to the young roots of mosses and lichens. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 77 Kjibrilla is a little bundle of annular ducts, or sometimes of spiral vessels, encased in woody fibre, and covered by a lax cellular integument : it is in direct communication with the vas- cular system of the root, of which it is, in fact, only a sub- division; and its apex consists of extremely lax cellular tissue and mucus. This apex has the property of absorbing fluid with great rapidity, and has been called by M. De Candolle the Spongiole. It must not be considered a particular organ ; it is only the newly formed and forming tender tissue. In Pandanus the spongioles of the aerial roots consist of numerous very thin exfoliations of the epidermis, which form a sort of cup fit for holding water in. The proportion borne by the root to the branches is ex- tremely variable : in some plants it is nearly equal to them, in others, as in Lucerne (Medicago sativa), the roots are many times larger and longer than the stems ; in all succu- lent plants and Cucurbitaceae they are much smaller. When the root is divided into a multitude of branches and fibres, it is cal \edji brous : if the fibres have occasionally dilatations at short intervals, they are called nodulose. When the main root perishes at the extremity, it receives the name of prcemorse, or bitten off: frequently it consists of one fleshy elongated centre tapering to the extremity, when it is termed fusiform (or tap- rooted by the English, and pivotante by the French) ; or it dilates immediately below the surface of the earth into a globose form, when it is named turnip-shaped ', as in the common turnip ; if it is terminated by several distinct buds, as in some herbaceous plants, it is called many-headed {midticeps). The roots of many plants are often fleshy, and composed of lobes, which appear to serve as reservoirs of nutriment to the fibrillse that accompany them ; as in 44 many terrestrial Orchideous plants, Dahlias, &c. These must not be confounded either with tubers or bulbs, as they have been by some writers, but are rather to be con- sidered a special form of the root, to which the name of Pseudo-tuber {Jig. 44.) would" not be inapplicable. In Orchis the pseudo- tubers are often palmated or lobed ; in the 78 ORGANOGRAPHY. BOOK I. Dahlia, and many Asphodeleae, they hang in clusters, or are fasciculated. In internal structure the root differs little from the stem, except in being often extremely fleshy ; the cellular system being subject to an unusually high degree of developement in a great many plants, as the Turnip, the Parsnep, and other edible roots. In Endogenous plants, the mutual arrange- ment of the cellular and vascular systems of the root and stem is absolutely the same; but in Exogenous plants there is never any trace of pith in the root. Sect. IV. Of the Appendages of the Axis. From the outside of the stem, but connected immediately with its vascular system, arises a variety of thin flat expan- sions, arranged with great symmetry, and usually falling off after having existed for a few months. These are called, col- lectively, appendages of the axis ; and, individually, scales, leaves, bracteae, flowers, sexes, and fruit. They must not be confounded with mere expansions of the cuticle, such as ra- menta, already described (p. 41.), from which they are known by having a connection with the vascular system of the axis. Till lately, botanists were accustomed to consider all these as essentially distinct organs ; but, since the appearance of an admirable treatise by Goethe in 1790, On the metamorphosis of plants, proofs of their being merely modifications of one common type, the leaf, have been gradually discovered ; so that that which, forty years ago, was considered as the romance of a poet, is now universally acknowledged to be an indisputable truth. It is not my intention to enter into much separate discussion of this doctrine ; proof of it will be more conveniently adduced as the different modifications of the appendages of the axis come separately under consideration. The leaf, as the first that is formed, the most perfect of them all, and that which is most constantly present, is properly considered the type from which all the others are deviations, and is that with the structure of which it is first necessary to become acquainted. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 79 45 § 1. Of the "Leaf. 46 47 The leaf is an expansion of the bark at the base of a leaf- bud, prior to which it is developed. In most plants it con- sists of cellular tissue filling up the interstices of a net-work of fibres that proceed from the stem, and ultimately separating from the bark by an articulation ; in many Monocotyledonous plants, Ferns, and Mosses no articulation exists, and the base of the leaf only separates from its parent stem by rotting away. This difference of organisation has given rise to a distinc- tion, on the part of Oken, between the articulated leaves of Dicotyledones and the inarticulated leaves of Monocotyledones and Acotyledones : the former he calls true leaves, and dis- tinguishes by the name of Lcinb ; the latter he considers foliaceous dilatations of the stem, analogous to leaves, and calls Blatt. A leaf consists of two parts; namely, its stalk, which is called the petiole (fg. 46. a), and its expanded surface, which is called the lamina (fg. 46. c, b, d) : in ordinary language the latter term is not employed, but in very precise descriptions it is indispensable. The point where the base of the upper side of a leaf joins the stem is called the axilla ; any thing which arises out of that point is said to be axillary. If a branch or other process proceeds from above the axilla, it is called supra-axillary ,• if from below it, infra- axillary. 80 ORGANOGRAPHY. BOOK I. The scar formed by the separation of a leaf from its stem is called the cicatricula. The withered remains of leaves, which, not being articulated with the stem, cannot fall off, but decay upon it, are called reliquiae or induvice (debris, Fr.), and the part so covered is said to be induviatus. When leaves are placed in pairs on opposite sides of a stem (j%.51.), and on the same plane, they are called op- posite : if more than two are opposite, they then form what is called a whorl, or verticillus, and are said to be whorled, or verticillate: but if they arise at re- gular distances from each other round the stem, and not from the same plane> they are then called alternate. In plants having Exogenous stems, the first leaves, — namely, those which are present in the embryo itself (cotyledons), — are uniformly opposite; but those subsequently developed are either opposite, verticillate, or alternate in different species : on the contrary, in Endogenous plants, the embryo leaf is either solitary, or, if there are two, they are alternate ; and those subsequently developed are usually alternate also, but few cases occurring in which they are opposite. Hence some have formed an opinion that the normal position of the leaves of Exogenous plants is opposite, or verticillate ; and that when they are alternate, this arises from the extension of a nodus ; while that of Endogenae is alternate, the verticilli being the result of the contraction of internodia. But it seems more probable that the normal position of all leaves is alternate, and their position upon the stem an elongated spiral, as is in many cases exceedingly apparent, as, for instance, in the genus Pinus, in Pandanus, which is actually named Screw-pine, in consequence of the resem- blance its leaves bear to a screw, and in the Pine apple ; the Apple, the Pear, the Willow, the Oak, will also be found to indicate the same arrangement, which is only less apparent because of the distance between the leaves, and the irregu- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 61 larity of their direction. If, in the Apple tree, for instance, a line be drawn from the base of one leaf to the base of another, and the leaves be then broken off, it will be found that a per- fectly spiral line will have been formed. Upon this supposition, opposite or verticillate leaves are to be considered the result of a peculiar contraction or non-developement of internodia, and the consequent confluence of as many nodi as there are leaves in the whorl. The Rhododendron ponticum will fuiuiish the student with an illustration of this : on many of its branches the leaves are some alternate and some opposite ; and many intermediate states between these two will be perceivable. In many plants, the leaves of which are usually alternate, there is a manifest tendency to the appi-oximation of the nodi, and consequently to an opposite arrangement of the leaves, as in Solanum nigrum, and many other Solaneae * ; while, on the other hand, leaves that are usually opposite separate their nodi and become alternate, as in Erica mediterranea : but this is more rare. The best argument in support of the hypothesis that all verticilli arise from the contraction of internodia and con- fluence of nodi, is, however, to be derived from flowers, which are several series of verticilli, as will be seen hereafter. In plants with alternate leaves, the flowers often change into young branches, and then the verticilli of which they consist are broken, the nodi separate, and those parts that were before opposite become alternate; while, in monstrous Tulips, the verticilli of which the flower consists are plainly shown to arise from the gradual approximation of leaves, that in their unchanged state are alternate. In this normal state leaves are obviously distinct, both from each other and from the stem. But, in some cases, adhesions of various kinds occur, and give them a new character. Thus, in Cardui, and many other thistle-like plants, the elongated bases of the leaves adhere to the stem, and become what is called decurrent. In Bupleurum perforatum the lobes of the base of the leaf not only cohere with the stem, but, projecting beyond it, grow together, so as to resemble a leaf through Introduction to the Natural S3 stem of Botany, p. 231. G 82 ORGANOGRAPHY. BOOK I. which the stem has pierced : this is called being -perfoliate. Frequently two opposite leaves grow together at the base, as in Caprifolium perfoliatum ; to this modification the latter term is often also applied, but that of connate is what more properly belongs to it. The anatomical structure of the leaf is this: — From the medullary sheath diverges a bundle of woody tissue, accom- panied by spiral vessels : this passes through the bark, and proceeds, at an angle more or less acute, to a determinate distance from the stem, branching off at intervals, and, by numerous ramifications, forming a kind of net-work. At the point of the stem whence the bundle of fibrovascular tissue issues, the cellular tissue of the bark also diverges, accom- panying the fibrovascular tissue, expanding with its ramifi- cations, and filling up their interstices. The tissue that proceeds from the medullary sheath, after having passed from the origin of the leaf to its extremity, doubles back upon itself, forming underneath the first a new layer of fibre, which, upon its return, converges just as the first layer diverged, at length combines into a single bundle, corre- sponding in bulk and position to that which first emerged, and finally discharging itself into the liber. If, therefore, a section of the leaf and stem be carefully made at a nodus, it will be found that the bundle of woody tissue which forms the frame-work of the leaf communicates above with the medullary sheath, and below with the liber. This is easily seen in the spring, when the leaves are young; but is not so visible in the autumn, when their existence is drawing to a close. The double layer of fibrovascular tissue is also perceptible in a leaf which has laid during the winter in some damp ditch, where its cellular substance has decayed, so that the cohesion between the upper and lower layers is destroyed: they can then be easily separated. The curious Indian leaves which have the property of opening, upon slight violence, like the leg of a silk stocking, so that the hand may be thrust between their upper and lower surfaces, derive that singular separa- bility from an imperfect union between the layer of excurrent and recurrent fibre. M. De Candolle remarks, that when the fibres expand to form the limb of a leaf, they may (whether this phenomenon occurs at the extremity of a petiole, or at CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 83 the point of separation from the stem,) do so after two different systems : they may either constantly preserve the same plane when the common flat leaves are formed ; or they may expand in any direction, when cylindrical, or swollen, or triangular leaves are the result. (Organogr. p. 270.) The cellular tissue of which the rest of the leaf is composed is parenchyma, which Link then calls diachijma, or that im- mediately beneath the two surfaces cortex, and the intermediate substance diploe. M. De Candolle calls these two, taken to- gether, the mesaphyllum. The whole is protected, in leaves exposed to air, by a coating of cuticle, furnished with stomata ; but in submersed leaves the parenchyma is naked, no cuticle overlaying it. The general nature of the parenchymatous part of leaves has been very well explained, both by Link and others, and figured by Dr. Mohl, in 1828. (Uber die Poren des Pflanzcn- zellgewebes, tab. i. fig. 4, &c.) But the most complete a°count is that of M. Adolphe Brongniart, in 1830 (A/males des Sc. vol. xxi. p. 420.), of which the principal part of what follows is an abstract. The cuticle is a layer of cellules adhering firmly to each other, and sometimes but slightly to the subjacent tissue, from which they are entirely different in form and nature : in form, for the cellules are depressed, and, in consequence of the variety of outline that they present, form meshes either regular or irregular ; and in nature, because these cellules are perfectly transparent, colourless, and probably filled with air, — for the manner in which light passes through them proves that they do not contain dense fluid. They scarcely ever contain any organic particles, and are probably but little permeable either to fluids or gaseous matter; while, on the other hand, the cellules of the subjacent parenchyma are filled with the green substance that determines the colour of the leaf. The cuticle is not always formed of a single layer of cellules, but in some cases consists of two, or even three. No trace whatever is discoverable of vessels either terminating in or beneath the cuticle; M. Brongniart states this most explicitly, and my own observations are entirely in accordance with his : an opinion, therefore, which some distinguished botanists have g 2 84- ORGANOGRAPHY. BOOK I. entertained, that spiral vessels terminate in the stomata (D. C. Omanogr. p. 272, &c), must hereafter be abandoned. At the margin of a leaf the cuticle is generally harder than elsewhere, and sometimes becomes so indurated as to assume a flinty texture, as in the Aloe, and many other plants. Stomata (p. 33.) are found upon various parts of the cuti- cle : in some plants only on that of the under side of leaves, in others on the upper also ; in floating leaves upon the latter only. When leaves are so turned that their margins are directed towards the earth and the heavens, the two faces are then alike in appearance, and are both equally furnished with stomata. In succulent leaves they are said to be either alto- o-ether absent or very rare ; but this is a statement that requires confirmation. According to the observations of M. De Can- dolle (Organogr. p. 272.), they are, in the Orange and the Mesembryanthemum, as ten in the former to one in the latter. The parenchyma is, if casually examined, or even if viewed in slices of too great thickness, apparently composed of heaps of little green cells, arranged with little order or regularity ; but, if very thin slices are taken and viewed with a high mag- nifying power, it will be seen that nothing can be more perfect than the plan upon which the whole structure is con- trived, and that, instead of disorder, the most wise order pervades the whole. Upon this subject I extract the words of M. Adolphe Brongniart : — " There exists beneath the upper cuticle two or three layers of oblong blunt vesicles, placed perpendicular to the surface of the leaf, and generally much less in diameter than the cells of the cuticle; so that they are easily seen through it. These vesicles, which appear specially destined to give solidity to the parenchyma of the leaf, have no other intervals than the little spaces that result from the contact of this sort of cylinder: nevertheless, in plants that have stomata on the upper surface of their leaves, as is the case in most herbaceous plants, and in such as float on the surface of water, there exist here and there among the vesicles some large spaces, through which the stomata communicate with the interior of the leaf. This parenchyma is entirely different from what is found beneath the cuticle of the lower side. There, instead of con- sisting of regular cylindrical vesicles, it is composed of irre- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 85 gular ones, often having two or three branches, which unite with the limbs of the vesicles next them, and so form a reticu- lated parenchyma; the spaces between whose vesicles are much larger than the vesicles themselves. It is this reticulated tissue, with large spaces in it (to which the name of cavernous or spongy parenchyma might not im- properly be applied), that, in most cases, occupies at least half the thickness of the leaves between the veins. The arrangement of the vesicles is very obvious if the lower cuticle of certain leaves be lifted up with the layer of parenchyma that is applied against it; it may then be seen that these anastomosing vesicles form a net with large meshes, — a sort of grating inside the cuticle. It must not, however, be supposed that this structure, which I have remarked in several ferns, and in a great many dicotyledonous plants, is without ex- ception. In many monocotyledonous and succulent plants we have some remarkable modifications of this structure. Thus, in the Lily, and several plants of the same family, the vesicles of parenchyma that are in contact with the lower cuticle are lengthened out, sinuous, and toothed, as it were, at the sides : these projections join those of the contiguous vesicle ; and a number of cavities is the consequence, which render this sort of parenchyma permeable to air. An analogous arrangement exists in the lower parenchyma of Galega. In the Iris, there is scarcely any space between the oblong and polyedral vesi- cles which form the parenchyma ; but it is remarked, that the subjacent parenchyma is wanting at every point where the cuticle is pierced by a stoma. In such succulent plants as I have examined, the spaces between the cellules of parenchyma are very small ; but, nevertheless, here and there, there are often larger cavities, which either correspond directly with the stomata, or are in communication with them. The same thino- happens in plants with floating leaves, where the stomata placed on the upper surface correspond with the layer of cylindrical and parallel vesicles ; in such case there are, here and there, between these vesicles, empty spaces which almost always correspond to the points where the stomata exist, and which permit the air to penetrate between the vesicles as far as the middle of the parenchyma of the leaf. G 3 86 ORGANOGRAPHY. BOOK I. Thus much M. Brongniart; who adds, that in submersed leaves there is no cuticle, but the whole consists of solid parenchyma alone, in which there are no other cavities than such as are necessary to float the leaves. The veins, being elongations of the medullary sheath, neces- sarily consist of woody fibre and spiral vessels, to which are sometimes added annular ducts. In submersed leaves spiral vessels are often wanting, the veins consisting of nothing but woody fibre. In these veins M. Schultz finds what he calls vessels of the latex, or of the nutritive fluid ; but it is difficult to understand, either from his figures or descriptions, which kind of tissue in particular he means to designate by that name. M. Adolphe Brdngniart says, the latex vessels are the vasa propria; but what are the vasa propria of leaves, in which there is nothing but woody fibre, spiral vessels, and ducts ? Such are the general anatomical characters of leaves; but it must be borne in mind, that, in different species, they undergo a variety of remarkable modifications. These arise either from the addition of parenchyma when leaves become succulent, or from the non-developement of it when they become membranous, or from the total suppression of it, and even of the veins also in great part, as in those which are called ramentaceous, such as the primordial leaves of the genus Pinus. I have dwelt thus much at length upon the structure of the leaf, because it is by far the most important part of a plant, and that of which the functions are the best ascertained. Let us now turn our attention to the modifications of the leaf. It has already been seen that a leaf may consist of two distinct parts ; the petiole, or stalk, and lamina, or leaf itself: both of these demand separate consideration. The lamina, or limbus, as it is called by some, is subject to many diversities of figure and division ; most commonly it forms an approach to oval, being longer than broad. When speaking of the leaf, it is usual to take the opportunity of explaining the terms employed by botanists to distinguish varieties of figure ; but, as those terms are equally applicable to any other part with a similar dilated surface, it has appeared CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 87 to me expedient to include them in Glossology, where they will accordingly be found. That extremity of the lamina which is next the stem is called its base ; the opposite extremity, its apex ; and the line representing its two edges, the margin or circumscription. If the lamina consists of one piece only, the leaf is said to be simple, whatever may be the depth of its divisions : thus, the entire lamina of Box, the serrated lamina of the Apple, the toothed lamina of Coltsfoot, the r uncinate lamina of Taraxacum, the pinnatifid lamina of Hawthorn (which is often divided almost to its very midrib), are all considered to belong to the class of simple leaves. But if the petiole branches out, separating the cellular tissue into more than one distinct portion, each forming a perfect lamina by itself, such a leaf is often said to be compound, whether the divisions be two, as in the conjugate leaf of Zygophyllum, or inde- finite in number, as in the many varieties of pinnated leaves. Nevertheless, a more accurate notion of a compound leaf is found to consist in its divisions being articulated with the petiole, by which it is much better distinguished from the simple leaf than by the number of its divisions. Thus, the pinnated leaf of a Zamia, and the pedate leaf of an Arum, both in this sense belong to the class of simple leaves ; while the solitary lamina of the Orange, the common Berberry, &c. are referable to the class of compound leaves. This distinction is of some importance to the student of natural affinities ; for, while division, of whatever degree it may be, may be expected to occur in different species of the same genus or order (pro- vided there is no articulation), it rarely happens that truly compound leaves, — that is to say, such as are articulated with their petiole, — are found in the same natural assemblage with those in which no articulation exists. In speaking of the surface of a leaf it is usual to make use of the word pagina. Thus, the upper surface is called pagina superior ; the lower surface, pagina inferior. The upper surface is more shining and compact than the under, and less generally clothed with hairs; its veins are sunken; while those of the lower surface are usually prominent. The cuticle readily separates from the lower surface, but with difficulty from the g 4 88 ORGANOGRAPHY. BOOK I. upper. There are frequently hairs upon the under surface while the upper is perfectly smooth ; but there is scarcely any instance of the upper surface being hairy while the lower is smooth. The ramifications of the petiole among the cellular tissue of the leaf are called veins, and the manner of their distribution is termed venation. This influences in a great degree the figure and general appearance of the foliage, and requires a more careful consideration than it generally receives in ele- mentary works. The vein which forms a continuation of the petiole and the axis of the leaf is called the midrib or costa : from this all the rest diverge, either from its sides or base. If other veins similar to the midrib pass from the base to the apex of a leaf, such veins have been named nerves; and a leaf with such an arrangement of its veins has been called a nerved leaf. If the veins diverge from the midrib towards the margin, ramifying as they proceed, such a leaf has been called a venous or reticu- lated leaf. This is the sense in which these terms were used by Linnaeus ; but Link and some others depart from so strict an application of them, calling all the veins of a plant nerves, whatever may be their origin or direction. Till within a few years the distribution of veins in the leaf had not received much attention; the terms just mentioned had been contrived to express certain of the most striking forms of venation ; but the application of these was far from being sufficiently precise. Many improvements have been pro- posed by modern botanists ; it however appears to me that the whole nomenclature of venation is essentially defective, and requires complete revision. My ideas upon this subject have been already laid before the public in the Botanical Register for Sept. 1826, page 1004.; and, as I am not aware that any objection to them has yet been taken, I shall repeat them here, in a form better adapted to an elementary work than that under which they first appeared. The objections that I take to the present modes of distin- guishing veins are these: — 1st, That the veins are very im- properly, as I think, called nerves, either in all cases, as by Link, which is bad, or in certain cases only, when they have CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 89 a particular size or direction, as by Linnaeus and his followers, which is worse. Nothing is more destructive of accurate ideas in natural history than giving names well understood in one kingdom of nature to organs in another kingdom of an entirely different kind, unless it is the, perhaps, more repre- hensible practice of giving two names conveying totally differ- ent ideas to the same organ in the same kingdom of nature. Thus, when the veins of a plant are termed nerves, it is neces- sarily understood that they exercise functions of a similar nature to those of the nerves of animals : if otherwise, why are they so called ? But they exercise no such functions, being, beyond all doubt, mere channels for the transmission of fluid. Again, if one portion of the skeleton of a leaf is called a vein, and another portion a nerve, this apparently precise mode of speaking leads yet more strongly to the belief (especially when such a distinction is seen admitted into works which are said to be of the highest authority in science), that the structure and function of those two parts are as widely different as the structure and function of a vein and a nerve in the animal economy ; else why should such nice caution be taken to dis- tinguish them ? But it must be confessed that there is no difference whatever, except in size, between the veins and nerves of a leaf. Let us, then, abandon a term which is one of those relics of a barbarous age, which it is the duty of modern science to expel. My second objection is caused by the vague manner in which the veins of leaves are at present described ; whence it happens that no precise idea can be attached to the different terms that have been contrived to designate particular forms of venation. A third objection is this, — that, while slight modifications in the arrangement of the veins have received distinctive names, others of much greater importance, and of a more decided character, have received no distinctive appellation whatever. For these reasons, the practical weight of which I have long experienced, it has occurred to me that the follow- ing changes in the language used in speaking of venation will be found better, at least, than that for which they are sub- stituted, if they are not entirely what could be desired. It has been usual to call that bundle of vessels only which passes directly from the base to the apex of a leaf the costa, or 90 ORGANOGRAPHY. BOOK I. midrib. This term I would extend to all main veins which proceed directly from the base to the apex, or to the points of the lobes. There is no difference in size in these costae ; and in lobed leaves, which may be understood as simple leaves, approaching composition, each costa has its own particular set of veins. The costa (jig. 52, 7.) sends forth, alternately right and left along its whole length, ramifications of less dimensions than itself, but more nearly approaching it than any other veins: 52 53 ^7 these I would call vencs primaries (jig- 52, 3.). They diverge from the costa at various angles, and pass to the margin of the leaf, curving towards the apex in their course, and finally, at some distance within the margin, forming what is called an anastomosis, or junction, with the back of the vena primaria, which lies next them. That part of the vena primaria which is between the anastomoses thus described, having a curved direction, may be called the vena arcuata. Between this latter and the margin, other veins, proceeding from the venae arcuatae, with the same curved direction, and of the same magnitude, occasionally intervene : they may be distinguished by the name of venae externa ( jig. 52, 1 .). The margin itself and these last are connected by a fine net-work of minute veins, which I would distinguish by the name of venules marginales. From the costa are generally produced, at right angles with it, and alternate with the vena? primariae, smaller veins ; which may be considered imperfect venae primariae, and may not im- properly be named vence costales (jig. 52, 5.). The venae primariae are themselves connected by fine veins, which anas- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 91 tomose in the area between them. These veins, when they immediately leave the venae primariae, I call venules pro- price {Jig. 52, 4.); and where they anastomose, venules com- munes. The area of parenchyma, lying between two or more veins or veinlets, I name with the old botanists intervenium. These distinctions may to some appear over-refined ; but I am convinced that no one can accurately describe a leaf without the use of them, or of equivalent terms yet to be invented. Upon these principles leaves may be conveniently divided into the following kinds : — 1. Veinlcss (avenium), when no veins at all are formed, except a slight approach to a costa, as in Mosses, Fuci, &c. Leaves of this description exist only in the lowest tribes of foliaceous plants, and must not be confounded with the fleshy or thickened leaves common among the higher orders of vegetation, in which the veins are by no means absent, but only concealed within the substance of the parenchyma. (See No. 10.) Of this M. De Candolle has two forms, — first, his folia nullinervia, in which there is not even a trace of a costa, as in Ulva ,• and second, his folia fahinervia, in which a trace of a costa is perceptible. These terms appear to me unneces- sary ; but, if they be employed, the termination nervia must be changed to venia. 2. Equal-veined {ceqnalivenium\ when the costa is perfectly formed, and the veins are all of equal size, as in Ferns. This kind of leaf has not been before distinguished : it may be considered intermediate between those without veins and those in which vena3 primariae are first apparent. The veins are equal in power to the venulae propriae of leaves of a higher class. 3. Straight-veined (rectivenium). In this the veins consist only of venae primariae, generally very much attenuated, and arising from towards the base of the costa, with which they lie nearly parallel : they are connected by venulae propriae ; but there are no venulae communes. The leaves of Grasses and of Palms and Orchideous plants are of this nature. This form has been called by Link paralleli and convergenti-nervo- sum, according to the degree of parallelism of the venae pri- mariae ; and to these two he has added what he calls venuloso- 92 ORGANOGRAPHY. BOOK I. nervosum, when the venae primariae are connected by venulae propriae: but as this is always so, although it is not in all cases equally apparent, the term is superfluous. Ach. Richard calls this form later inervium, and De Candolle rectinervium ; from which I do not find it advisable to distinguish his rupti- nervium, which indicates the straight-veined leaf, when the veins are thickened and indurated, as in the Palm tribe. 4. Curve-veined (curvivenium). This is a particular modifi- cation of the last form, in which the venae primariae are also parallel, simple, and connected by unbranched venulae pro- priae; do not pass from near the base to the apex of the leaf, but diverge from the costa along its whole length, and lose themselves in the margin. This is the folium hinoideum and venuloso-hinoideum of Link, the f penninervium of A.Rich- ard, and they, curvinervium of De Candolle. It is common in Scitamineae. It is not improbable that both this and the last ought to be regarded as peculiar modifications of petiole (a kind of phyllodia), rather than as true leaves analogous to those next to be described. 5. Netted (reticulatum). Here the whole of the veins that constitute a completely developed leaf are present, arranged as I have above described them, there being no peculiar com- bination of any class of veins. This is the common form of the leaves of Dicotyledones, as of the Lilac, the Rose, &c. It is the folium venosum of Linnaeus, the^ indirecte vcnosum of Link, they! mixtinervium of A. Richard, and the^ retinervium of De Candolle. If the venae externae and venulae margi- nales are conspicuous, Link calls this form combinate venosum,- but if they are indistinct, he calls it evanescente venosum. 6. Ribbed (costaticm). In this three or more costae proceed from the base to the apex of the leaf, and are connected by branching venae primariae of the form and magnitude of venulae propriae, as in Melastoma. This must not be con- founded with the straight -veined leaf, from which it may in all cases of doubt be distinguished by the ramified veins that connect the costae. This is a very material difference, which has never been properly explained. Linnaeus and his followers confound the two forms ; but modern writers separate them : although it must be confessed that it is difficult to discover CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 93 their distinctions from the characters hitherto assigned to them. Link calls these leavesj^ ncrvata, A. Richard f. basi- nervia, and De Candolle f. triplinervia and f. quintupli- nervia. If a ribbed leaf has three costae springing from the base, it is said to be three-ribbed (tri-costatum, trinerve of authors) ; if five, Jive-ribbed^ and so on. But if the ribs do not proceed exactly from the base, but from a little above it, the leaf is then said to be triply-ribbed (triplicostatiwi), as in the Helianthus. 7. Falsely ribbed (pseudocostattim), is when the venae arcuatae and venae externae, both or either, in a reticulated leaf, become confluent into a line parallel with the margin, as in all Myr- taceae. This has not been before distinguished. 8. Radiating (radiatum), when several costae radiate from the base of a reticulated leaf, to its circumference, as in lobed leaves. This and the following form the f. directe venosum of Link : it is the f. digit inervium of A. Richard. Hither I refer without distinguishing them the f. pedali- nervia, palminervia, and peltinervia of M. De Candolle ; the differences of which do not arise out of any peculiarity in the venation, but from the particular form of the leaves themselves. 9. Feather-veined (penniveniam), when the venae primariae of a reticulated leaf pass in a right line from the costa to the margin, as in Castanea. This has the same relation to the radiating leaf that the curve-veined bears to the straio-ht- veined ; it is the folium pennivenium of M. De Candolle. 10. Hidden-veined (introvenium). To this I refer all leaves the veins of which are hidden from view by the parenchyma being in excess, as in the Hcn/a, and many others. Such a leaf is often inaccurately called veinless. M. De Candolle calls a leaf of this nature, in which the veins are dispersed through a large mass of parenchyma, as in Mesembryanthe- raum, vaginervium. It is often necessary to explain the direction that the venae primariae take when they diverge from the costa: this may be denoted by measuring the angle which is formed by the costa and the diverging vein, and can either be stated in distinct words, or by applying the following terms thus : — if 94- ORGANOGRAPHY. BOOK I. the angle formed by the divergence is between 10° and 20°, the vein may be said to be nearly parallel (subparallela) ; if between 20° and 40°, diverging ; between 40° and 60°, spread- ing,- between 60° and 80°, divaricating; between 80° and 90°, right-angled; between 90° and 120°, oblique; beyond 120°, reflexed (retrqflexa). The petiole {Jig- 55 a — b.) is the part which connects the la- mina with the stem, of which it was considered by Linnaeus as a part. It consists of one or more bundles of fibrovascular tissue surrounded by cellular tissue. Its figure is generally half cy- lindrical, frequently channelled on the surface presented to the heavens ; but in most mo- nocotyledonous plants it is perfectly cylindrical. If the petiole is entirely absent, which is often the case, the leaf is then said to be sessile. Generally the petiole is simple, and con- tinuous with the axis of the leaf; sometimes it is divided into several parts, each bearing a separate leaf or leaflet (Jbliolus) : in such case it is by some said to be compound: each of the stalks of the leaflets being called petiolules (ramastra, Jungius). In all simple leaves the petiole is continuous with the axis of the lamina, from which it never separates ; in all truly com- pound leaves the petiole is articulated with each petiolule; so that when the leaf perishes, it separates into as many portions as there are leaflets, as in the Sensitive plant : hence, when- ever an apparently simple leaf is found to be articulated with its petiole, as in the Orange, such a leaf is not to be considered a simple leaf, but the terminal leaflet of a pinnated leaf, of which the lateral leaflets are not developed. This is a most important difference, and must be borne constantly in mind by all persons who are engaged in the investigation of natural affinities. It is an occult sign which must never be neglected. At the base of the petiole, where it joins the stem, and upon its lower surface, the cellular tissue increases in quan- tity, and produces a protuberance or gibbosity, which Ruellius, CHAF. II. COMPOUND ORGANS IN FLOWERING PLANTS. 95 and after him Link, called the pulvinus, and M. De Candolle coussinet {%fig. 55, a.). At the opposite extremity of the petiole, where it is connected with the lamina, a similar swelling is often remarkable, as in Sterculia, Mimosa sensitiva, and others: this is called the struma, or, by the French, bourrelet (>y%. 55, b.). Occasionally the petiole embraces the branch from which it springs, and in such case is said to be sheathing ,- and is even called a sheath or vagina, as in grasses {Jig- 54>, a.). When the lower part only of the petiole is sheathing, as in Umbelliferse, that part is sometimes called the jpericladium. In grasses there is a peculiar membranous process at the top of the vagina, between it and the lamina, which has received the name of ligula {Jig- 54, b.) (languette, Fr. ; collare, Rich.) ; the nature of this process has not yet been determined. In the Asparagus, the petiole has the form of a small sheath, is destitute of lamina, and surrounds the base of certain small branches having the appearance of leaves : such a petiole has been named hypophijllium by Link. In Trapa natans, Pontedera crassipes, and other plants, the petiole is excessively dilated by air, and acts as a bladder to float the leaves : except in this state of dilatation, it differs in no wise from common petioles : it has, nevertheless, received the name of vesicula from M. De Candolle, who considers it the same as the bladdery expansions of Fuci. The petiole is generally straight : occasionally it becomes rigid and twisted, so that the plant can climb by it. It has been said that the figure of the petiole usually ap- proaches more or less closely to the cylindrical : this, however, is not always the case. In many plants, especially of an herba- ceous habit, it is very thin, with foliaceous margins ; it is then called winged. There are, moreover, certain leafless plants, as the greater number of species of Acacia, in which the petiole becomes so much developed as to assume the appear- ance of a leaf, all the functions of which it performs. Petioles of this nature have received the name of Phyllodia [Jig. 56.). They may always be distinguished from true leaves by the fol- lowing characters : — 1. If observed when the plant is very young, they will be found to bear leaflets. 2. Both their sur- faces are alike. 3. They very generally present their margins 96 ORGANOGRAPHY. BOOK I. to the earth and heavens, — not their surfaces. 4. They are always straight-veined ; and, as they only occur among dico- tyledonous plants which have reticulated leaves, this peculiarity alone will characterise them. But, besides the curious transformation undergone by the petiole when it becomes a phyllodium, there are several others still more remarkable : among these the first to be noticed is the tendril ( Vrille, Fr.; Cirrhus, Linn.; Capreolus and Clavicula of the old botanists). It is one of the contrivances employed by nature to enable plants to support themselves upon others that are stronger than themselves. It was included by Lin- naeus among what he called fulcra ,- and has generally, even by very recent writers, been spoken of as a peculiar organ. But, as it is manifestly in most cases a particular form of the petiole, I see no reason for regarding it in any other light. It may, indeed, be a modification of the inflorescence, as in the Vine ; but this, I conceive, is an exception, showing, not that the cirrhus is not a modification of the petiole, but that any part may become cirrhose. In some cases, the petiole of a compound leaf is elongated, branched, and endowed with the power of twisting round any small body that is near it, as in the Pea : it then becomes what is called a cirrhus petiolaris. At other times, it branches off on each side at its base below the lamina into a twisting ramifica- tion, as in Smilax horrida ; when it is called a cirrhus peduncxi- laris. At other times it passes, in the form of midrib, beyond the apex of a simple leaf, twisting and carrying with it a portion of the parenchyma, as in Gloriosa superba ; when it is said to be a cirrhus foliar is. M. De Candolle refers to tendrils the acu- minate, or rather caudate, divisions of the corolla of Strophan- thus, under the name of cirrhus corollaris ,• but these do not appear to me to possess any of the requisites of a tendril. As another modification of the petiole, I am disposed to consider, with Link (Elem. 202.), the singular form of leaf in Sarracenia and Nepenthes, which has been called Ascidium or Vasculum {outre De Candolle). This consists of a fistular green body, occupying the place and performing the functions of a leaf, and closed at its extremity by a lid termed the operculum. To me it appears that the ascidium itself, or fistular part, is CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 97 57 the petiole, and the operculum the lamina of a leaf in an extraordinary state of transformation. Look, for example, at Dionsea muscipula; in this plant the leaf consists of a broad winged petiole, articulated with a collapsing lamina, the mar- gins of which are pectinate and inflexed. Only suppose the broad winged petiole to collapse also, and that its margins» when they meet, as they would in consequence of collapsion, cohere; a fistular body would then be formed, just like the ascidium of Sarracenia ; and there would be no difficulty in identifying the acknowledged lamina of Dionaea with the operculum of Sarracenia also. From Sarracenia the transition to Nepenthes would perhaps not be considered improbable. 58 98 ORGANOGRAPHY. BOOK I. The student must not, however, suppose that all pitchers are petioles, because those of Nepenthes and Sarracenia are so. Those of the curious Dischidia Rafflesiana (Jig. 58.), figured by Dr. Wallich in his Plantce Asiatics Rariores, are leaves, the margins of which are united. The pitchers of Marcgraavia and Norantea (fg. 59.) are bracteae in the same state. Spines of the leaves are formed either by an elongation of the woody tissue of the veins, or by a contraction of the paren- chyma of the leaves : in the former case they project beyond the surface or margin of the leaf, as in the Holly (Ilex aquifolium) : in the latter case they are the veins them- selves become indurated, as in the palmated spines of Berberis vulgaris. The spiny pe- tiole of many Leguminous plants is of the same nature as the latter. So strong is the tendency in some plants to assume a spiny state, that in a species of Prosopis from Chili, of which I have a living specimen now before me, half the leaflets of its bi- pinnate leaves have the upper half converted into spines. 2. Of Stipulae. f CHAP. II. COiMPOUND ORGANS IN FLOWERING PLANTS. 99 At the base of the petiole, on each side, is frequently seated a small appendage, most commonly of a texture less firm than the petiole, and having a subulate termination. These two appendages are called stipulce. They either adhere to the base of the petiole or are separate ; — they either endure as long as the leaf, or fall off before it ; — they are membranous, leathery, or spiny ; — finally, they are entire or lacineated. By Link they have been called Paraphyllia ,• an unnecessary term. When they are membranous, and surround the stem like a vagina, cohering by their anterior margins, as in Poly- gonum ( Jig. 60.), they have been termed ocJirea by Willdenow. Of this the fibrous sheath at the base of the leaves of Palms, called reticulum by some, may possibly be a modification. In pinnated leaves there are often two stipulae at the base of each leaflet as well as at the base of the common petiole : stipulae, under such circumstances, are called stipcllcc. The exact analogy of stipulae is not well made out. M. De Candolle seems, from some expressions in his Organographies to suspect their analogy with leaves ; while, in other places in the same work, it may be collected that he rather con- siders them special organs. I am clearly of opinion that, notwithstanding the difference in their appearance, they are really accessory leaves : first, because occasionally they are transformed into leaves, as in Rosa bracteata, in which I have seen them converted into pinnated leaves ; secondly, because they often are undistinguishable from leaves, of which they obviously perform all the functions, as in Lathyrus, Lotus, and many other Leguminosae : and, finally, because there are cases in which buds develope in their axilla, as in Salix ; a property peculiar to leaves and their modifications. M. De Candolle, in suggesting, after Seringe, that the tendrils of Cu- curbitaceae are modified stipulae, assigns the latter a tendency to a transformation exclusively confined either to the midrib of a leaf, or to a branch ; and they cannot be the latter. It is sometimes difficult to distinguish from true stipulae, certain membranous expansions, or ciliae, or glandular append- ages of the margin of the base of the petiole, such as are found in Ranunculaceae, Apocyneae, Urn bell iferae, and many other plants. In these cases the real nature of the parts is h 2 100 ORGANOGRAPHY. BOOK I only to be collected from analogy, and a comparison of them with the same part differently modified in neighbouring species. M. De Candolle remarks, that no Monocotyledonous plants have stipulae; but they certainly exist, at least in Fluviales and Aroideae. The ligula of grasses, a membranous appendage at the apex of their sheathing petiole, which some have con- sidered stipulae, should rather be understood as a membranous expansion analogous to the corona of some Caryophylleae, such as Silene. It has been already noted, that when they surround the stem of a plant they become an ochrea ,• in this case their anterior and posterior margins are united by cohesion ; a property that they possess in common with all modifications of leaves, and of which different instances may be pointed out in Magnoliaceae, where the back margins only cohere, in cer- tain Cinchonaceae, in which the anterior margins of the stipulae of opposite leaves are united, and in a multitude of other plants, Of Bracteae. 65 All the parts that have hitherto been subjects of enquiry are called orgems of vegetation ,• their duty being exclusively to perform the nutritive parts of the vegetable economy. Those which are about to be mentioned are called organs of fructification ■, their office being to reproduce the species by a process in some respects analogous to that which takes place CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 101 in the animal kingdom. The latter are, however, all modifi- cations of the former, as will hereafter be seen : and as the subject of this division is in itself a kind of proof; bracteae not being exactly organs either of vegetation or reproduction, but between the two. Botanists call Bractca either the leaf from the axilla of which a flower is developed, such as we find in Veronica agrestis ; or else all those leaves that are found upon the inflo- rescence, and are situated between the true leaves and the calyx. There are, in reality, no exact limits between bracteae and common leaves ; but in general the former may be known by their situation immediately below the calyx, by their smaller size, din°ex*ence of outline, colour, and other marks. They are generally entire, however much the leaves may be divided; frequently scariose, either wholly or in part; often deciduous before the flowers expand ; but rarely very much dilated, as in Origanum, Dictamnus, and a few other plants. It is often more difficult to distinguish bracteae from the sepals of a polyphyllous calyx than even from the leaves of the stem. In fact, there is in many cases no other mode than ascertaining the usual number of sepals in other plants of the same natural order, and considering every leaf-like appendage on the outside of the usual number of sepals as bracteae. In Camellia, for example, if it were not known that the normal number of sepals of kindred genera is five, it would be im- possible to determine the number of its sepals. When the bracteae are very small, they are called bracteola ; or if they are of different sizes upon the same inflorescence, the smallest receive that name. It rarely occurs that an inflorescence is destitute of bracteae. In Cruciferae this is a frequent cha- racter, and is observed by Link to indicate an extremely irregular structure. When bracteae do not immediately sup* port a flower or its stalk, they are called empty {vacua). As a general rule, it is to be understood, that whatever inter- venes between the true leaves and the calyx, whatever be their form, coloui', size, or other peculiarity, comes within the meaning of the term. Under particular circumstances bracteae have received the following peculiar names : — h 3 102 ORGANOGRAPHY. BOOK I. When they are empty, and terminate the inflorescence, they form a coma, as in Salvia Horminum, In this case they are generally enlarged and coloured. If they are verticillate, and surround several flowers, they constitute an involucrum. In Umbelliferous plants, the bracteae which surround the general umbel are called an universal involucrum ; and those which surround the umbellules a partial involucrum, or involucellum. In Compositae, the involucrum often consists of several rows of imbricated bracteae, and has received a variety of names, for none of which does there appear to be the least occasion. Linnaeus called it calyx communis, Necker perigynavdra communis, Richard periphor- anthium, Cassini periclinium. There is often found at the base of the involucrum of Compositae an exterior rank of bracteae, which Linnaeus called calyculus ,• and such involucra as were so circumstanced calyx calyculatus. M. Cassini re- stricts the term involucrum to this; but it seems most con- venient to call these exterior bractese bracteolce, and to say that an involucrum in which they are present is basi bracteo- latus, bracteolate at the base. Another and very remarkable form of the involucrum is the cupula {Jig. 66.). It consists of bracteae not developed till after flowering, when they cohere by their bases, and form a kind of cup. In the Oak the cupula is woody, entire, and scaly, with indurated bracteae: in Fagus it forms a sort of coriaceous valvular spurious pericarpium : in Corylus {Jig. 64.) it is folia- ceous and lacerated : in Taxus it is fleshy and entire, with no appearance of bracteae. The name squama or scale is usually applied to the bracteae of the amentum ; it is also occasionally used to indicate any kind of bracteae which has a scaly appearance. The bracteae which are stationed upon the receptacle of Compositae, between the florets, have generally a membranous texture and no colour, and are called palece, Englished by some botanists chaff of the receptacle. The French call this sort of bracteae paillette, Cassini squamelles {Jig. 63.). In Palms and Aroideae there are seated, at the base of the spadix, large, coloured bracteae, in which the spadix is in aestivation wholly enwrapped, and which may perhaps perform CHAP. U. COMPOUND ORGANS IN FLOWERING PLANTS. 103 in those plants the office of corolla. This is called the spat/ia {Jig. 84.). Link considers it a modification of the petiole ! (Elemeiita, p. 253.) 67 The most remarkable arrangement of bracteae takes place in Grasses, in which they occupy the place of calyx and corolla, and have received a great variety of names from different systematic writers. In order to explain distinctly the application of these terms, I must describe with some minuteness the structure of a locusta or spicula, as the partial inflorescence of Grasses is denominated. Take, for example, any common Bromus ; each locusta will be seen to have at its base two opposite empty bracteae (Jig. 67, b.), one of which is attached to the rachis a little above the base of the other: these are the gluma of Linnaeus and most botanists, the gluma exterior or calycinalis of some writers, the tegmen of Palisot de Beauvois, the lepicena of Richard, the ccetonium of Trinius, and, finally, the jperistachyum of Panzer. Above the gluma are several florets sitting in denticulations of the rachis (Jig. 67, c): each of these consists of one bractea, with the midrib quitting the lamina a little below the apex, and elongated into a bristle called the wwn, beard, or arista, and of another bractea facing the first, with its back to the rachis, bifid at the apex, with no dorsal vein, but with its edges inflexed, and a rib on each side at the line of inflexion (Jig. 67, a.). These bracteae are the corolla of Linnaeus, the calyx of Jussieu, the pierianthium of Mr. Brown, the gluma interior or corollina and perigonium of some, the stragulum of Palisot de Beauvois, the gluma of h 4 104 ORGANOGRAPHY. BOOK I. Richard, the bale or Glumella of De Candolle and Desvaux, the palecc of others. When the arista proceeds from the very apex of the bracteae, and not from below it, it is denominated in the writings of Palisot a seta. Within the last-mentioned bracteae, and opposite to them, are situated two extremely minute colourless fleshy scales {Jig. 67, e.), which are some- times connate : these are named corolla by Micheli and Dumortier, nectarium by Linnaeus, squamula by Jussieu and Brown, glumella by Richard, glumellula by Desvaux and De Candolle, lodicula by Palisot de Beauvois. Amidst these conflicting terms it is not easy to determine which to adopt. I recommend the exterior empty bracteae to be called glumce-, those immediately surrounding the fertilising organs palece ; and the minute hypogynous ones, scales or squamidce. The pieces of which these three classes of bracteae are composed are called valves or valvulce by the greater part of botanists ; but as that term has been thought not to convey an accurate idea of their nature, Desvaux has proposed to sub- stitute that of spathella, which is adopted by M. De Candolle. Palisot proposed to restrict the term gluma to the pieces of the gluma, and to call the pieces of the perianthium palcce. Richard called the pieces of both gluma and perianthium palece, and the squamulae paleolce. It seems to me most con- venient to use the term valvula, because it is more familiar to botanists than any other, and because I do not see the force of the objection which is taken to it. In the genus Carex two bracteae (j%. 67, e, //..) become con- fluent at the edges, and enclose the pistillum, leaving a passage for the stigmata at their apex. They thus form a single urceo- late body named urceolus or perigynium. M. De Candolle justly observes, in his Theorie, that some botanists call this nec- tarium, although it does not produce honey ; others capsula, although it has nothing to do with the fruit ; but he does not seem to me more correct than those he criticises in arranging the urceolus among his miscellaneous appendages of the floral organs, which are " ni organes genitaux ni tegumens." I believe I was the first who explained the true nature of the urceolus, in my translation of Richards's Analyse du Fruity printed in 1819. (p. 13.) CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 105 At the base of the ovarium of Cyperaceae are often found little filiform appendages, called hypogynous setce by most botanists. These are probably of the nature of the squamulae of Grasses, and have been named perisporwn by some French writers. Bracteae are generally distinct from each other, and imbri- cated or alternate. Nevertheless, there are some striking exceptions to this ; as remarkable instances of which may be cited Althaea and Lavatera among Malvaceae, all Dipsaceae, and some Trifolia, particularly my Tr. cyathiferum {Hooker, Fl. Borcali-Amcr.), in all which the bracteae are accurately verticillate, and their margins confluent, as in a true calyx. 4. Of the Flower. 69 70 71 The Flower is a terminal bud enclosing the organs of reproduction by seed. By the ancients the term flower was restricted to what is now called the corolla; but Linnaeus wisely extended its application to the union of all the organs which contribute to the process of fecundation. The flower, therefore, as now understood, comprehends the calyx, the corolla, the stamens, and the pistillum, of which the two last only are indispensable. The calyx and corolla may be wanting, and a flower will nevertheless exist ; but if neither stamens nor pistillum nor their rudiments are to be found, no assem- blage of leaves, whatever may be their form or colour, or how much soever they may resemble the calyx and corolla, can constitute a flower. 106 ORGANOGRAPHY. BOOK I. The flower, when in the state of a bud, is called the alabas- trus [bouton of the French), a name used by Pliny for the rose-bud. Some writers say alabastrum, forgetting, as it would seem, that that term was used by the Romans for a scent-box, and not for the bud of a flower. Link calls the parts of a flower generally, whether united or connate, ?noria, whence a flower is bi-polymorious (Elem., 243.) ; but I know of no writer who employs these terms, which indeed are quite superfluous. The flowers of an anthodium, which are small, and some- what different in structure from ordinary flowers, are called Jlorets (Jfosculi ; elytriculi of Necker ; Jleuron of the French). The period of opening of a flower is called its anthcsis; the manner in which its parts are arranged with respect to each other before opening is called the cestivation. ^Estivation is the same to a flower-bud as vernation (p. 53.) is to a leaf- bud : the terms expressive of its modifications are to be sought in Glossology. This term aestivation is applied separately to the parts of which a flower may consist ; thus, we speak of the aestivation of the calyx, of the corolla, of the stamens, and of the pistillum ; but never of the aestivation of a flower, col- lectively. 5. Of the Inflorescence. Inflorescence is a term contrived to express generally the arrangement of flowers upon a branch or stem. The part which immediately bears the flowers is called the peduncidus or peduncle, and is to be distinguished from any portion of a branch by not producing perfect leaves; those which are found upon it called bractecc being much reduced in size and figure from what are borne by the rest of the plant. The term peduncle, although it may be understood to apply to all the parts of the inflorescence that bear the flowers, is only made use of practically, to denote the immediate support of a single solitary flower, and is therefore confined to that part of the inflorescence which first proceeds from the stem. If it is divided, its principal divisions are called branches ; and its ultimate ramifications, which bear the flowers, are named pedicels. There are also other names which are applied to modifications of the peduncle. CHAP. II. COMPOUND ORGANS IN FLOWERING TLANTS. 107 In plants which are destitute of stem, it often rises above the ground, supporting the. flowers on its apex, as in the Cowslip. Such a peduncle is named a scape (hampe, Fr.). Some botanists distinguish from the scape the pedunculus radi- calism confining the former term to the peduncle which arises from the central bud of the plant, as in the Hyacinth ; and applying the latter to a peduncle proceeding from a lateral bud, as in Plantago media. When a peduncle proceeds in a nearly right line from the base to the apex of the inflorescence, it is called the rachis, or the axis of the inflorescence. This latter term was used by Palisot de Beauvois to express the rachis of Grasses, and is perhaps the better term of the two, especially as the term rachis is applied by Willdenow and others, without much necessity it must be confessed, to the petiole and costa of Ferns. In the locustae of Grasses the rachis has an unusual toothed flexuose appearance, and has received the name of scobina from M. Dumortier. If it is reduced to a mere bristle, as in some of the single-flowered locustae, the same writer then distinguishes it by the name of acicida. I mention these and similar terms, in order that nothing which can even remotely lead to information may be omitted ; but I cannot recommend their adoption. When the part which bears the flowers is repressed in its de- velopement, so that, instead of being elongated into a rachis, it forms a flattened area on which the flowers are arranged, as in Compositae, it becomes what is called a receptacle; or, in the lan- guage of some botanists, the receptacle of the Jloisoer (Jig. 72.). 108 ORGANOGRAPHY. BOOK I. When the receptacle is not fleshy, but is surrounded by an involucrum, it is called the clinanthium (the thalamus of Tournefort), as in Compositae, or, in the language of M. Richard, phoranthium ; the former term is that generally adopted. But if the receptacle is fleshy, and is not enclosed within an involucrum, as in Dorstenia and Ficus (Jig. 73.), it is then called by Link Hypanthodium ; the same writer formerly named it Amphanthium, a term now abandoned. According to the different modes in which the inflorescence is arranged, it has received different names, the right applica- tion of which is of the first importance in descriptive botany. If flowers are sessile along a common axis, as in Plantago, the inflorescence is called a spike (e*pi, Fr.), (Jig. 76.) ; if they are pedicellate, under the same circumstances, they form a raceme (grappe, Fr.), (Jig. 77.) as in the Hyacinth : the raceme and the spike differ, therefore, in nothing, except that the flowers of the latter are sessile, of the former pedicellate. These are the true characters of the raceme and spike, which have been confused and misunderstood in a most extraordinary manner by some French writers. 79, a. CHAP. IJ. COMPOUND ORGANS IN FLOWLIUNG PLANTS. 10.0 80 81 82 83 84 85 86 When the flowers of a spike are destitute of calyx and corolla, the place of which is taken by bracteae, and when with such a formation the whole inflorescence falls off" in a single piece, either after flowering or ripening the fruit, as in Cory 1 us, Salix, &c, such an inflorescence is called an amentum or catkin [chaton, Fr. ; Catuhis, lulus, nucamentum, of old writers), [Jig. 8 1 .) Linnaeus considered the catkin to be an elongated filiform receptacle, analogous to that of Compositae, in which he is followed by Sir James Edward Smith, Link, and others. This opinion arises from a distinction being drawn between the axis of a spike and the receptacle of Compositae ; but, as I have already stated, the latter can be considered in no other light than that of a depressed axis or rachis : so that, when the amentum is said not to be a true axis, but an elongated receptacle, a difference is drawn between words rather than tilings ; for if a receptacle is only a depressed axis, an elongated receptacle is necessarily a return to the common form of the axis. If a spike consists of flowers destitute of calyx and corolla, the place of which is occupied by bractaea, supported by other bracteae which enclose no flowers, and when with such a form- ation the rachis, which is flexuose and toothed, does not fall off with the flowers, as in Grasses, each part of the inflo- rescence so arranged is called a spicula or locust a {ejnllet, Dec; jiaquct, Tournefort). Link is of opinion that the rachis of a spicula, as well as that of the amentum, is a kind of receptacle. 110 ORGANOGRAPHY. BOOK I. When the flowers are closely arranged around a fleshy rachis, which is enclosed in the kind of bracteae called a spatha (see p. 103.), the inflorescence is termed a spadix (spa dice or poi?ifo?i, Fr.), (Jig. 84-.). This is only known to exist in Aroideae and Palms. The raceme has been said to differ from the spike only in its flowers being pedicellate: to this must be added, that the pedicels are all of nearly equal length ; but in many plants, as Alyssum saxatile, the lower pedicels are so long that their flowers are elevated to the same level as that of the upper- most flowers ; a corymbus is then formed (Jig. 86.). This term is frequently used in an adjective sense, to express a similar arrangement of the branches of a plant or of any other kind of inflorescence : thus, in Stevia, the branches are said to be corymbose; in others, the panicle is said to be corymbose; and so on. When corymbose branches are very loose and irregular, they have given rise to the term muscar'mm ; a name formerly used by Tournefort, but not now employed. If the expansion of an apparent corymb is centrifugal, in- stead of centripetal ; that is to say, commences at the centre, and not at the circumference, as inDianthus Carthusianorum, we then have the fasciculus (Jig. 82.); a term which may not incorrectly be understood as synonymous with compound co- rymbus. The modern corymbus must not be confounded with that of Pliny, which was analogous to our capihdum. When the pedicels all proceed from a single point, as in Astrantia, and are of equal length, or corymbose, we have what is called an umbel (Jig. 79.). If each of the pedicels bears a single flower, as in Eryngium, the umbel is said to be simple (fg. 79, a.) ; but if they divide and bear other umbels, as in Heracleum, the umbel is called compound; and then the assemblage of umbels is called the umbella universalis, while each of the secondary umbels, or the umbellules, is named an umbella partialis. The peduncles which support the partial umbels are named radii. The late M. Richard confined the word umbel to the compound umbel, and named the simple umbel sertulum (bouquet); but this was an unnecessary change. Suppose the flowers of a simple umbel to be deprived of their pedicels, and to be seated on a receptacle or enlarged CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. Ill axis, and we have a capitulum or head, named glomus by some, glomerulus by others. If this is flat, and surrounded by an involucrum, the compound flower, as it is inaccurately called by the school of Linnaeus, of Compositae, is produced; which is often named by modern botanists anthodium; a term invented by Ehrhart, and to which there seems to be no objection. It was called cephalanthium by Richard, and calathidium by Mirbel. The flowers or florets borne by the anthodium in its circumference are usually ligulate, and differ- ent from those produced within the circumference. Those in the former station are called Jlorets of the ray, and those in the latter Jlorets of the disk. I have said that the school of Linnaeus inaccurately calls the anthodium a compound flower, from which opinion I should think that few persons would at the present clay dissent, unless they applied the same term to the umbella, the spica, and all other forms of inflorescence, of which the anthodium is palpably a mere modification. Professor Link, however, has in a late work defended the nomenclature of Linnaeus ; urging, that the rays of an anthodium may be considered a sort of corolla, the florets of the disk a representation of the stamens and pistillum ; and that this mode of viewing the subject is much confirmed by the property possessed by the rays of many Compositae, of closing at night, or in cloudy or rainy weather. What this sort of argument may be worth, I profess not to understand ; but it seems to me that we may as well call a branch clothed with leaves a compound leaf, or a flock of sheep a compound sheep, as the cluster of flowers of an anthodium a compound flower. All the forms of inflorescence which have been yet men- tioned are to be considered as reductions of the spike or raceme. Those which are now to be described are decom- positions, more or less irregular, of the raceme. The first of these is the panicle and its varieties. The simple panicle differs from the raceme in bearing branches of flowers where the raceme bears single flowers, as in Poa (fg. 80.); but it often happens that the rachis itself separates into irregular branches, so that it ceases to exist as an axis, as in some Oncidiums. This is called by Willdenow a deliquescent 112 ORGANOGRAPHY. BOOK I. panicle. When the panicle was very loose and diffuse, the older botanists named it a juba; but this is obsolete. If the lower branches of a panicle are shorter than those of the middle, and the panicle itself is very compact, as in Syringa, it then receives the name of thyrsus. Suppose the branches of a deliquescent panicle to become short and corymbose, with a centrifugal expansion indicated by the presence of a solitary flower seated in the axillae of the dichotomous ramifications, and a clear conception is formed of what is called a cyme. This kind of inflorescence is found in Sambucus, Viburnum, and other plants ( and will be spoken of more particularly hereafter. It is sometimes elongated in the form of a little cord, as in the Hazel nut, and many Cruciferae : such a part is called the umbilical card [funiculus umbilicalis, podospermium). The swelling of the ovarium after fertilisation is termed grossijication. The style {tuba of old authors) is that elongation of the ovarium which supports the stigma (Plate V. fig. l.f). It is 140 ORGANOGRAPHY. BOOK I. frequently absent, and then the stigma is sessile : it is not more essential to a pistillum than the petiole to a leaf, or the unguis to a petal, or the filament to a stamen. Anatomically considered, it consists of a column of one or more bundles of vascular tissue, surrounded by cellular tissue ; the former com- municating on the one hand with the stigma, and on the other with the vascular tissue of the ovarium. It is usually taper, often filiform, sometimes very thick, and occasionally angular: rarely thin, flat, and coloured, as in Iris and in Canna. In some plants it is continuous with the ovarium, the one passing insensibly into the other, as in Digitalis; in others it is articulated with the ovarium, and falls off, by a clean scar, immediately after fertilisation has been accom- plished, as in the Scirpus. Its usual point of origin is from the apex of the ovarium ; nevertheless, cases occur, in which it proceeds from the side, as in Alchemilla, or even from the base, as in Labiatae and Boi'agineas. In these cases, how- ever, it is to be understood that the geometrical and organic apices are different, the latter being determined by the origin of the style. For this reason, when the style is said to pro- ceed from the side or base of the ovarium, it would be more correct to say that the ovarium is obliquely inflated or dilated, or gibbous at the base of the style. The surface of the style is commonly smooth ; but in Com- positse, Campanulaceas, and others, it is often densely covered with hairs, called collectors, which seem intended as brushes to clear the pollen out of the cells of the anthers. In Lobelia these hairs are collected in a whorl below the stigma ; in Goodenovise they are united into a cup, in which the stigma is enclosed, and which is called the indusium (Plate V. fig. 13.6.). Many styles which appear to be perfectly simple, as for instance those of the Primrose, the Lamium, the Lily, or the Borage, are in reality composed of several grown together ; as is indicated by the lobes of their stigma, or by the number of cells or divisions of their ovarium. In Malva an example may be seen of a partial union only of the styles, which are distinct upwards, but united below. In speaking of styles in this latter state, botanists are apt to describe them as divided CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 141 in different ways, which is manifestly an inaccurate mode of expression. The stigma is the upper extremity of the style, without a cuticle; in consequence of which it has almost uniformly either a humid or papillose surface. In the first case it is so in consequence of the fluids of the style being allowed to flow up through the intercellular passages of the tissue, there being no cuticle to repress and conceal them ; in the latter case the papillae are really the rounded sides of vesicles of cellular tissue. When perfectly simple, it is usually notched on one side, the notch corresponding with the side from which the placenta arises : see the stigma of Rosa, Prunus, Pyrus, and others. If it belongs to a single carpellum (p. 143.), it is either undivided, or its divisions, if any, are all placed side by side, as in some Euphorbiaceas, Crocus, &c. ; but if it is formed by the union of the stigmas of several carpella, its lobes are either opposite each other, as in Mimulus, or placed in a verticillus, as in Geranium. Such being the case, it is always to be understood that an apparently simple ovarium, to which two or more opposite stigmata belong, is really of a compound nature, some of its parts being abortive, as in Compositae. Nothing is, properly speaking, stigma, except the secreting surface of the style : it very often, however, happens, that the term is carelessly applied to certain portions of the style. For example, in the genus Iris the three petaloid lobed styles in the centre are called stigmata ; while the stigma is in reality confined to a narrow humid space at the back of each style ; in Labiatse, my friend Mr. Bentham has shown that what is called a two-lobed stigma is a two-lobed style, the points only of the lobes of which are stigmatic : and in Lathyrus, and many other Papilionaceous plants, Linnaean botanists call the hairy back of the style the stigma ; while, in fact, the latter is confined to the mere point of the style. Nevertheless, there are certain stigmata in which no denuded or secreting surface can be detected. Of this nature is that of Tupistra, in which the apparent stigma is a fungous mass with a surface of the same nature as that of the style ; 14-2 ORGANOGRAPHY. BOOK I. in such a stigma the mode of fertilisation forms a very inte- resting problem, which botanists have yet to solve. In almost all cases the stigmata are perfectly distinct from all surrounding bodies, and are freely exposed to the influence of the pollen ; but in Asclepiadeae they adhere to the anthers in a solid mass, of which the angles only that are in contact with the cells of the anther are free and sus- ceptible of fertilisation. The centre of a stigma consists of tissue of a peculiar character, which communicates directly with the placenta, and which is called the stigmatic tissue. It is more lax than that which surrounds it, and serves for the conveyance of the fertilising matter of the pollen into the ovula. Such is a general view of the more remarkable peculiarities of the female system. This part, however, bears so important an office in the functions of vegetation, is so valuable as a means of scientific arrangement, and is liable to such a great variety of modifications, that it will be necessary now to con- sider it in another and more philosophical point of view. For we have yet to consider the structure of the compound pis- tillum, and to learn to understand the exact nature of its cells, and dissepiments, and placentae, and the precise relation that these parts bear to each other; and also to prove that the necessary consequence of the laws under which pistilla are constructed is, that they can be subject to only a parti- cular course of modification, within which every form must absolutely, and without exception, fall. This enquiry would, perhaps, be less important if none but structure of a very regular and uniform kind were to exist ; but, considering the numberless anomalies that the pistillum exhibits, it becomes at once one of the most difficult and most essential parts of a student's investigation. In the days of Linnaeus and Gasrtner, and even in those of the celebrated L. C. Richard, nothing whatever was known of this matter, and consequently the writings of those car- pologists are a mere tissue of ingenious misconceptions. Nor did the subject become at all intelligible until the admirable Treatise upon Vegetable Metamorphosis, which had been published by Goethe in 1 790, but which had long been neg- *3 CHAP.IT. compound organs in flowering plants. 143 lected, was again brought into notice, and illustrated by the skilful demonstrations of De Candolle, Turpin, Du Petit Tho uars, and others. 120 121 According to these writers, the pistillum is either the modification of a single leaf, or of one or more whorls of such leaves, which are technically called carpella. Each carpellum has its own ovarium, style, and stigma, and is formed by a folded leaf, the upper surface of which is turned inwards, the lower outwards, and the two margins of which develope one or a greater number of buds, which are in a rudimentary state, and are called the ovules. A very distinct idea of the manner in which this occurs may be obtained from the carpellum of a double cherry, in which the pistillum loses its normal carpellary character, and reverts to the structure of the leaf. In this plant the pis- tillum is a little contracted leaf the sides of which are pressed face to face, the midrib elongated, and its apex discoloured, or a little distended. If we compare this with the pistillum of a single cherry, the mai'gins of the leaf with the ventral suture, the elongated midrib with the style, the discoloured distended apex with the stigma, they will be found to cor- respond exactly. In this case there is an indisputable identity of origin and nature between the ovarium and the lamina of a leaf, — between the little suture that occupies one angle of the car- pellum of a cherry, and the line of union of the two edges of the leaf, — and between the elongated midrib, with its dis- 14-4? ORGANOGRAPHY. BOOK I. tended apex, and the style and stigma. There can be no doubt that the plan of all carpella is the same ; so that the ovarium is the lamina of a leaf, the style an elongated midrib, and the stigma the denuded, secreting, humid apex of the latter. Such being the origin of the carpellum, its two edges will correspond, one to the midrib, the other to the united mar- gins of the leaf. These edges often appear in the carpellum like two sutures, of which that which corresponds to the midrib is called the dorsal, that which corresponds to the united margins is named the ventral suture. It is at some point of the ventral suture that is formed the placenta, which is a copious developement of cellular sub- stance, out of which the ovules or young seeds arise. It, the placenta, originates from both margins of the carpellary leaf, — but as they are generally in a state of cohesion, there appears to be but one placenta, — nevertheless if, as sometimes happens, the margins of the carpellary leaf do not unite, there will be two obvious placenta? to each carpellum. Now, as the stigma is the termination of the dorsal suture, it occupies the same position as that suture with regard to the two placentae ; con- sequently the normal position of the two placentas of a single carpellum will, if they are separate, be right and left of the stigma. This is a fact very important to bear in mind. Pistilla consisting of but one carpellum are simple ; of several, are compound. If the carpella of a compound pis- till um are distinct entirely or in part, they are apocarpous, as in Caltha ; if they are completely united into an undivided body, as in Pyrus, they are syncarpous. That syncarpous pistilla are really made up of a number of united carpella is easily shown, as Goethe has well remarked, in the efenus Nijrella, in which N. orientalis has the carpella partially united, while N. damascena has them completely so. In the latter case, however, the styles are distinct ; they and the stigmata are all consolidated in a single body, when the pistillum acquires its most complete state of complication, as in the Tulip ; which is, however, if carefully examined, nothing but an obvious modification of such a pistillum as that of Nigella damas- cena. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 145 There is this important conclusion that is deducible from the foregoing considerations : viz., that, as the carpella are modified leaves, they are necessarily subject to the same laws of arrangement, and to no others, as leaves developed round a common axis upon one or several planes. For no axiom appears more incontestible in botany, than that all modifications of a given organ are controlled essentially in the same way, and by the same influences, as the organ itself in an unmodified state : and hence every theory of the structure of fruit which is not reducible to that which would be applicable to the structure of whorls of leaves is vicious of necessity. I shall proceed to demonstrate the perfect accordance of the carpellary theory of structure in every point with these principles. The placenta arises from the two margins, either distinct or more usually combined, of a leaf folded inwards. When a leaf is folded inwards, its margins will point towards the stem or axis around which it is developed ; and in a whorl of leaves such inflected margins would all be collected round a common centre; or, if the axis were imaginary, in consequence of the whorl being terminal, would be placed next each other, in a circle of which the back of the leaves would represent the circumference. Therefore the placentae will always be turned towards the axis, or will actually meet there, forming a common centre ; and, which is a very important consequence of this law, if one carpellum only, with its single placenta, be formed in a flower, the true centre of that flower will be indicated by the side of the car- pellum occupied by the placenta. Proofs of this may be found in every blossom : but particularly in such as habitually having but one carpellum occasionally form two, as the Wis- teria sinensis, Alchemilla arvensis, Cerasus acida, &c. ; in these the second carpellum, when added, does not arise by the side of the first, but opposite to it, the face of its placenta being in front of that of the habitual carpellum. A fourth proof of this uniform direction of the placentas towards the axis, is afforded by those pistilla in which a great number of carpella is developed in several rows, as in the Strawberry and the Ranunculus : in all these the placentas will be, with- out exception, found directed towards the axis, and conse- L 0. 146 ORGANOGRAPHY. BOOK I. 122 quently towards the back of every row, except the inner. For ex- ample, in the following diagram \ let O be the axis, bb placenta?, Cjf \^-A \cc the backs of carpella ; the pla- r — i^y ^ f centae, /; b, of the inner row will "~ be next the centre O ; the pla- centa?, bb, of the second row will be next the backs, c c, of the first row ; and so on. If the order of developement of leaves were exactly followed in that of the stamens and carpella, it would happen [that the latter would be invariably alternate with the inner row of sta- 123 mens ; for if a a (Jig. 123.) is the sta- tion of five stamens, bb would be the situations of the carpella: this relative position is therefore considered'the normal one, and is in fact that which usually exists in perfectly regular flowers ; but as all the parts of a flower are subject to deviations, either real or apparent, from what is considered their normal state, in consequence of the non-developement of some parts, or the excessive developement of others, it frequently happens that the carpella either bear no apparent relation to the stamens or are opposite them. In Papilionaceous plants, for example, whei'e only one carpellum is present, it is difficult to say that it bears any exact relation to the stamens, although it is pro- bable that its position is really normal with regard to them ; and so also in Rosaceous plants, with numerous carpella, no exact relation can be proved to exist between the latter and the stamens, unless it may be said to be indicated by those genera, such as Spiraea, in which the carpella are reduced to five ; and, finally, in such plants as Delphinium, in which the carpella are three, while the floral envelopes and male system are divided upon a quinary plan, it is manifest that no alter- nation can exist between the stamens and carpella. As the vessels and petals most commonly consist each of a CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 147 single whorl of parts, so the pistillum is more frequently com- posed of one whorl of carpella than of more. There are, however, certain families in which several whorls are pro- duced one within the other, as in Fragaria, Ranunculus, Magnolia, Annona, and the like. In these cases it mostly happens that the carpella are either entirely separate or nearly so ; but it sometimes happens that syncarpous pistilla are habitually produced with more than one whorl of car- pella, and consequently of cells, as Nicotiana multivalvis, and some varieties of the genus Citrus. In such instances the placentae of the outer series will necessarily be applied to the backs of the inner series, as has been just demonstrated. This mutual relation of the different rows of carpella is sometimes observed when the receptacle from which they arise is either convex or concave : in the former state the outer series will obviously be lowermost, and in the latter uppermost; a circumstance that leads to no intricacy of struc- ture when the carpella are distinct, but which may cause an exceedingly anomalous structure in syncarpous pistilla, espe- cially when accompanied by other unusual modifications of structure. There can be no doubt that the true nature of the composition of the pomegranate is to be explained upon this principle. In order to make these considerations more clear, let figs. 124, 125, and 126. represent — -Jig. 124. a convex receptacle, with distinct carpella ; fig. 125. a concave one, with the same; and^. 126. a concave one, with the carpella con- 124 125 126 solidated. In these, a a are the outer row of carpella, bb the next, and dd the central row. The relative position of these, as the receptacle is convex or concave, will now be apparent. L 2 148 ORGANOGRAPHY. BOOK I. I have stated that the placenta, however simple it may ap- pear to be, is really the result of the union of two united mar- gins of a carpellary leaf: it is, therefore, essentially double; and, accordingly, we find that in polyspermous ovaria the ovula are almost always arranged in two rows, as in the Pea and Bean, the Quince, the Paeony, &c. ; nevertheless there are instances in which the placentae occupy a considerable portion of the wall of the ovarium, and bear the ovula in a great many rows, but in no certain order, as in Nymphaea ; and, on the other hand, some plants have the placentae so little developed, that not more than one ovulum is generated between the two placentae, as in Boragineae, Labiatae, Umbel! i- ferae, Stellatae, Compositae, and many others. There can be no doubt, however, that all the latter cases are mere instances of suppressed structure, in consequence of the general incom- pleteness of developement. When two leaves are developed upon a stem, they are always opposite, and never side by side. As carpella are modified leaves they necessarily obey this law ; and, conse- quently, when a pair of carpella form a bilocular ovarium, the separation of the two cells is directly across the axis of the flower. The partitions that are formed in ovaria, by the united sides of cohering carpella, and which separate the inside into cells, are called dissepiments or septa. It is extremely im- portant to bear in mind, not only that such is really their origin, but that they cannot possibly have any other origin, in order to form an exact idea of the structure of pistilla. Now, as each dissepiment is thus formed of two united sides, it necessarily consists of two plates, which are, in the ovarium state, often so completely united, that their double origin is undiscoverable, but which frequently separate in the ripe Pericarpium. This happens in Rhododendron, Euphorbia, Pentstemon, and a multitude of other plants. The consider- ation of this circumstance leads to certain laws which cannot be subject to exception, but which are of great importance ; the principal of which are these : — 1 . All dissepiments are vertical and never horizontal. — For 15 CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 149 L-J f 12? if a, b mfg. 127. represents the ' side of one carpellum and c, d that of another, the dissepiment a, c, b, d formed by this union will have precisely the same di- rection as that of the carpella, and can never acquire any other ; and the same would be true of the sides f,./and g, k, if they formed themselves into dissepiments by uniting with other carpella: consequently a partition in any cell in the direction of *, k could not be a dissepiment, but would be of a different nature. 2. They are uniformly equal in number to the carpella out of which the pistillum is formed. — Suppose the triangle A, B, C represented a transverse section of an ovarium formed by the union of three carpella o, o, o ; then d, e, f would be the dissepiments, and could not be either more or less. 3. They proceed directly from the pla- centa. — As the placenta is the margin of the carpellary leaf, and as the dissepiment is the side of the carpellary leaf, it is evident that a dissepiment cannot exist apart from the placenta. Hence, when any partition exists in an ovarium which is not connected with the pla- centa, it follows that such a partition is not a dissepiment, however much it may otherwise resemble one. 4. They are alternate with placenta, formed by the cohesion of the margins of the same carpellum, and opposite to placentae, formed by the co- hesion of the contiguous margins of differ- ent carpella. — Let the triangle A, B, C represent a transverse section of a three- celled ovarium of which d, e, f are the dissepiments: the dissepiments d and e will alternate with the placentae m, g, both belonging to the carpellum A; but l 3 150 OUGANOGRAPHY. BOOK I. the dissepiment d will be opposite the placentae m, I, formed by the cohesion of the contiguous margins of the carpella A and B. 5. A single carpellum can have no dissepiment whatever. 6. The dissepiment will always alternate with the stigma ; — for the stigma is the extremity of the mid-rib of the carpellary leaf, or of the dorsal suture of the carpellum; and the sides of either of these (which form dissepiments) will be right and left of the stigma, or in the same position with regard to the latter organ as the sides of the lamina of a leaf to its apex. Let the triangle a, b, c represent a 13° transverse section of a three-celled ^ ovarium, of which d, e,f are the dis- sepiments. The stigmata would oc- cupy a position equal to that of the spaces s, s, s, and would consequently be alternate with d, e,f, the dissepi- ments : they could not possibly be placed opposite d, e,f, upon any prin- ciple of structure with which we are acquainted. This law proves, that neither the membrane which separates the two cells of a Cruciferous siliqua, nor the vertical plate that divides the ovarium of Astragalus into two equal portions, are dissepiments ; both are expansions of the placenta, or some other part, in different degrees. Such is the structure of an ovarium in its most common state ; certain deviations from it remain to be explained. We have seen that when carpella become syncarpous, they form a pistillum, the ovarium of which has as many cells and dissepiments as there are carpella employed in its construc- tion. But sometimes the united sides of the carpella do not project so far into the cavity of the ovarium as to meet in the axis, as in the Poppy; and then an ovarium is the result, which, although composed of many carpella, is nevertheless one- celled [fig. 133.) In such case the dissepiments project a short distance only beyond the inner lining, or paries, of the ova- rium, and, bearing on their edges the placenta?, the latter are said to be parietal. In other plants, such as Corydalis, Viola, and Orchis, the carpella are not folded together at all, but are spread open and united by their edges (Jig. 132.) : in that case © CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 151 the placentae do not project at all into the cavity of the ova- rium, but are still more strictly parietal than the last. 131 132 133 Another class of anomalies of a still more remarkable cha- racter, is that in which the dissepiments are obliterated, while the placentae remain a distinct mass in the centre of the ovarium, as in Lychnis ; forming what is called a. free central placenta (Jig. 131.). But, if we examine these plants at a very early period of their formation, long before the flowers expand, the explanation of the anomaly will be obvious. Such plants are, at that time, constructed upon the ordinary plan, with their dissepiments meeting in the centre and forming there a fungous placenta ; but subsequently the shell of the ovarium grows more rapidly than the dissepiments, and breaks away from them; while the excessive growth of the placenta after- wards destroys almost all trace of them : their previous pre- sence is only to be detected by lines upon the shell of the ovarium, or by a separation of the mass of ovula into distinct parcels upon the placenta. All partitions whose position is at variance with the fore- going laws are spurious. Such spurious dissepiments are caused by many circumstances, the chief of which are the following:, — the}' are caused by expansions of the placenta, as in Cruciferae, when they form a partition stretching from one side to the other of the fruit ; or they are mere dilatations of the lining of the pericarpium, as in Cathartocarpus Fistula, in which they are horizontal ; or they are internal expansions of the dorsal or ventral suture, as in Amelanchier, Astragalus, and Thespesia, in which they are distinguishable from their dissepiments by not bearing the placentae, and by being op- posite the stigma, or by projecting beyond the placentae ; or, finally, they are caused by the sides of the ovarium projecting l 4- 152 ORGANOGRAPHY. BOOK I. into the cavity, uniting and forming many supernumerary cells, as in Diplophractum. 11. Of the Receptacle. The part upon which the carpella are seated is the apex of the peduncle, or the summit of the floral branch, of which the carpella are the termination. Usually this part, which is called the receptacle, is flat, or merely a vanishing point; but in other cases it is very much dilated, and then assumes a variety of curious appearances. This receptacle is called torus, or thalamus as w-ell as receptaculum, and, in Greek compounds, has the name of Clinium. In Annonaceae and Magnoliacese it elevates itself from the base of the calyx, and bears the numerous stamens peculiar to these orders : here it is called Gonophore (Gonophorum) by De Candolle. In Caryophylleae the receptacle is elongated, and bears on its summit the petals and stamens : M. De Can- dolle calls this form Anthophore (Anthophorum). When the receptacle bears only the ovarium, and is not a support to either corolla or stamens (Plate V. fig. 1. a.) it is called Car- pophore, or Gipwphore (Carpophorum, Gynophorum) : this may either be a simple rounded stalk to the fruit, and of the same texture with it, as in Capparis, Phaca, and others, — when Richard calls it Basigynium or Podogynium, and Ehrhart llie- caphore, — or it may be succulent and much dilated, so as to resemble the receptacle of a Composita, bearing at the same time many ovaria, as in the Strawberry and Raspberry, when R-ichard calls it Polyphorum : most commonly such a recep- tacle is sufficiently described by the adjective fleshy. In Geranium it is remarkable for being lengthened into a taper- ing cone, to which the styles adhere, in the form of a beak ; and in Nelumbium it is excavated into a number of cavitities, in which the ovaria are half hidden. 12. Of the Ovulum. The ovulum (Plate V. fig. 16. to 26.) is a small, semipellu- cid, pulpy body, borne by the placenta, and gradually chang- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 15S ing into a seed. Its internal structure is exceedingly difficult to determine, either in consequence of its minuteness, or of the extreme delicacy of its parts, which are easily torn and crushed by the dissecting knife. It is doubtless to this cir- cumstance chiefly, that the anatomy of the ovulum was almost unknown to botanists of the last century, and that it has only begun to be understood within ten or twelve years, durino- which it has received ample illustration from several skilful observers. Mr. Brown, indeed, claims to have pointed out its real nature so long ago as 1814; but the brief and incom- plete terms then used by that gentleman, in the midst of a long description of a single species, in the Appendix to Captain Flinders' Voyage, unaccompanied as they were by any explan- atory remarks, prove indeed that he knew something of the subject, but by no means entitle him to the credit of having, at that time, made the world acquainted with it. The late Mr. Thomas Smith seems to deserve the credit of hav- ing first made any general remarks upon the subject : of what extent they exactly were is not known, as his discoveries, in 1818, were communicated, as it would seem, in convers- ation only ; but it is to be collected from Mr. Brown's state- ment that they were of a highly important nature. Since that period the structure of the ovulum has received much atten- tion from Messrs. Brown, in England ; Turpin and Adolphe Brongniart, in France; and Treviranus, in Germany ; by all of whom the subject has been greatly illustrated. It is, how- ever, to the learned M. Mirbel, — who, by collecting the dis- coveries of others, examining their accuracy, snd combining them with numerous admirable observations of his own, has given a full account of the gradual developement and the different modifications of the ovulum — that we are indebted for by far the best description of that important organ. His two papers read before the Academy of Sciences at Paris, in 1828 and 1829, are a perfect model of candour and patient investigation, and form the basis of what is here about to be recorded on the subject. I regret, however that the space which can now be devoted to the explanation of the structure of the ovulum is by no means such as its intricacy and interest demand. 154 ORGANOGRAPHY. BOOK I. As the ovula are the production of the placentae, they necessarily originate in the margins of the carpellary leaf; and hence they have not only been compared to the buds found upon the margins of some true leaves, in a theoretical point of view, but it also has been attempted to be shown that they are analogous to them in structure. Of the truth of the former there can be little doubt; for, to say nothing of such plants as Bryophyllum, which habitually form buds on the margins of the leaves, the case of Malaxis paludosa, first recorded by Professor Henslow, in which the edge of the leaf is frosted by little micrpscopical points, which are neither exactly ovula nor exactly buds, seems a sufficient proof: but that in structure they are analogous to buds is by no means well made out. M. Turpin indeed has attempted, with his usual ingenuity, to demonstrate an analogy between the bracteae of Marcgraavia and the outer integument or primine of the ovulum ; but his statement cannot be regarded, as a sufficient evidence, of the fact. It is, therefore, only safe, at present, to regard the ovulum as analogous to the marginal buds of leaves in nature and position, but not in structure. In almost all cases the ovulum is enclosed within an ova- rium, as would necessarily happen in consequence of the con- volute nature of the carpellary leaves ; but if the convolution is imperfect, as in Reseda, the ovula are partially naked ; and if it does not exist at all, as in Cycadeae and Coniferae, the ovula are then entirely naked ; and, instead of being fertilised by molecules conveyed through the stigma and the style, as in other plants, are exposed to the direct influence of the pollen. This was first noticed by Mr. Brown; and, although since contradicted, is no doubt perfectly true. When the ovula are attached to the placenta by a kind of cord, that cord is called the funiculus (Plate 5. fig. 26, a.), and is a mere prolongation of the placenta. In the beginning, the ovulum is a pulpy excrescence (Plate 5. fig. 16.), appearing to be perfectly homogeneous, with no trace of perforation or of envelopes. But, as it advances in growth, it is gradually (Plate 5. fig. 17 to 21.) enclosed in two sacs or integuments, which are open only at their apex; where, in both these sacs, a passage exists, called CIIAr. II. COMPOUND ORGANS IN FLOWERING TLANTS. 155 the foramen (Plate 5. fig. 21, a.) ; or, in the language of M. Mirbel, Exostome (fig. 25, a.), in the outer integument, and Endostome (fig. 25, &.), in the inner integument. The central part is a fleshy, pointed, pulpy mass, called the nucleus, or micelle (Plate V. fig. 19, 20. a, 22. b, 23. c, 24. d, 25. e, 27. e). The outermost of the sacs (Plate V. fig. 22. c, 23. a, 25. c) is called the primine. It is either merely a cellular coating, or it is traversed by numerous veins or bundles of tubes ; these are sometimes very apparent, as in the Orange tribes ; and M. Mirbel seems disposed to think that they often exist in a rudimentary state when they are not visible. Usually it is nearly as long as the secondine, but sometimes is remark- ably shorter, as in the Euphorbia Lathyris when very young (Plate V. fig. 22.). The outermost but one of the sacs (Plate V. fig. 23. b, 20. b, 25. d.) is called the second 'ine ; it immediately reposes upon the primine, and often contracts an adhesion with it, so that the two integuments become confounded. In order to ascertain its existence, it is, therefore, often necessary to examine the ovulum at a very early period of its growth. It is probable that it always exists ; but Myrica, Alnus, Corylus, Quercus, and Juglans have been named by M. Mirbel as plants in which the secondine is not perceptible (Plate V. fig. 24.). Its point is usually exerted beyond the foramen of the primine. The nucleus (Plate V. fig. 22. b, 18, 19, 20. a, 24. d, 25. e) is a pulpy conical mass, enclosed by the primine and secondine, and often covered up by them ; but frequently protruded beyond the latter, and afterwards, at a subsequent period of its growth, again covered by them. Sometimes its cuticle separates in the form of a third coating of the ovulum, called the tercine. These three parts, the primine, the secondine, and the nucleus, have all an organic connection at some one point of their surface. That point is, in ovula whose parts do not undergo any alteration of direction in the course of their growth, at the base next the placenta ; so that the nucleus is, like a cone, growing from the base of a cup, the base of which is connected with the hiluni through another cup like 1 56 ORGANOGRAPHY. BOOK I. itself (Plate V. fig. 23.). The axis of such an ovulum, which M. Mirbel calls Orthotropous, is rectilinear, as in Myrica, Cistus, Urtica, &c. ; and the foramen is at the end of the ovulum most remote from the hilum. But sometimes, while the base of the nucleus and that of the outer sacs continue contiguous to the hilum, the axis of the ovulum instead of remaining rectilinear is curved down upon itself (Plate V. fig. 26, 27-); so that the foramen, in- stead of being at the extremity of the ovulum most remote from the hilum, is brought almost into contact with it. Ex- amples of this are found in Papilionaceous plants, Caryophyl- leous plants, Mignonette, &c. M. Mirbel, who first distin- guished these, calls them Campulitropous. In both these modifications the base of the ovulum and the base of the nucleus are the same. In a third class the axis of the ovulum remains rectilinear ; but one of the sides grows rapidly, while the opposite side does not grow at all, so that the point of the ovulum is gra- dually pushed round to the base; while the base of the nucleus is removed from the hilum to the opposite extremity (Plate V. fig. 16 — 21.); and when this process is completed the whole of the inside of the ovulum is reversed; so that the apex of the nucleus, and consequently the foramen, corresponds with the base of the ovulum. Such ovula as these M. Mirbel terms Anatropous ,• they are very common : examples may be found in the Almond, the Apple, the Ranunculus, the Cu- cumber, &c. When the base of the nucleus is thus removed from the base of the ovulum, a communication between the two is always maintained by means of a vascular cord, called the raphe (Plate V. fig. 24.*?, 25./). This raphe, which ori- ginates in the placenta, runs up one side of the ovulum, until it reaches the base of the nucleus ; and there it expands into a sort of vascular disk, which is called the chalaza (Plate V. fig. 24./ 25. g.). As the chalaza is uniformly at the base of the nucleus, it will follow that, in Orthotropous and Campu- litropous ovula it is confounded with the hilum ; while it is only distinguished in Anatropous ones, in which alone it is distinctly to be recognised. It has been remarked that the raphe, or vascular extension chap.it. compound organs in flowering plants. 157 of the placenta, always occupies the side next the ventral suture of the ovarium ; and that when, as in Euonymus, it is turned towards the dorsal suture, that circumstance arises from an alteration in the position of the ovulum subsequent to its being fertilised. It has already been stated that the passage through the primine and secondine is called the foramen ; or the exos- tome, when speaking of that of the primine ; and the endos- tome, in speaking of the secondine. Upon these M. Mirbel remarks, — " These two orifices are at first very minute ; but they gradually enlarge ; and when they have arrived at the maximum of dilatation they can attain, they contract and close up. This maximum of dilatation is so considerable in a great number of species, in proportion to the size of the ovulum, that, to give an exact idea of it, I would compare it not to a hole, as those express themselves who have hitherto spoken of the exostome and endostome, but to the mouth of a goblet or of a cup. It may therefore be easily understood that to perceive either the secondine or the nucleus, it is not necessary to have recourse to anatomy. I have often seen, most distinctly, the primine and secondine forming two large cups, one of which encompassed the other without entirely covering it, and the nucleus extending itself in the form of an elongated cone beyond the secondine, to the bottom of which its base was fixed." In practical botany the detection of the foramen is often a matter of great importance ; for it enables an observer to judge from the ovulum of the direction of the radicle of the future embryo : it having been ascertained by many observ- ations that the radicle of the embryo is almost always pointed to the foramen. A partial exception to this law exists, how- ever, in Euphorbiaceae, in many of which M. Mirbel has noticed that, after fertilisation, the axis of the nucleus and the endostome is inclined five or six degrees, without the exos- tome changing its position ; by this circumstance the foramen of the secondine and that of the primine cease to correspond, and the radicle, instead of pointing when formed to the exos- tome, is directed to a point a short distance on one side of it. Besides the two external integuments, M. Mirbel has re- 158 ORGANOGRAPHY. BOOK I. marked the occasional presence of three others peculiar to the nucleus, which he calls the tercine, quartine, and quintine. The former is the external coat of the nucleus, and is very generally, if not universally, present. As I am almost unac- quainted either with it or the two latter, I can add nothing to the following remarks of M. Mirbel upon the subject : — " The quartine and quintine are productions slower to show them- selves than the preceding. The quartine is not very rare, although no one has previously indicated it ; as to the quin- tine, which is the vesicula amnios of Malpighi, the additional membrane of Mr. Brown, and the sac of the embryo of M. Adolphe Brongniart, I am far from thinking that it only exists in a small number of species, as Mr. Brown seems to suppose. If no one has noticed the quartine, it is, no doubt, because it has been confounded with the tercine ; nevertheless these two envelopes differ essentially in their origin and mode of growth. I have only discovered the quartine in ovula of which the tercine is incorporated at an early period with the secondine ; and I think that it is only in such cases that it exists. At its first appearance it forms a cellular plate, which lines all the internal surface of the wall of the cavity of the ovulum ; at a later period it separates from the wall, and only adheres to the summit of the cavity : at this period it is a sac, or rather a perfectly close vesicle. Sometimes it rests finally in this state, as in Statice ; in other cases it fills with cellular tissue, and becomes a pulpy mass ; under this aspect it is seen in Tulipa gesneriana. All this is the reverse of what takes place in the tercine ; for this third envelope always begins by being a mass of cellular tissue, (and at that time it has the name, as we have seen, of nucleus,) and generally finishes by becoming a vesicle. " I have remarked the fifth envelope, or quintine, in many species ; its general characters are such as to prevent its being mistaken. Its complete developement takes place only in a nucleus which remains full of cellular tissue, or in a quartine that has filled with the same. At the centre of the tissue is organised, as in a womb, the first rudiment of the quintine ; it is a sort of delicate intestine, which holds by one end to the summit of the nucleus, and by the other end to the chalaza. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 159 The quintine swells from top to bottom ; it forces back on all sides the tissue that surrounds it, and it often even invades the place occupied by the quartine or the nucleus. A very delicate thread, the suspensor, descends from the summit of the ovulum into the quintine, and bears at its extremity a globule which is the nascent embryo." It is apparently this quintine that Mr. Brown describes, in the ovulum of the Orchis tribe, as a thread consisting of a simple series of short cells, the lowermost joint or cell of which is probably the original state of what afterwards, from enlargement and deposit of granular matter, becomes the opaque speck, or rudiment, of the future embryo. (Observ. on the Organs, fyc. of Orch. and Asclepiad. pp. 18, 19.) " The existence," continues M. Mirbel, " of a cavity in the quartine, or, indeed, the destruction of the internal tissue of the nucleus, at the period when the quintine developes, becomes the cause of some modifications in the manner of existence of this latter integument. The quintine is never seen, in certain Cucurbitacese, adhering to the chalaza : it is nevertheless evident that the adhesion has existed. The quintine, distended at its upper part, and suspended like a lustre from the top of the cavity, still presents at its lower end a portion of a rudimentary intestine become distinct ; the separation occurred very early, in consequence of the tearino- of the tissue of the nucleus. The quintine of Statice is reduced to a sort of cellular placenta, to the lower surface of which the embryo is attached. This abortion of the quintine arises from the quartine havino- a large internal cavity, which prevents the young quintine from placing itself in communication with the chalaza, and taking that developement which it acquires in a multitude of other species." The fluid matter contained within the nucleus is called the liquor amnios, and is supposed to be what nourishes the embryo during its growth. When an ovulum grows erect from the base of the ovarium, it is called erect ; when from a little above the base, ascending . when it hangs from the summit of the cavity, it is jpendidous ; and when from a little below the summit, it is suspended. 160 ORGANOGRAPHY. BOOK I. 134 13. Of the Fruit. 135 136 CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 1G1 163 o 164 165 a 163 b 166 165 b The fruit is the ovarium or pistilhim arrived at maturity; but, although this is the sense in which the term is strictly applied, yet in practice it is extended to whatever is combined with the ovarium when ripe. Thus the pine-apple fruit con- sists of a mass of bracteae, calyces, corollae, and ovaria ; that of the nut, the acorn, and many others, of the superior dry calyx and ovarium ; that of the apple of succulent superior calyx, and corolla, and ovarium ; and that of the strawberry-blite of a succulent inferior calyx and dry ovarium. The fruit being the matured ovarium, it should exhibit upon some part of its surface the traces of a style or stigma ; and this mark will, in many cases, enable the student to dis- tinguish minute fruits from seeds. Many fruits were formerly called naked seeds, such as those of Umbelliferae, Labiatae, and Boragineae, and the grain of corn ; but now, that atten- tion has been paid to the gradual developement of organs, such errors have been corrected. In cases where a trace of the style cannot be discovered, anatomy will generally show whether a minute body is a seed or fruit, by the presence, in the latter case, of two separable and obviously organically distinct coatings to the nucleus of the seed ; but in other cases, when the pericarpium and the integuments of the seeds are combined in a single covering, and when no trace of style remains, as sometimes happens, nothing can be deter- mined as to the exact nature of a given body without follow- ing it back in its growth to its young state. This, however, M 162 ORGANOGRAPHY. BOOK I. may be stated, that naked seeds, properly so called, are not known to exist in more than three or four orders in the whole vegetable kingdom ; viz. in Coniferae and Cycadeae, where the ovula also are naked, and in Peliosanthes Teta and Leontice, in which the ovula, originally enclosed in an ovarium, rupture it at an early period after fertilisation, and subsequently continue naked until they become seeds. Such being the case, it follows that all the laws of structure which exist in the ovarium are equally to be expected in the fruit ; and this fact renders a repetition in this place of the general laws of formation unnecessary. Nevertheless, as, in the course of the advance of the ovarium to maturity, many changes often occur which contribute to conceal the real structure of the fruit, it is in all cases advisable, and in many absolutely necessary, to examine the ovarium, in order to be certain of the exact construction of the fruit itself. These changes are caused by the abortion, non-developement, obli- teration, addition, or union of parts. Thus the three-celled six-ovuled ovarium of the oak and the hazel becomes, by the non-developement of two cells and five ovules, a fruit with one seed ; the three-celled ovarium of the cocoa-nut is converted into a one-celled fruit by the obliteration of two cells and their ovula ; and the two-celled ovarium of some Pedalineae becomes many-celled by a division and elongation of the placentae. In a very early state the ovarium of the Lychnis and of the primrose consists of five cells, each with a placenta having a number of ovula ; by degrees the dissepiments are ruptured and obliterated by the rapid growth of the shell of the ovarium ; and it finally becomes a fruit with only one cell, and a large fungous placenta in the middle. In Cathartocarpus fistula a one-celled ovarium changes into a fruit, having each of its many seeds lodged in a separate cell, in consequence of the formation of numerous horizontal membranes which in- tercept the seeds. A still more extraordinary confusion of parts takes place in the fruit of the pomegranate after the ovarium is fertilised ; and many other cases might be men- tioned. Every fruit consists of two principal parts, the pericardium and the seed, the latter being contained within the former. CHAT. II. COMPOUND OIIGANS IN FLOWERING PLANTS. 163 When the ovarium is inferior, or coheres with the calyx, the latter and the pericarpium are usually so completely united as to be inseparable and undistinguishable : in such cases it is usual to speak of the pericarpium without reference to the calyx, as if no such union had taken place. Botanists call a fruit, the pericarpium of which adheres to the calyx, an infe- rior fruit (fructus injerus) ; and that which does not adhere to the calyx, a superior fruit (fructus superus). But M. Des- vaux has coined other words to express these ideas : a supe- rior fruit he calls autocarpien ,■ an inferior fruit, heterocarpien ; terms wholly unnecessary and unworthy of adoption. Every thing which in a ripe fruit is on the outside of the real integuments of the seed belongs to the pericarpium. It consists of three different parts, the epicarpium, the sarcocar- pium, and the endocarpium ; terms contrived by Richard, and very useful in practice. The epicarpium is the external integument or skin ; the endocarpium, called put amen by Gaertner, the inner coat or shell ; and the sarcocarpium, the intermediate flesh. Thus, in the peach, the separable skin is the epicarpium, the pulpy flesh the sarcocarpium, and the stone the endocarpium or putamen. In the apple and pear the epicarp is formed by the cuticle of the calyx, and the sarcocarpium is confluent with the remainder of the calyx in one fleshy body. The pericarpium is extremely variable in size and texture, varying from the dimension of a single line in length to the magnitude of two feet in diameter : and from the texture of a delicate membrane to the coarse fabric of wood itself, through various cartilaginous, coriaceous, bony, spongy, succulent, or fibrous gradations. The base of the pericarpium is the part where it unites with the peduncle ; its apex is where the style was : hence the organic and apparent apices of the fruit are often very different, especially in such as have the style growing from their sides, as in Rosaceae and Chrysobalaneae, Labiatae and Boragineae. When a fruit has arrived at maturity its pericarpium either continues perfectly closed, when it is indehiscent, as in the hazel-nut, or separates regularly round its axis, either wholly m 2 164 ORGANOGRAPHY. BOOK I. A d or partially, into several pieces : the separation is called dehis- cence, and such pieces valves ; and the axis from which the valves separate in those cases where there is a distinct axis, is called the columella. When the dehiscence takes place through the dissepiments it is said to be septicidal ; when through the back of the cells it is called loeidicidal ; if along the inner edge of a simple fruit it is called sutural ,• if the dissepiments are separated from the valves the dehiscence is named septif vagal. In septicidal dehiscence the dissepiments 167 divide into two plates and form the sides of v/\d each valve, as in Rhododendron, Menziesia, / \ &c. Formerly botanists said that in this sort t— s— a of dehiscence the valves were alternate with the dissepiments, or that the valves had their margins turned inwards. This may be understood from Jig. 167., which represents the relative position of parts in a transverse section of a fruit with septi- cidal dehiscence ; v being the valves, d the dissepiments, and a the axis. In loculicidal dehiscence the dissepiments form the middle of each valve, as in the lilac, or in the diagram 168., where the letters have the same value as above. In this it was formerly said that the dissepiments were opposite the valves. In septifragal dehiscence the dissepiments adhere to the axis and separate from the valves, as in Convolvulus ; or in the dia- gram 169., lettered as before. In sutuval dehiscence there are no disse- piments, the fruit being composed of only A one carpellum, as the pea. Besides these regular forms of valvular dehiscence, there is a mode which obtains in a very few plants, called civcumscissile. This occurs by a transverse circular separation, as in Ana- gallis ; in Jeffersonia it only takes place half round the fruit. Valvular dehiscence, which is by far the most common mode by which pericarpia open, must not be confounded with 168 CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 165 either rujrturing or solubility, — irregular and unusual con- trivances of nature for facilitating the dispersion of seeds. In valvular dehiscence the openings have a certain reference to the cells, as has been already shown ; but neither rup- turing nor solubility bear any distinct relation to the cells. Rupturing consists in a spontaneous contraction of a portion of the pericarpium, by which its texture is broken through and holes formed, as in Antirrhinum and Campanula. Solubi- lity arises from the presence of certain transverse contractions of a one-celled pericarpium, through which it finally separates into several closed portions, as in Ornithopus. For the nature of the placenta and funiculus umbilicalis, see the observations under ovarium. These parts, which are mere modifications of each other, essentially appertain to the pericarpium, in which the former often acquires a spongy dilated substance, occasionally dividing the cells by spurious dissepiments, and often giving to the fruit an appearance much at variance with its true nature. In some seeds, as Euonymus europseus, it becomes exceedingly dilated around each seed, forming an additional envelope, called arillus. The true character of this organ was unknown till it was settled by Richard : before his time the term was applied, not only in its true sense to an enlargement of the placenta, but also to the endocarpium of certain Rubiaceae and Rutaceae, to the testa of Jasminum, Orchideas, and others, and even to the perianthium of Carex. A very remarkable instance of the arillus is to be found in the nutmeg, in which it forms the part called the mace, surrounding the seed. It is never developed until after the impregnation of the ovulum. Having thus explained the structure of the pericarpium, it is in the next place necessary to enquire into the nature of its modifications, which in systematic botany are of considerable importance. It is on the one hand very much to be regretted that the terms employed in this department of the science, which is that of Carpology, have been often used so vaguely as to have no exact meaning; while, on the other hand, they have been so exceedingly multiplied by various writers, that the phraseology of Carpology is a mere chaos. In practice but a small number of terms is actually employed ; but it can- m 3 166 ORGANOGRAPHY. BOOK I. not be doubted that, if it were not for the inconvenience of overburdening the science with terms, it would conduce very much to clearness of description if botanists would agree to make use of some very precise and uniform nomen- clature. What, for instance, can be more embarrassing than to find the term nut applied to the superior plurilocular pericarpium of Verbena, the gland of Corylus, and the achenia of Rosa and Borago ; and that of berry to the fleshy envelope of Taxus, the polyspermous inferior fruit of Ribes, the succulent calyx of Blitum, and several other things ? So much discordance, indeed, exists in the application of terms expressive of the modifications of fruit, that it is quite indispensable to give the definitions of some of the most eminent writers upon the subject in their own words, in order that the meaning attached by those authors to carpological terms, when employed by themselves, may be clearly under- stood. In the phraseology of writers antecedent to Linnaus, the following are the only terms of this description employed ; viz. — 1. JBacca, a berry, any fleshy fruit. 2. Acinus, a bunch of fleshy fruit, especially a bunch of grapes. 3. Caclirys, a cone, as of the pine tree. 4. Pilula, a cone like the Galbulus of modern botanists. 5. Folliculus (Fuchs), any kind of capsule. 6. Grossus, the fruit of the fig unripe. 7. Siliqua, the coating of any fruit. In his " Philosophia Botanica" Linnaeus gives the follow- ing definitions of the terms he employs : — 1. Capsula, hollow, and dehiscing in a determinate manner. 2. Siliqua, two-valved, with the seeds attached to both sutures. 3. Legumen, two-valved, with the seeds attached to one suture only. 4. Conceptaculum, one-valved, opening longitudinally on one side, and distinct from the seeds. 5. Drupa, fleshy without valves, containing a nut. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 167 6. Pomum, fleshy without valves, containing a capsule. 7. Bacca, fleshy without valves, containing naked seeds. 8. Strobilus, an amentum converted into a pericarpium. Gartner has the following, with definitions annexed to them : — 1. Capsula, a dry, membranous, coriaceous, or woody peri- carpium, sometimes valveless, but more commonly dehiscing with valves. Its varieties are, — a. Utriculus, an unilocular one-seeded capsule, very thin and transparent, and constantly valvular ; as in Chenopodium, Atriplex, Adonis. b. Samara, an indehiscent, winged, one or two-celled cap- sule ; as Ulmus, Acer, Liriodendron. c. Folliculus, a double one-celled, one-valved, membran- ous, coriaceous capsule, dehiscing on the inside, and either bearing the seed on each margin of its suture, or on a recep- tacle common to both margins ; as Asclepias, Cinchona, and Vinca. 2. Nux, a hard pericarpium, either indehiscent or never dividing into more than two valves ; as in Nelumbium, Bora- ginese, and Anacardium. 3. Coccum is a pericarpium of dry elastic pieces or coccula, as in Diosma, Dictamnus, Euphorbia. 4. Drupa is an indehiscent pericarpium with a variable rind, very different in substance from the putamen, which is bony ; as in Lantana, Cocos, Sparganium, Gaura, &c. 5. Bacca, any soft pericarpium, whether succulent or other- wise; provided it does not dehisce into regular valves, nor contain a single stone adhering to it. Of this the following are kinds : — a. Acinus, a soft, succulent, semi-transparent unilocular berry, with one or two hard seeds, as the grape, Rivina, Rhipsalis, Rubus, Grossularia, &c. b. Pomum, a succulent or fleshy, two or many celled berry, the dissepiments of which are fleshy or bony, and coherent at the axis ; as Pyrus, Crataegus, Cydonia, Sapota, and others. c. Pepo, a fleshy berry with the seeds attached at a dis- m 4 168 ORGANOGRAPHY. BOOK I. tance from the axis upon the parietes of the pericarp ; as Cucumis, Stratiotes, Passiflora, Vareca, and others. To the term bacca all other succulent fruits are referred which belong to neither Acinus, Pomum, nor Pepo ; as Gar- cinia, Caryophyllus, Cucubalus, Hedera. 6. Lcgumen, the fruit of Leguminosae. 7. Siliqua and Silicula, the fruit of Cruciferae. Willdenaw defines those employed by him in the following manner : — 1. Utriculus, a thin skin enclosing a single seed. Adonis, Galium, Amaranthus. 2. Samara, a pericarpium containing one, or at most two seeds, and surrounded by a thin membrane, either along its whole circumference or at the point, or even at the side. Ulmus, Acer, Betula. 3. Follicuhis, an oblong pericarpium bursting longitudi- nally on one side, and filled with seeds. Vinca. 4. Capsida, a pericarpium consisting of a thin coat contain- ing many seeds, often divided into cells, and assuming various forms. Silene, Primula, Scrophularia, Euphorbia, Magnolia. 5. Nux, a seed covered with a hard shell which does not burst. Corylus, Quercus, Cannabis. 6. Drupa, a nut covered with a thick succulent or carti- laginous coat. Primus, Cocos, Tetragonia, Juglans, Myris- tica, Sparganium. 7. Bacca, a succulent fruit containing several seeds, and not dehiscing. It encloses the seeds without any determinate order, or it is divided by a thin membrane into cells. Ribes, Garcinia, Hedera, Tilia. Rubus has a compound bacca. 8. Pomum, a fleshy fruit that internally contains a capsule for the seed. It differs from the celled berry in having a per- fect capsule in the heart. Pyrus. 9. Pepo, a succulent fruit which has its seeds attached to the inner surface of the rind. Cucumis, Passiflora, Stratiotes. 10. Siliqua, a dry elongated pericarp, consisting of two halves held together by a common permanent suture. Cru- ciferae. Silicula is a small form of the same. 11. Legumen, a dry elongated pericarpium, consisting of CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 169 two halves or valves externally forming two sutures. Legu- minosae. 12. Lomentanii a legumen divided internally by spurious dissepiments, not dehiscing longitudinally, but either remain- ing always closed, as Cassia Fistula, or separating into pieces at transverse contractions along its length, as Ornithopus. The following are enumerated as spurious fruits. 13. Strobi/us, an Amentum the scales of which have be- come woody. Pinus. 14. Spurious capsule. Fagus, Rumex, Carex. 15. Spurious nut. Traps, Coix, Mirabilis. 16. Spurious drupe. Taxus, Anacardium, Semecarpus. 17. Spurious bacca. Juniperus, Fragaria, Basella. By this author the names of fruits are, perhaps, more loosely and inaccurately applied than by any other. Professor Link objects to applying particular names to variations in anatomical structure ; observing, " that botanists have strayed far from the right road in distinguishing these terms by characters which are precise and difficult to seize. Terms are only applied to distinct parts, as the leaf, peduncle, calyx, and stamens, and not to modifications of them. Who has ever thought of giving a distinct name to a labiate or papilionaceous corolla, or who to a pinnated leaf?" But this sort of reasoning is of little value if it is considered that the fruit is subject to infinitely greater diversity of structure than any other organ, and that names for these modifications have become necessary, for the sake of avoiding a minute explan- ation of the complex differences upon which they depend. Besides, to admit, as Professor Link actually does, such names as capsula, &c. is abandoning the argument; and when the following definitions, which this learned botanist has proposed, are considered, I think that little doubt need exist as to whether terms should be employed in the manner recommended by himself, or with the minute accuracy of the French. According to Professor Link, the following are the limits of Carpological nomenclature : — 1. Capsula^ any dry membranous or coriaceous pericarp. 2. Capsella, the same, if small and one-seeded. 170 ORGANOGRAPHY. BOOK I. 3. Nux, externally hard. 4. Nucula, externally hard, small, and one-seeded. 5. Drupa, externally soft, internally hard. 6. Pomum, fleshy or succulent, and large. 7. Bacca, fleshy or succulent, and small. 8. Bacca sicca, fleshy when unripe, dry when ripe, and then distinguishable from the capsule by not being brown. 9. Legwnen, "I , . c . , , ~ r.-,- r the pericarps or certain natural orders. 10. Sihqua, J ' t 11. Amphispermium, a pericarpium which is of the same figure as the seed it contains. In more recent times there have been three principal attempts at classing and naming the different modifications of fruit ; namely, those of Richard, Mirbel, and Desvaux. These writers have all distinguished a considerable number of varia- tions, of which it is important to be aware for some purposes, although their nomenclature is not much employed in practice. But, in proportion as the utility of a classification of fruit con- sists in its theoretical explanation of structure rather than in a strict applicability to practice, it becomes important that it should be founded upon characters which are connected with internal and physiological distinctions rather than with external and arbitrary forms. Viewing the subject thus, it is not to be concealed that, notwithstanding the undoubted experience and talent of the writers just mentioned, their carpological systems are essentially defective. Besides this, each of the three writers has felt himself justified in contriving a nomenclature at variance with that of his predecessors, for reasons which it is difficult to comprehend. I have attempted to adjust the synonyms of carpological writers, and have also ventured to propose a new arrange- ment, in which those names which seem to be most legitimate are retained in every case, their definitions only being altered ; previously to which I shall briefly explain the methods of Messrs. Richard, Mirbel, and Desvaux. The Arrangement of Richard. Class 1. Simple fruits. § 1. Dry. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 171 * Indehiscent. * * Dehiscent. § 2. Fleshy. Class 2. Multiplied fruits. Class 3. Aggregate or compound fruits. The Arrangement of Mirbel. Class 1. Gymnocarpiens. Fruit not disguised by the ad- herence of any other organ than the calyx. Ord. 1. Carcerulaires. Pericarpium indehiscent, but sometimes with apparent sutures, generally dry, superior or inferior, mostly unilocular and monosper- mous, sometimes plurilocular and polyspermous. Ord. 2. Capsalaires. Pericarpium dry, superior or inferior, opening by valves, but never separating into distinct pieces or cocci. Ord. 3. Dieresiliens. Pericarpium superior or inferior, dry, regular, and monocephalous (that is, having one common style), composed of several distinct pieces arranged systematically round a central real or imaginary axis, and separating at maturity. Ord. 4. Etairionaires. Pericarps several, irregular, superior, one or many-seeded, with a suture at the back. Ord. 5. Cenobionaires. A regular fruit divided to the base into several acephalous pericarpia ; that is to say, not marked on the summit by the stigmatic scar, the style having been inserted at their base. Ord. 6. Dnipacees. Pericarpium indehiscent, fleshy externally, bony internally. Ord. 7. Bacciens. Succulent, many-seeded. Class 2. Angiocarpiens. Fruit seated in envelopes not form- ing part of the calyx. The Arrangement of Desvaux. Class 1. Pericarpium dry. Ord. 1. Simple fruits. 172 ORGANOGRAPHY. BOOK I. § Indehiscent. § § Dehiscent. Ord. 2. Dry compound fruits. Class 2. Pericarpium fleshy. Ord. 1. Simple fruits. - Ord. 2. Compound fruits. In explanation of the principles upon which the classifica- tion of fruit which I now venture to propose is founded, it will of course be expected that I should offer some observ- ations. In the first place, I have made it depend primarily upon the structure of the ovarium, by which the fruit is of necessity influenced in a greater degree than by any thing else, the fruit itself being only the ovarium at maturity. In using the terms simple and compound, I have employed them precisely in the sense that has been attributed to them in my remarks upon the ovarium ; being of opinion that, in an arrangement like the following and those which have pre- ceded it, in which theoretical rather than practical purposes are to be served, the principles on which it depends should be conformable to the strictest theoretical rules of structure. A consideration of the fruit without reference to the ovarium necessarily induces a degree of uncertainty as to the real nature of the fruit ; the abortion and obliteration, to which almost every part of it is more or less subject, often disguising it to such a degree that the most acute carpologist would be unable to determine its true structure from an examination of it in a ripe state only. In simple fruits are stationed those forms in which the ovaria are multiplied so as to resemble a compound fruit in every respect except their cohesion, they remaining simple. But, as the passage which is thus formed from simple to compound fruits is deviated from materially when the ovaria are placed in more than a single series, I have found it advisable to constitute a particular class of such under the name of aggregate fruit. Care must be taken not to confound these with the fourth class containing col- lective fruits, as has been done by more carpologists than one. While the true aggregate fruit is produced by the ovaria of a single flower, a collective fruit, if aggregate, is produced by the ovaria of many flowers ; a most important difference. As CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 173 the pericarpium is necessarily much affected by the calyx when they adhere so as to form a single body, it is indispensable, if a clear idea is to be attached to the genera of carpology, that inferior or superior fruits should not be confounded under the same name ; for this reason I have in all cases founded a dis- tinction upon that character. In order to facilitate the knowledge of the limits of the genera of carpology, the following analytical table will be found convenient for reference. It is succeeded by the cha- racters of the genera in as much detail as is necessary for the perfect understanding of their application. Class I. Fruit simple. APOCARPI. One or two-seeded. Membranous, - - Dry and bony, Fleshy externally, bony internally, Many-seeded. Dehiscent. One-valved, - Two-valved, Indehiscent, - Class II. Fruit aggregate. AGGREGATI. Ovaria elevated above the calyx. Pericarpia distinct, - Pericarpia cohering into a solid mass, Ovaria enclosed within the fleshy tube of the calyx, Class III. Fruit compound. SYNCARPI. Sect. I. Superior. A. Pericarpium dry externally. Indehiscent. Utricui.us. achenium. Drupa. Foixiculus. Legumen. lomenium. Et,erio. SyNCARPIUM. Cynarrhodum. One-celled, - - Caryopsis. Many-celled. Dry internally. Apterous - Carcerulus. Winged, - Samara. Pulpy internally, - Amphisarca. Dehiscent. By a transverse suture, - Pyxidium. By elastic cocci, - Regma. By a longitudinal suture, - CONCEPTACULUM. By valves. Placenta; opposite the lobes of the stigma. Linear, - SlLIQUA. Roundish, - SlLICULA. 174 ORGANOGRAPHY. BOOK f. Placentae alternate with the lobes of the stigma. Valves separating from the replum, Ceratium. Sect. 2. A. Replum none, Pericarpium fleshy. Indehiscent. Sarcocarpium separable, Sarcocarpium inseparable, Dehiscent, - Inferior. Pericarpium dry. Indehiscent. Capsula. Hesperidium. Nuculanium. Trtma. Cells two or more, - Cremocarpium. Cell one. Surrounded by a cupulate involucrum, Glans. Destitute of a cupula, - Cypsela. Dehiscent or rupturing, - DlPLOTEGIA. B. Pericarpium fleshy. Epicarpium hard. Seeds parietal, - Pepo. Seeds not parietal, - Balausta. Epicarpium soft. Cells obliterated ; or unilocular, - Bacca. Cells distinct, - - POMUM. Class IV. Collective fruits. ANTHOCARPI. Single. Perianthium indurated, dry, - Dicr,Esiuji. Perianthium fleshy, - - Sphalerocar- Aggregate. PIUM. Hollow, - - Syconus. Convex. An indurated amentum, - Strobilus. A succulent spike, - Sorosis. Class I. Fruit simple. APOCARPI. Ovaria strictly simple ; a single series only produced by a single flower. I. Utriculus, Gcertner (Cystidium, Link.) One-celled, one or few-seeded, superior, membranous, frequently dehiscent by a transverse incision. This differs from the pyxis in texture, being strictly simple, i. e. not proceeding from an ovarium with obliterated dissepiments. Example. — Amaranthus, Chenopodium. II. Achenium ; (Akenium, of many ; Spermidium ; Xylodium, Desv.; The- cidium, Mirb. ; Nux, Linn.) One-seeded, one-celled, superior, indehiscent, hard, dry, with the integu- ments of the seed distinct from it. Linnaeus includes this among his seeds, defining it " semen tectum epider- mide ossea." I have somewhere seen it named Spermidium ; a good term if it were wanted. M. Desvaux calls the nut of Anacardium a Xylodium. Examples. — Lithospermum, Borago. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 17.5 III. Drupa Drupe. —Jig, 163. One-celled, one or two-seeded, superior, indehiscent, the outer coat (nau- cum) soft and fleshy, and separable from the inner or endocarpium (the stone), which is hard and bony ; proceeding from an ovarium which is perfectly simple. This is the strict definition of the term drupa, which cannot strictly be applied to any compound fruit, as that of Cocos, certain Verbenaceae, and others, as it often is. Fruits of the last description are generally carcerules with a drupaceous coat. The stone of this fruit is the Kux of Richard, but not of others. Examples. — Peach, Plum, Apricot. IV. Foluculus. — Follicle (Hemigyrus, Desvaux; Plopocarpium, Desv.), fig- I*). One-celled, one or many-seeded, one-valved, superior, dehiscent by a suture along its face, and bearing its seeds at the base, or on each margin of the suture. This differs from the legumen in nothing but its having one valve instead of two. The Hemigyrus of Desvaux is the fruit of Proteaceae, and differs from the follicle in nothing of importance. When several follicles are in a single flower, as in Nigella and Delphinium, they constitute a form of fruit called Plopocarpium by Desvaux, and admitted into his Etaerio by Mirbel. Examples. — Preonia, Banksia, Nigella. V. Legumen. — Pod (Legumen, Linn.; Gousse, Fr.), fig. 136, 137. One-celled, one or many-seeded, two-valved, superior, dehiscent by a suture along both its face and its back, and bearing its seeds on each margin of the ventral suture. This differs from the follicle in nothing except its dehiscing by two valves. In Astragalus two spurious cells are formed by the projection inwards of either the dorsal or ventral suture, which forms a sort of dissepiment; and in Cassia a great number of transverse diaphragms (phragmata) are formed by projections of the placenta. Sometimes the legumen is indehiscent, as in Cathartocarpus, Cassia fistula, and others ; but the line of dehiscence is in such species indicated by the presence of sutures. When the two sutures of the legumen separate from the valves, they form a kind of frame called replum, as in Carmichaelia. Examples. — Bean, Pea, Clover. VI. Lomentum. — (Legumen lomentaceum, Rich.) Differs from the legumen in being contracted in the spaces between each seed, and there separating into distinct pieces, or indehiscent, but divided by internal spurious dissepiments, whence it appears at maturity to consist of many articulations and divisions. Example. — Ornithopus. Class II. Fruit aggregate. A G G RE G A TI. Ovaria strictly simple ; more than a single series produced by eachjlower. VII. Etaerio, Mirb. — (" Polychorion, Mirb. ; " Polysecus, Desvaux; Amalthea, Desv. ; Erythrostomum, Desvaux), Jig. 161. Ovaries distinct; pericarpia indehiscent, either dry upon a dry receptacle, as Ranunculus, dry upon a fleshy receptacle, as strawberry, or fleshy upon a 176 ORGANOGRAPHY. BOOK I. dry receptacle, as Rubus. The last is very near the syncarpium, from which it differs in the ovaria not coalescing into a single mass. It is Desvaux's Erythrostomum. This term is applied less strictly by M. Mirbel, who admits into it dehiscent pericarpia, not placed upon an elevated receptacle, as Delphi- nium and Paeonia ; but the fruit of these plants is better understood to be a union of several follicules within a single flower. If there is no elevated receptacle, we have Desvaux's Amalthea. The parts of an Etaerio are Achenia. Examples. Ranunculus, Fragaria, Rubus. VIII. Syncarpium. — (Syncarpium, Rich. ; Asimina, Desv.) Ovaries cohering into a solid mass, with a slender receptacle. Examples. Annona, Magnolia. IX. Cyxarrhodum. — (Cynarrhodum, Officin. Desvaux.) Ovaries distinct ; pericarpia hard, indehiscent, enclosed within the fleshy tube of a calyx. Examples. Rosa, Calycanthus. Class III. Fruit compound. SYNCARPI. Ovaria compound. Sect. 1. Fruit superior. A. Pericarpium dry. X. Caryopsis. — (Cariopsis, Rich. ; Cerio, Mirb.) One-celled, one-seeded, superior, indehiscent, dry, with the integuments of the seed cohering inseparably with the endocarpium, so that the two are undis- tinguishable ; in the ovarium state evincing its compound nature by the pre- sence of two or more stigmata ; but nevertheless unilocular, and having but one ovulum. Examples. Wheat, Barley, Maize. XI. Reqma, Mirb. ; — (Elaterium, Rich. ; Capsula tricocca, L.) Three or more celled, few-seeded, superior, dry, the cells bursting from the axis with elasticity into two valves. The outer coat is frequently softer than the endocarpium or inner coat, and separates from it when ripe; such regmata are drupaceous. The cells of this kind of fruit are called cocci. Example. Euphorbia. XII. Carcerulus, Mirb. ; — (Dieresilis, Mirb. ,- Ca?nobio, Mirb. ; Synochorion, Mirb.; Sterigmum, Desvaux ; Microbasis, Desvaux ,- Polexostylus, Mirb.; Sarcobasis, Dec, Desv. ; Baccaularius, Desv. ) Many-celled, superior ; cells dry, indehiscent, few-seeded, cohering by a common style round a common axis. From this the Dieresilis of Mirbel does not differ in any essential degree. The same writer calls the fruit of Labiatai (Jig. 162.), which Linnaeus and his followers mistake for naked seeds, Caano- bio : it differs from the Carcerulus in nothing but the low insertion of the style into the ovaria, and the distinctness of the latter. Examples. Tilia, Tropaeolum, Malva. XIII. Samara, Geertn. ; — Key. (Pteridium, Mirb. ; Pterodium, De$v.),Jig. 143. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 177 Two or more celled, superior; cells few-seeded, indeliiscent, dry; elongated into wing-like expansions. This is nothing but a modification of the Carcerule. Examples. Fraxinus, Acer, Ulmus. XIV. Pvxidium (Pyxidium, Ehr., Rich., Mirb.; Capsula circumscissa, L.), Jig- 152. One-celled, many-seeded ; superior, or nearly so ; dry, often of a thin tex- ture ; dehiscent by a transverse incision, so that when ripe the seed and their placenta appear as if seated in a cup, covered with a lid. This fruit is one- celled by the obliteration of the dissepiments of several carpella, as is apparent from the bundles of vessels which pass from the style through the pericarpium down into the receptacle. Example. Anagallis. XV. Conceptaculum (Conceptaculum, Linn.; Double Follicule, Mirb.), Jig. 138, 139. Two-celled, many-seeded, superior, separating into two portions, the seeds of which do not adhere to marginal placentae, as in the folliculus, to which this closely approaches, but separate from their placentae, and lie loose in the cavity of each cell. Examples. Asclepias, Echites. XVI. Siliqca, Linn. Jig. 155, 156, 157. One or two-celled, many-seeded, superior, linear, dehiscent by two valves separating from the replum ; seeds attached to two placentas adhering to the replum, and opposite to the lobes of the stigma. The dissepiment of this fruit is considered a spurious one formed by the projecting placentas, which some- times do not meet in the middle ; in which case the dissepiment or phragma has a slit in its centre, and is said to he fenestrate. XVII. — Silicula. Linn. This differs from the latter in nothing but its figure, and in containing fewer seeds. It is never more than four times as long as broad, and often much shorter. Examples. Thlaspi, Lepidium, Lunaria. XVIII. Ceratium. — (Capsula siliquiformis, Dec. ; Conceptaculum, Desv.) One-celled, many-seeded, superior, linear, dehiscent by two valves separat- ing from the replum ; seeds attached to two spongy placenta? adhering to the replum, and alternate with the lobes of the stigma. Differs from the siliqua in the lobes of the stigma being alternate with the placentae, not opposite. This, therefore, is regular, while that is irregular in structure. Examples. Glaucium, Corydalis, Hypecoum. XIX. Capsula, Capsule, Jig. 145, 146. 150, 151. 134, 135. One or many-celled, many-seeded, superior, dry, dehiscent by valves, always proceeding from a compound ovarium. The valves are variable in their nature : usually they are at the top of the fruit, and equal in number to the cells; sometimes they are twice the number; occasionally they resemble little pores or holes below the summit, as in the Antirrhinum. Examples. Digitalis, Primula, Rhododendron. XX. Amphisarca. — (Amphisarca, Desv.) Many-celled, many-seeded, superior, indehiscent ; indurated or woody exter- nally, pulpy internally. N 178 ORGANOGRAPHY. BOOK I. Examples. Omphalocarpus, Adansonia, Crescentia. B. Pericarpium fleshy. XXI. Tryma (Tryma, Watson.) Superior, by abortion one-celled, one-seeded, with a two-valved indehiscent endocarpium, and a coriaceous or fleshy, valveless sarcocarpium. Example. Juglans. XXII. Nucula^ium. — (Nuculanium, Rich. ; Bacca, Desvaux.) Two or more celled, few or many-seeded, superior, indehiscent, fleshy, of the same texture throughout, containing several seeds, improperly called nucules by the younger Richard. This differs scarcely at all from the berry, except in being superior. Examples. Grape, Achras. XXIII. Hesperidium. — (Hesperidium, Desv. Rich.) Many-celled, few-seeded, superior, indehiscent, covered by a spongy separ- able rind ; the cells easily separable from each other, and containing a mass of pulp, in which the seeds are imbedded. The pulp is formed by the cellular tissue, which forms the lining of the cavity of the cells : this cellular tissue is excessively enlarged and succulent, is filled with fluid, and easily coheres into a single mass. The external rind is by M. De Candolle supposed to be an elevated discus of a peculiar kind, analogous to that within which the fruit of Nelumbium is seated ; and perhaps its separate texture and slight connexion with the cells of the fruit seem to favour this supposition. But it is difficult to reconcile with such an hypothesis the continuity of the rind with the style and stigma, which is a sure indication of the identity of their origin ; and it is certain that the shell of the ovarium and the pericarpium are the same. The most correct explanation of this structure is to consider the rind a union of the epicarp and sarcocarp, analogous to that of the drupa. Example. Orange. Sect. 2. Fruit inferior. A. Pericarpium dry. XXIV. Glans (Glans, Linn. Desv. ; Calybio, Mirb. ,• Nucula, Desvaux), Jig. 164. One-celled, one or few-seeded, inferior, indehiscent, hard, dry; proceeding from an ovarium containing several cells and several seeds, all of which are abortive but one or two ; seated in that kind of persistent involucre called a cupule. The pericarpium is always crowned with the remains of the teeth of the calyx ; but they are exceedingly minute, and are easily overlooked. Sometimes the gland is solitary, and quite naked above, as in the common oak ; sometimes there is more than one completely enclosed in the cupule, as the beech and sweet chestnut. Examples. Quercus, Corylus, Castanea. XXV. Cypsela (Akena, Necker ; Akenium, Rich.; Cypsela, Mirb.; Ste- phanoum, Desv.), fig. 147, 148. One-seeded, one-celled, indehiscent, with the integuments of the seed not cohering with the endocarpium ; in the ovarium state evincing its compound nature by the presence of two or more stigmata ; but nevertheless unilocular, and having but one ovulum. Such is the true structure of the Achenium ; but, as that term is often applied to the simple superior fruits, called Nux by CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 179 Linnrcus, I have thought it hetter, in order to avoid confusion, to adopt the name Cypsela. Examples. All Composite. XXVI. Cremocarpium (Cremocarpium, Mirb. ; Polakenium, or Pentake- nium, Ttich. ; Carpadelium, Desv.), fig. 153, 154. 158, 159. Two to five-celled, inferior ; cells one-seeded, indehiscent, dry, perfectly close at all times ; when ripe separating from a common axis. M. Mirbel confines the application of Cremocarpium to Umbellifera; ; but it is better to let it apply to all fruits which will come within the above definition. It will then be the same as Richard's Polakenium, excluding those forms in which the fruit is superior. The latter botanist qualifies his term Polakenium according to the number of cells of the fruit: thus when there are two ceWs'itis diakenium, three triakenium, and so on. M. De Candolle calls the half of the fruit of Umbellifera; ?nericarpc Examples. Umbellifera?, Aralia, Galium. XXVII. Diplotegia (Diplotegia, Desv.), fig. 144. One or many-celled, many-seeded, inferior, dry, usually bursting either by pores or valves. This differs from the Capsule only in being adherent to the calyx. Examples. Campanula, Leptospermum. B. Peri carpi urn fleshy. XXVIII. Pomum, Apple or Pome (Melonidium, Rich. ,- Pyridium, Mirb. ; Pyrenarium, Desvaux ; Antrum, Mcench.) fig. 165. Two or more celled, few-seeded, inferior, indehiscent, fleshy; the seeds dis- tinctly enclosed in dry cells, with a bony or cartilaginous lining, formed by the cohesion of several ovaria with the sides of the fleshy tube of a calyx, and some- times with each other. These ovaria are called parietal by M. Richard. Some forms of Nuculanium and this differ only in the former being distinct from the calyx. Examples. Apple, Cotoneaster, Crataegus. XXIX. Pefo. — (Peponida, Rich.) One-celled, many-seeded, inferior, indehiscent, fleshy ; the seeds attached to parietal pulpy placenta;. This fruit has its cavity frequently filled at maturity with pulp, in which the seeds are imbedded ; their point of attachment is, how- ever, never lost. The cavity is also occasionally divided by projections of the placenta into spurious cells, which has given rise to the belief that in Pepo Ma- crocarpus there is a central cell, which is not only untrue but impossible. Examples. Cucumber, Melon, Gourd. XXX. Bacca, Berry (Bacca, L. ; Acrosarcum, Desvaux), fig. 160. Many-celled, many-seeded, inferior, indehiscent, pulpy ; the attachment of the seeds lost at maturity, when they become scattered in the substance of the pulp. This is the true meaning of the term berry ; which is, however, often otherwise applied, either from mistaking nucules for seeds, or from a mis- apprehension of the strict limits of the term. Example. Ribes. XXXI. Balausta. — (Balausta, Officin. Rich.) Many-celled, many-seeded, inferior, indehiscent ; the seeds with a pulpy N 2 ^- 180 ORGANOGRAPHY. BOOK I. coat, and attached distinctly to their placentae. The rind was called Malicorium by Ruellius. Example. Pomegranate. Class IV. Collective Fruits. Anthocarpi. Fruit of which the principal characters are derived from the incrassated floral envelopes. XXXII. Diclesium. — (Dyclesium, Desvaux ; Sclerantlium, Mcench ; Catacle- sium, Desvaux ; Sacellus, Mirb.) Pericarpium indehiscent, one-seeded, enclosed withi n an indurated perian- thium. Examples. Mirabilis, Spinacia, Salsola. XXXIII. Sphalerocarpum. — (Sphalerocarpum, Desv. ; Nux baccata of authors. ) Pericarpium indehiscent, one-seeded, enclosed within a fleshy perianthium. Examples. Hippophae, Taxus, Blitum, Basella. XXXIV. Stconus. — (Syconus, Mirb.) A fleshy rachis, having the form of a flattened disk, or of a hollow receptacle, with distinct flowers and dry pericarpia. Examples. Ficus, Dorstenia, Ambora. XXXV. Strobilus, Cone (Conus, or Strobilus, Rich., Mirb. ; Galbulus, Gtertn. ; Arcesthide, Desvaux; Cachrys, Fuchs ; Pilula, Pliny), fig. 166. An amentum, the carpella of which are scale-like, spread open, and bear naked seeds ; sometimes the scales are thin, with little cohesion ; but they often are woody, and cohere into a single tuberculated mass. The Galbulus differs from the Strobilus only in being round, and having the heads of the carpella much enlarged. The fruit of the Juniper is a Gal- bulus, with fleshy coalescent carpella. Desvaux calls it Arcesthide. Example. Pinus. XXXVI. Sorosis. — (Sorosis, Mirb.) A spike or raceme converted into a fleshy fruit by the cohesion in a single mass of the ovaria and floral envelopes. Examples. Ananassa, Moms, Artocarpus. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 181 14. Of the Seed. 17 6 181 184 185 As the fruit is the ovarium arrived at maturity, and is therefore subject to the same laws of structure as the latter; so is the seed the ovulum in its most perfect and finally organised state, and constructed upon exactly the same plan as the ovulum. But as the fruit, nevertheless, often differs from the ovarium in the suppression, or addition, or modi- fication of certain portions, so is the seed occasionally altered from the precise structure of the ovulum, in consequence of changes of like nature. The seed is a body enclosed in a pericarpium, is clothed with its own integuments, and contains the rudiment of a future plant. It is the point of developement at which vege- tation stops, and beyond which no increase, in the same direction with itself, can take place. In a young state it has already been spoken of under the name of ovulum ; to which I also refer for all that relates to the insertion of seeds. That side of a seed which is most nearly parallel with the axis of a compound fruit, or the ventral suture or sutural line of a simple fruit, is called the face, and the opposite side the back. In a compound fruit with parietal placentas, the pla- centa is to be considered as the axis with respect to the seed ; and that part of the seed which is most nearly parallel with the placenta, as the face. Where the raphe is visible, the face is indicated by that. When a seed is flattened lengthwise it is said to be com- N 3 182 ORGANOGRAPHY. BOOK I. pressed, when vertically it is depressed; a difference which it is of importance to bear in mind, although it is not always easy to ascertain it : for this purpose it is indispensable that the true base and apex of the seed should be clearly under- stood. The base of a seed is always that point by which it is attached to the placenta, and which receives the name of hilum ; the base being found, it would seem easy to deter- mine the apex, as a line raised perpendicularly upon the hilum, cutting the axis of the seed, ought to indicate the apex at the point where the line passes through the testa : but the apex so indicated would be the geometrical, not the natural apex ; for discovering which with precision in all seeds, the natural and geometrical apex of which do not correspond, another plan must be followed. If the testa of a seed be carefully examined, it will usually be found that it is com- posed in great part of lines representing rows of cellular tissue, radiating from some one point towards the base, or, in other words, of lines running upwards from the hilum and meet- ing in some common point. This point of union or radiation is the true apex, which is not only often far removed from the geometrical apex, but is sometimes even in juxtaposition with the hilum, as in mignonette : in proportion, therefore, to the obliquity of the apex of the seed will be the curve of its axis, which is represented by a line passing through the whole mass of the seed from the base to the apex, accurately follow- ing its curve. If the lines above referred to are not easily distinguished, another indication of the apex resides in a little brown spot or areola, hereafter to be mentioned under the name of chalaza. Where there is no indication either exter- nally or internally of the apex, it may then be determined geometrically. The integuments of a seed are called the testa ; the rudi- ment of a future plant, the embryo (Plate VI. fig. 1. b, &c.) ; and a substance interposed between the embryo and the testa, the albumen (fig. 1. a, 5. «, &c). The testa, called also lorica by Mirbel, perisperme and episperme by Richard, and spermodermis by De Candolle, according to some consists, like the pericarpium, of three portions; viz. 1. the external integument, tunica externa of CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 183 WilldenoWj testa of De Candolle ; 2. the internal integu- ment, tunica interna of Willdenow, endopleura of De Can- dolle, hilofere and tegmen of Mirbel ; and, 3. of an inter- vening substance answering to the sarcocarpium, and called sarcodcrmis by De Candolle : this last is chiefly present in seeds with a succulent testa, and by many is considered a portion of the outer integument, which is the most accurate mode of understanding it. The outer integument is either membranous, coriaceous, crustaceous, bony, spongy, fleshy, or woody ; its surface is either smooth, polished, rough, or winged, and sometimes is furnished with hairs, as in the cotton and other plants, which, when long and collected about either extremity, form what is called the coma (sometimes also, but improperly, the pappus). It consists of cellular tissue disposed in rows, with or without bundles of vessels intermixed : in colour it is usually of a brown or similar hue : it is readily separated from the inner integument. In Maurandya Barclayana it is formed of reticulated cel- lular tissue ; in Collomia linearis and others it is caused by elastic spirally twisted fibres enveloped in mucus, and springing outwards when the mucus is dissolved ; in Casu- arina it (or the inner integument) contains a great quantity of spirally fibrous cellules. In the genus Crinum it is of a very fleshy, succulent character, and has been mistaken for albumen, from which it is readily known by its vascularity. According to Mr. Brown, a peculiarly anomalous kind of partition, which is found lying loose within the fruit of Banksia and Dryandra, without any adhesion either to the pericarpium or the seed, is a state of the outer integument. It is said that in those genera the inner membrane (secondine) of the ovulum before fertilisation is entirely exposed, the primine being dimidiate and open its whole length ; and that the outer membranes (primines) of the two collateral ovula, although originally distinct, finally contract an adhesion by their corresponding surfaces, and together constitute the anomalous dissepiment. But it may be reasonably doubted whether the integument here called secondine is not primine, and the sup- posed primine arillus. n 4 184 ORGANOGRAPHY. BOOK I. The inner membrane (secondine) of the ovulum, however, in general appears to be of greater importance as connected with fecundation, than as affording protection to the nucleus at a more advanced period. For in many cases, before im- pregnation, its perforated apex projects beyond the aperture of the testa, and in some plants puts on the appearance of an obtuse, or even dilated stigma ; while in the ripe seed it is often either entirely obliterated, or exists only as a thin film, which might readily be mistaken for the epidermis of a third membrane, then frequently observable. " This third coat (tercine) is formed by the proper membrane or cuticle of the nucleus, from whose substance in the unim- pregnated ovulum it is never, I believe, separable, and at that period is very rarely visible. In the ripe seed it is distin- guishable from the inner membrane only by its apex, which is never perforated, is generally acute and more deeply co- loured, or even sphacelated." M. Mirbel has, however, justly remarked that the pri- mine and the secondine are, in the seed, very frequently con- founded ; and that therefore the word testa is better employed, as one which expresses the outer integument of the seed without reference to its exact origin, which is practically of little importance. The tercine is also, no doubt, often absent. He observes, that these mixed integuments often give rise to new kinds of tissue ; that in Phaseolus vulgaris the testa consists, indeed, of three distinct layers, but of those the innermost was the primine ; and that the others, which repre- sent nothing that pre-existed in the ovulum, have a horny consistence, and are formed of cylindrical cellules, which elongate in the direction from the centre to the circumference. And this is probably the structure of the testa of many Legu- minosse. It sometimes happens that the endopleura (or tercine ?) thickens so much as to have the appearance of albumen, as in Cathartocarpus fistula. In such a case as this it is only to be distinguished from albumen by gradual observation from the ovulum to the ripe seed. With regard to the quartine and quintine, one of them is occasionally present in the form of a fleshy sac that is inter- posed between the albumen and the ovulum, and envelopes CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 185 the latter. It is what was called the vitellus by Gaertner, and which M. Richard, by a singular prejudice, considered a dila- tation of the radicle of the embryo, to his macropodal form of which he referred the embryo of such plants. Instances of this are found in Nymphaea and its allies, and also in Scita- mineae, peppers, and Saururus. Mr. Brown, who first ascer- tained the fact, considers this sac to be always of the same nature and origin, and as the vesicula colliquamenti or amnios of Malpighi. The end by which the seed is attached to the placenta is called the hilum or umbilicus (Plate VI. fig. 5. c, 17. e, 11. c, &c.) ; it is frequently of a different colour from the rest of the seed, not uncommonly being black. In plants with small seeds it is exceedingly minute, and recognised with difficulty; but in some it is so large as to occupy fully a third part of the whole surface of the seed, as in the horse-chestnut, Sa- poteae, and others. Seeds of this kind have been called nauca by Gaertner. In grasses the hilum is indicated by a brownish spot situated on the face of the seed, and is called by M. Richard spilus. The centre of the hilum, through which the nourishing vessels pass, is called by Turpin the omphalodium. Sometimes the testa is enlarged in the form of irregular lumps or protuberances about the umbilicus ; these are called strophiolce or carunculee ; and the umbilicus, about which they are situated, is said to be strophiolate or carun- culate. M. Mirbel has ascertained that in Euphorbia Lathy- ris the strophiola is the fungous foramen of the primine ; and it is probable that such is often the origin of this tubercle : but at present we know little or nothing upon the subject. The foramen in the ripe seed constitutes what is called the micropyle : it is always opposite the radicle of the embryo ; the position of which is therefore to be determined without dissection of the seed, by an inspection of the micropyle, — often a practical convenience. In some seeds, as the asparagus, Commelina, and others {Jig. 185.), there is a small callosity at a short distance from the hilum : this callosity gives way like a little lid at the time of germination, emitting the radicle, and has been named by Gaertner the embryolega. At the apex of the seed in the orange and many other 186 ORGANOGRAPHY. BOOK I. plants may be perceived upon the testa a small brown spot, formed by the union of certain vessels proceeding from the hilum : this spot is the chalaza (Plate VI. fig. 11. b). In the orange it is beautifully composed of dense bundles of spiral vessels and spiral ducts, without woody fibre. The vessels which connect the chalaza with the hilum constitute a parti- cular line of communication, called the raphe : in most plants it consists of a single line passing up the face of the seed ; but in many Aurantiaceae and Guttiferae it ramifies elegantly in every direction upon the surface of the testa. The raphe is always a true indication of the face of the seed; and it is very remarkable that the apparent exceptions to this rule only serve to confirm it. Thus, in some species of Euony- mus in which the raphe appears to pass along the back, an ex- amination of other species shows, that the ovula of such species are in fact resupinate ; so that with them the line of vascularity representing the raphe is turned away from its true direction by peculiar circumstances. In reality, the chalaza is the place where the secondine and the primine are connected ; so that in orthotropous seeds, or such as have the apex of the nucleus at the apex of the seed, and in which, consequently, the union of the primine and secondine takes place at the hilum, there can be no apparent chalaza, and consequently no raphe : the two latter can only exist as distinct parts in anatropous seeds, when the base of the nucleus corresponds to the geo- metrical apex of the seed. Hence, also, there can never be a chalaza without a raphe, nor a raphe without a chalaza. Something has already been said about the arillus {Jig. 186. and 187.) when speaking of the ovulum ; but it more properly comes under con- sideration along with the ripe seed. As a general rule it may be stated, that every thing proceeding from the placenta and not forming part of the seed is referable to the arillus. Even in plants like Hib- bertia volubilis and Euonymus europaeus, in which it is of un- usual dimensions, it is scarcely 186 187 CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 187 visible in the unimpregnatecl ovarium ; and it is stated by Mr. Brown, that he is not acquainted with any case in which it covers the foramen of the testa before impregnation. The mass enclosed within the true testa or outer integu- ment is called the nucleus ; and consists either of albumen and embryo, or of the latter only. The albumen {jperispermium, Juss. ; endospermium, Rich. ; medulla seminis, Jungius ; secimdince internee, Malpighi) (Plate VI. fig. 5. a, l.a, 9. a, &c), when present, is a body enclosing the embryo, and interposed between it and the in- tegument of the seed : it is of various degrees of hardness, varying from fleshy to bony, or even stony, as in some palms. It is in all cases destitute of vascularity, and has been usually considered as the amnios in an indurated state ; but Mr. Brown is of opinion that it is formed by a deposition or secretion of granular matter in the cellules of the amnios, or in those of the nucleus itself. The albumen is often absent, frequently much smaller than the embryo, but also occasion- ally of much greater size. This is particularly the case in monocotyledones, in some of which the embryo scarcely weighs a few grains, while the albumen weighs many ounces, as in the cocoa-nut. It is almost always solid, but in An- nonaceae and the nutmeg tribe it is perforated in every direction by dry cellular tissue, which appears to originate from the remains of the nucleus in which the albumen has been deposited : in this state it is said to be ruminated. The embryo (or corculum) (Plate VI. fig. 1. b, &c.) is a fleshy body, occupying the interior of the seed, and consti- tuting the rudiment of a future plant. It is usually solitary, but there are instances of the presence of several in one seed. It was originally developed within the innermost membrane of the ovulum. In most plants one embryo only is found in each seed. It nevertheless occurs, not unfrequently, that more than one is developed within a single testa, as occasionally in the orange and the hazel nut, and very commonly in Coni- ferae, Cycas, the onion, and the misseltoe. Now and then a union takes place of these embryos. It is divided into three parts; viz. the radicle (Plate VI. fig. 2. b, &c.) {rhizoma or rostellum); cotyledons (fig. 2. a, &c), 188 ORGANOGIIAPHY. BOOK I. and plumula (or gemmuld) (fig. 2. c) ; from which is also by some distinguished the cauliculus or neck {collet, scapus, sca- pellus, or tigelle). Mirbel admits but two principal parts ; viz. the cotyledons, and what he calls the blasteme, which comprises radicle, plumula, and cauliculus. Upon certain remarkable differences in the structure of the embryo, modern botanists have divided the whole vegetable kingdom into three great portions, which form the basis of what is called the natural system. These are, 1. Dicoty- ledons ; 2. Monocotyledons ; and, 3. Acotyledons. In order to understand exactly the true nature of the embryo in each of these, it will be requisite first to describe it fully as it exists in dicotyledons, and then to explain its organisation in the two others. If a common Dicotyledonous embryo (Plate VI. fig. 2.), that of the apple for example, be examined, it will be found to be an obovate, white, fleshy body, tapering and solid at the lower end, and compressed and deeply divided into two equal opposite portions at the upper end ; the lower tapering end is the radicle, and the upper divided end consists of two cotyledons. Within the base of the cotyledons is just visible a minute point, which is the plumula. The imaginary line of division between the radicle and the cotyledons is the caulicidus. If the embryo be now placed in circumstances favourable for germination, the following phenomena occur : the radicle will become elongated downwards, forming a little root ; the cauliculus will extend upwards ; the cotyledons will elevate themselves above the earth and unfold ; and the plumula will lengthen upwards, and give bhth to a stem and leaves. Such is the normal or proper appearance of a dicoty- ledonous embryo. The exceptions to it chiefly consist, 1. in the cohesion of the cotyledons in a single mass, instead of their unfolding ; 2. in an increase of their number ; 3. in their occasional absence ; and, 4. in their inequality. A cohesion of the cotyledons takes place in those embryos, which Gaertner called pseudomonocoty- ledonous, and Richard macrocephalous. In Hippocastanum, the horse-chestnut, the embryo consists of a homogeneous undi- vided mass, with a curved horn-like prolongation of one side CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 189 directed towards the hilum. If a section be made in the direction of the axis of the horn-like prolongation through the whole mass of the embryo, a slit will be observable above the middle of the horn, at the base of which lies a little conical body. In this embryo the slit indicates the division between the two bases of a pair of opposite confluent cotyledons; the conical body is the plumula, and the horn-like prolongation is the radicle. In Castanea nearly the same structure exists, except that the radicle, instead of being curved and exserted, is straight, and enclosed within the projecting base of the two cotyledons ; and in Tropaeolum, which is very similar to Cas- tanea in structure, the bases of the cotyledons are slit into four little teeth enclosing the radicle. The germination of these seeds indicates more clearly that the cotyledonary body consists of two and not of one cotyledon ; at that time the bases of the cotyledons, which had been previously scarcely visible, separate and elongate, so as to extricate the radicle and plumula from the testa, within which they had been con- fined. In number the cotyledons vary from two to a much more considerable number. Ceratophyllum has constantly four, of which two are smaller than the others ; in Coniferse they vary from two to more than twelve. Instances of the absence of cotyledons occur, 1 . In Cus- cuta (Plate VI. fig. 19.), to which they may be supposed to be denied in consequence of the absence of leaves in that genus ; 2. in Lentibularise ; 3. in Cyclamen, in which the radicle enlarges exceedingly : to these a fourth instance has by some been added in Lecythis, of which M. Richard gives the fol- lowing account. The kernel is a fleshy almond-like body, so solid and homogeneous that it is extremely difficult to dis- cover its two extremities until germination takes place : at that period one of the ends forms a little protuberance, which sub- sequently bursts through the integuments of the seed, and extends itself as a root; the other end produces a scaly plu- mula, which in time forms the stem. The great mass of the kernel is supposed by M. Richard to be an enlarged radicle. I, however, see no reason for calling the two-lobed part of the embryo (Plate VI. fig. 17. c) a plumula; it is merely coty- ledons. An inequality of cotyledons is the most unusual 190 ORGANOGRAPHY. BOOK I. circumstance with dicotyledons, and forms a distinct approach to the structure of monocotyledons : it occurs in Trapa and Sorocea, in which they are extremely disproportionate. In Cycas they are also rather unequal ; but the structure of that plant is essentially dicotyledonous. The embryo of Monocotyledons (Plate VI. fig. 1. B. &c.) is usually a solid, cylindrical, undivided, homogeneous body, slightly conical at each extremity, with no obvious distinction of radicle, plumula, or cotyledons. In germination the upper end swells and remains within the testa (fig. 10. C. b, &c.) ; the lower lengthens, opens, and emits from within one or more radicles ; and a thread-like green body is protruded from the upper part of the portion, which is lengthened beyond the testa. Here the portion remaining within the testa is a single cotyledon ; that which lengthens, producing radicles from within its point, is the cauliculus and radicle ; and the thread-like protruded green body is the plumula. If this is compared with the germination of dicotyledons, an obvious difference will be at once perceived in the manner in which the radicles are produced : in monocotyledons they are emitted from within the substance of the radicular extremity, and are actually sheathed at the base by the lips of the pas- sage through which they protrude ; while in dicotyledons they appear at once from the very surface of the radicular extre- mity, and consequently have no sheath at their base. Upon this difference in economy, Richard proposed to substitute the term Endorhizae for monocotyledons, and Exorhizae for dicotyledons. Some consider the former less perfect than the latter ; endorhizae being involute, or imperfectly developed, exorhizae evolute, or fully developed. Dumortier adds to these names endophyllous and exophyllous; because the young leaves of monocotyledons are evolved from within a sheath {coleopliyllum or coleoptilum), while those of dicotyledons are always naked. The sheath at the base of the radicle of monocotyledons is called the coleorhiza by Mirbel. Another form of monocotyledonous embryo is that of Aroideae and their allies, in which the plumula is not so intimately com- bined with the embryo as to be undistinguishable, but is indi- cated externally by a little slit above the base (Plate VI. CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 191 fig. 6. B. e), within which it lies until called into developement by germination. The exceptions to what has been now described ought, like those of dicotyledons, rather to be called remarkable modi- fications. Much stress has been laid upon them by several writers, who have thought it requisite to give particular names to their parts. To me, however, it appears far more advisable to explain their analogies without the unnecessary creation of new and bad names. In Graminece (Plate VI. fig. 4.) the embryo consists of a lenticular body lying on the outside of the base of the albumen on one side, and covered on its inner face by that body, and on its outer face by the testa : if viewed on the face next the testa, a slit will be observed of the same nature as that in the side of the embryo of Aroideee ; opening this cleft a small conical projection is discovered, pointing towards the apex of the seed. If the embryo be then divided vertically through the conical projection, it will be seen that the latter (c) is a sheath including other little scales resem- bling the rudiments of leaves ; that that part of the embryo which lies next the albumen (d), and above the conical body, is solid ; and that the lower extremity of the embryo (e) contains within it the indication of an internal radicle, as in other monocotyledons. In this embryo it is to be understood that the conical projection is the plumul 'a ,• that part of the embryo lying between it and the albumen, a single scutelliform coty- ledon ; and the lower point of the embryo, the radicle. In wheat there is a second small cotyledon on the outside of the embryo, inserted a little lower down than the scutelliform cotyledon. This last is called scutellwn by Gaertner, who considered it of the nature of vitellus. The late M. Richard considered the scutelliform cotyledon a particular modification of the radicle, which he called hypoblastus ; the plumula a form of cotyledon, called blastns ; the anterior occasional cotyledon a peculiar appendage, named epiblastus ; and the radicle a pro- tuberance of the cauliculus, called radiadoda. He further, in reference to this peculiar opinion, termed embryos of this description macropodous. In these ideas, however, Richard was manifestly wrong, as is now well known. From what has been stated, it is apparent that dicoty- 192 ORGANOGRAPHY. BOOK I. ledons are not absolutely characterised by having two coty- ledons, nor monocotyledons by having only one. The real distinction between them consists in their endorhizal or exorhizal germination, and in the cotyledons of dicotyledons being opposite or verticillate, while they are in monocoty- ledons solitary or alternate. Some botanists have, therefore, recommended the substitution of other terms in lieu of those in common use. M. Cassini suggests isodynamous or iso- brious for dicotyledons, because their force of developement is equal on both sides ; and anisodynamous or anisobrious for monocotyledons, because their force of developement is greater on one side than on the other. Another writer, M. Lestiboudois, would call dicotyledons exoj)tiles, because their plumula is naked ; and monocotyledons endoptiles, because their plumula is enclosed within the cotyledons ; but there seems little use in these proposed changes, which are moreover as open to objections as the terms in common use. The Acotyledonous embryo is not exactly, as its name seems to indicate, an embryo without cotyledons ; for, in that case, Cuscuta would be acotyledonous. On the contrary, it is an embryo, which does not germinate from two fixed in- variable points, namely, the plumula and the radicle, but indifferently from any point of the surface; as in some of the Arum tribe, and in all flowerless plants. For further illustrations of the embryo, consult Plate VI. and the explanation of its figures. The direction of the embryo is either absolute or relative. Its absolute direction is that which it has independently of the parts that surround it. In this respect it varies much in different genera ; it is either straight (Plate VI. fig. 5.), ar- cuate (fig. 9.), or falcate, uncinate, or coiled up (fig. 8.) (cy- clical), folded up, spiral (fig. 19.), or bent at right angles (Plate V. fig. 28.) (gromonical, Link), serpentine, or in figure like the letter S (sigmoid). Its relative position is determined by the relation it bears to the chalaza and micropyle of the seed ; or, in other words, upon the relation that the integuments, the raphe, chalaza, hilum, micropyle, and radicle bear to each other. If the sacs of the ovulum are in no degree inverted, but have their com- CHAP. II. COMPOUND ORGANS IN FLOWERING PLANTS. 193 mon point of origin at the hilum, there being (necessarily) neither raphe nor chalaza visible, the radicle will in that case be at the extremity of the seed most remote from the hilum, and the embryo inverted with respect to the seed, as in Cistus, Urtica, and others, where it is said to be antilropal. But if the ovulum undergoes that remarkable extension of one side already described in speaking of that organ, in which the sacs are so inverted that their orifice is next the hilum, and their base at the apex of the ovulum, then there will be a raphe and chalaza distinctly present; and the radicle will, in the seed, be at the end next the hilum, and the embryo will be erect with respect to the seed, or orthotropal, as in the apple, plum, &c. On the other hand, supposing that the sacs of the embryo suffer only a partial degree of inversion, so that their foramen is neither at the one extremity nor the other, there will be a chalaza and a short raphe ; and the radicle will point neither to the apex nor to the base of the seed, but the embryo will lie, as it were, across it, or be heterotropal, as is the case in the primrose. When an embryo is so curved as to have both apex and radicle presented to the hilum, as in Reseda, it is amphitropal. In the words of Gsertner, an embrvo is ascending when its apex is pointed to the apex of the fruit ; descending, if to the base of the fruit ; centripetal, if turned towards the axis of the fruit ; and centrifugal, if towards the sides of the fruit : those embryos are called wandering, or vagi, which have no evident direction. The cotyledons are generally straight, and placed face to face ; but there are numberless exceptions to this. Some are separated by the intervention of albumen (Plate VI. fig. 11.); others are naturally distant from each other without any intervening substance. Some are straight, some waved, others arcuate or spiral. When they are folded with their back upon the radicle, they are called incumbent ,- if their edges are presented to the same part, they are accumbent : terms chiefly used in speaking of Cruciferae. 194 ORGANOGRAPHY. BOOK I. 15. Of Naked Seeds. By naked seeds has been understood, by the school of Lin- naeus, small seed-like fruit, like that of Labiatae, Boraginea?, grasses, and Cyperaceae. But as these are distinctly covered by pericarpia, as has been shown above, the expression in the sense of Linnaeus is obviously incorrect, and is now abandoned. Hence it has been inferred that there is no such thing in existence as a naked seed ; that is to say, a seed which bears on its own integuments the organ of impregnation. To this proposition botanists had assented till the year 1825, when Mr. Brown demonstrated the existence of seeds strictly naked ; that is to say, from their youngest state destitute of pericarpium, and receiving impregnation through their inte- guments without the intervention of style or stigma, or any stigmatic apparatus. That learned botanist has demonstrated that seeds of this description are uniform in Coniferae and Cy- cadeae, in which no pericarpial covering exists. But we have no knowledge at present of such an economy obtaining in other plants, as a constant character. It does however happen, as the same observer has pointed out, that in particular species the ovarium is ruptured at an early period by the ovula, which thus, when ripe, become truly naked seeds ; re- markable instances of which occur in Ophiopogon spicatus, Leontice thalictroides, and Peliosanthes Teta* 195 CHAPTER III. OF THE COMPOUND ORGANS IN FLOWERLESS PLANTS. "We have now passed in review all the different organs which exist in the most perfectly formed plants ; that is to say, in those whose reproduction is provided for by the complicated apparatus of sexes and of fertilising organs. Let us next proceed to consider those lower tribes, some of which are scarcely distinguishable from animals, where there is no evi- dent trace of sexes, in which nothing constructed like seeds is to be detected, and which seem to have no other provision made for the perpetuation of their races than a dissolution of their cellular system. In what I may have to say about them, I shall not, however, do any thing more than give a mere enumeration and description of their organs. All speculative considerations are in this case left out of view : those who wish to be informed upon such points may consult the " Introduction to the Natural System of Botany." 1. Ferns. Filices, or ferns, are plants consisting of a number of leaves or fronds, as they are called, attached to a stem which is either subterraneous or elongated above the ground, some- times rising like a trunk to a considerable height. They are the largest of known vegetables in which no organs of fructi- fixation analogous to those of phaenogamous plants have been discovered. Their petioles, or stipes (rac/iis, W. ; peridroma, Necker), consist of sinuous strata of indurated, very compact, woody fibre, connected by cellular tissue ; and the wood of those which have arborescent trunks is formed by the cohe- sion of the bases of such petioles round a hollow or solid cellular axis. The organs of reproduction are produced from the back or under side of the fronds. In Poli/podiacece, or o 2 196 ORGANOGRAPHY. BOOK I. what are more commonly called dorsiferous ferns, they ori- ginate, either upon the cuticle or from beneath it, in the form of spots at the anastomoses, margins, or extremities of the veins. As they increase in growth they assume the appear- ance of small heaps of granules, called sorts if examined beneath the microscope these granules, commonly called cap- sules, thecce, or conceptacles, are found to be little brittle com- pressed bags formed of cellular membrane, partially sur- rounded by a thickened longitudinal ring {gyrus, annidus, gyroma), which at the vertex loses itself in the cellularity of the membrane, and at the base tapers into a little pedicel ; the thecae burst with elasticity by aid of their ring, and emit minute particles named sporules, from which new plants are produced; as from seeds, in vegetables of a higher order. Interspersed with these thecae are often intermixed articulated hairs ; and, in those genera in which the thecae originate beneath the cuticle, the sori, when mature, continue covered with the superincumbent portion of the cuticle, which is then called the indusium or involucnim {membramda, Necker; glan- dules squamosa:, Guettard). In Trichomanes and Hymeno- phyllum the thecae are seated within the dilated cup-like extremities of the lobes of the frond, and are attached to the vein which passes through their axis, which is then called their receptacle. In another tribe, called Gleichenece, the thecae have a transverse complete, instead of a vertical incom- plete ring, and they are nearly destitute of pedicels ; in a third tribe the sori occupy the whole of the under surface of the frond, which becomes contracted, and wholly alters its appear- ance : the thecae have no ring, and the cellular tissue of their membrane is not reticulated, but radiates regularly from the apex. In these it has been in vain endeavoured to discover traces of organs of fecundation. Nevertheless, as it was difficult for sexualists to believe that plants of so large a size were desti- tute of such organs, it has been considered indispensable that they should be found ; and, accordingly, while all seem to agree in considering the thecae as female organs, a variety of other parts have been dignified by the title of male organs : thus, Micheli and Hedwig found them in certain stipitate CHAP. III. COMPOUND ORGANS IN FLOWERLESS PLANTS. 197 glands of the frond ; Stashelin, Hill, and Schmidel, in the elastic ring ; Koelreuter, in the indusium ; Gleichen, in the stomata ; and Von Martius, in certain membranes enclosing the spiral vessels. None of these opinions are now adopted. In Ophioglosseae, a remarkable tribe of ferns, the fertile frond is rolled up in two lines parallel with its axis or midrib, and at maturity opens regularly by transverse valves along its whole length, emitting a fine powder, which, when magni- fied, is found to consist of particles of the same nature as the sporules found in the thecae of other ferns : here there are no thecae, the metamorphosed frond probably performing their functions. Such is my view of the structure of Ophioglosseae ; but by other botanists it is described as a dense spike of tWo- valved capsules, dehiscing transversely. 2. EquisetacecB. In these the organs of reproduction are arranged in an amentum, consisting of scales bearing on their lower surface an assemblage of cases, called t/iecce, follicidi, or i?ivolucrai which dehisce longitudinally inwards. In these thecae are contained two sorts of granules ; the one very minute and lying irregularly among a larger kind, each of which is wrapped in two filaments, fixed by their middle, rolled spir- ally, having either extremity incrassated, and uncoiling with elasticity. By Hedvvig the apex of the larger granules was supposed to be a stigma, and the thickened ends of the fila- ments anthers, the small granules being the pollen. At any rate it is certain that the larger granules, round which the elastic filaments are coiled, are the reproductive particles. 3. Lycopodiaceoe . These are leafy plants with the habit of gigantic mosses. Their leaves and stem have the same structure as those plants, except that the former are sometimes provided with stomata, and the latter with vessels. Their organs of reproduction are of two kinds : the one kidney-shaped two-valved cases, called thecce, conceptacles, or capsules, destitute of internal divisions, and filled with minute pow is a cup-like dilatation of the Podetium, bearing shields on its margin. 12. Soredia (globtdi, glomeruli), are heaps of powdery bodies lying upon any part of the surface of the thallus ; the bodies of which the soredia are composed are called conidia by Link, and propagula by others. 13. Cystula, or Cistella, a round closed apothecium, filled with sporules, adhering to filaments which are arranged like rays around a common centre, as in Sphaerophoron. 14. Pidvinuli, are spongy, excrescence-like bodies, sometimes rising from the thallus, and often resembling minute trees, as in Parmelia glomulifera. Greville. 15. Cyp/iellce, are pale tubercle-like spots on the under surface of the thallus, as in Sticta. Grev. 16. Lacunce, are small hollows or pits on the upper surface of the thallus. Grev. 206 ORGANOGRAPHY. BOOK I. 17. Nucleus pr oli gems, is a distinct cartilaginous body, coming out entire from the Apothecia, and containing the spo- rules. Grev. 18. Lamina proligcra, is a distinct body containing the sporules, separating from the apothecia, often very convex and variable in form, and mostly dissolving into a gela- tinous mass. Grev. 19. Fibrillar, are the roots. 20. Excipuhis, is that part of the thai! us which forms a rim and base to the shields. 21. Nucleus, is the disk of the shield which contains the sporules and their cases. 22. Asci, are tubes, in which the sporules are contained while in the nucleus. 23. Thallodes, is an adjective used to express an origin from the thallus: thus, margo thallodes signifies a rim formed by the thallus, excipulus thallodes a cup formed by the thallus. 24. Lorulum, is used by Acharius to express a filamentous, branched thallus. 25. Crusta is a brittle crustaceous thallus. 26. Gongyli, are the granules contained in the shields, and have been thought to be the sporules by which lichens are propagated : but this is doubted by Agardh. 8. Algce and Characece. These, with fungi, constitute the lowest order of vegetable developement : they vary in size from mere microscopic ob- jects to a large size, and are composed of cellular tissue in various degrees of combination ; some are even apparently ani- mated, and thus form a link between the two great kingdoms of organised matter. Their sporules are either scattered through the general mass of each plant, or collected in cer- tain places which are more swollen than the rest of the stem, and sometimes resemble the pericarpia of perfect plants. The terms used in speaking of the parts of Algae are the following : — CHAP. III. COMPOUND ORGANS IN FLOWERLESS PLANTS. 207 1. Gongylits ; a round hard body, which falls off the mother plant, and produces a new individual : this is found in Fuci. w. 2. Thallns ; the plant itself. 3. Apothecia ; the cases in which the organs of reproduction are contained. 4s Peridiola, Fr. ; the membrane by which the sporules are immediately covered. 5. Gramda ; large sporules, contained in the centre of many Algae ; as in Gloionema of Greville. Crypt. Jf.. 6. 30. 6. Pseudoperithecium j "i terms used by Fries to express such 7. Pseudokymeniwn i J- coverings of Sporidia as resemble 8. Pseudoperidhim ; J in figure the parts named peri- thecium, hymenium, and peridium in other plants: see those terms. 9. Sporidia; granules which resemble sporules, but which are of a doubtful nature. It is in this sense that Fries declares that he uses the word : vide Plant, homonom. p. 294. 10. Phycomater, Fries; the gelatine in which the sporules of Byssaceae first vegetate. 11. VesiculcE; inflations of the thallus, filled with air, by means of which the plants are enabled to float. 12. Hupha, Willd. ; the filamentous, fleshy, watery thallus of Byssacese. 13. Nucula ; one of the apothecia of Characeas; described by Greville to be a sessile, oval, solitary, spirally striated body, with a membranous covering, and the summit indistinctly cleft into five segments, containing sporules. 14. Globules; the second organ of Characea? ; the excellent observer last quoted describes it as a minute round body of a reddish colour, composed externally of a number of triangular (always ?) scales, which separate, and produce its dehiscence. The interior is filled with a mass of elastic, transversely undulated filaments. The scales are composed of radiating hollow tubes, partly filled with minute coloured granules, which freely escape when the tubes are injured : their nature is wholly unknown, and, I believe, hitherto unnoticed. 208 ORGANOGRAPHY. BOOK I» 15. Coniocystct; tubercle-like closed apothecia, containing a mass of sporules. 9. Fungi. The structure of these plants is yet more simple than that of Algae, consisting of little besides cellular tissue, among which sporules lie scattered. Some, of the lowest degree of developement, are composed only of a few cellules, of which one is larger than the rest, and contains the sporules; others are more highly compounded, consisting of myriads of cellules, with the sporules lying in cases, or asci. Notwithstanding the extreme simplicity of these plants, writers upon fungi have contrived to multiply the terms relating to them in a remark- able manner. The following are all with which I am ac- quainted : — 1. The pileus, or cap, is the uppermost part of the plant of an Agaricus, and resembles an umbrella in form. 2. The stipes, is the stalk that supports the pileus. 3. The volva, or wrapper, is the involucrum-like base of the stipes of Agaricus. It originally was a bag enveloping the whole plant, and was left at the foot of the stipes when the plant elongated and burst through it. 4. The velum, or veil, is a horizontal membrane, connecting the margin of the pileus with the stipes : when it is adnate with the surface of the pileus, it is a velum universale : when it extends only from the margin of the pileus to the stipes, it is a velum partiale. 5. The annulus, is that part of the veil which remains next the stipes, which it surrounds like a loose collar. 6. Cortina, is a name given to a portion of the velum which adheres to the margin of the pileus in fragments. 7. The hymenium, is the part in which the sporules imme- diately lie ; in Agaricus, it consists of parallel plates, called lamellce, or gills; these are adnate with the stipes, when the end next it cohere with it: when they are adnate, and at the same time do not terminate abruptly at the stipes, but are carried down it more or less, they are decurrent ; if they do not adhere to the stipes, they are said to he free. CHAP. III. COMPOUND ORGANS IN FLOWEIILESS PLANTS. 209 8. Stroma, is a fleshy body to which flocci are attached ; as in Isaria and Cephalotrichum. 9. Flood, are woolly filaments found mixed with sporules in the inside of many Gastromyci. The same name is also applied to the external filaments of Byssaceae. 10. Orbicidus, is a round flat hymenium contained within the peridium of some fungi ; as Nidularia. W. 11. Nucleus, is the central part of a perithecium. 12. Sporangium, is the external case of Lycoperdon and its allies. 13. Sporangiola, are cases containing sporidia. 14. Perithecium, is a term used to express the part which contains the reproductive organs of Sphaeria and its co-ordinates. 15. Peridium, is also a kind of covering of sporidia; peridio- Iujh is its diminutive. 16. Ostiolum, is the orifice of the perithecium of Sphaeria. 17. Sphcerula, is a globose peridium, with a central opening, through which sporidia are emitted, mixed with a gela- tinous pulp. 18. Capillitium, is a kind of purse or net, in which the sporules of some fungi are retained ; as in Trichia. W. 19. Trichidium, or pecten, is a tender, simple, or sometimes branched hair, which supports the sporules of some fungi; as Geastrum. W. 20. Asci, are the tubes in which the sporidia are placed; ascelli or thecce are the same thing. 21. Sporidia, are the immediate covering of sporules; spo- ridiola are sporules. 22. Thallus, or thalamus, is the bed of fibres from which many fungi arise. 23. Mycelia, are the rudiments of fungi, or the matter from which fungi are produced. 210 BOOK II. PHYSIOLOGY J OR, PLANTS CONSIDERED IN A STATE OF ACTION. GENERAL CONSIDERATIONS. We have thus far considered plants as inert bodies, having certain modifications of structure, and formed upon a plan, the simplicity and uniformity of which is among the most beautiful proofs of the boundless power and skill of the Deity. Our next business is to enquire into the nature of their vital actions, and to consider those phenomena in which the analogy that undoubtedly exists between plants and animals is most striking; in a word, to make ourselves acquainted with the exact nature of the laws of vegetable life. In explaining these things, it is not my purpose to notice all the different speculations that ingenious men have from time to time brought forward : for this would be incompatible with the plan of my work, and would be far more curious than useful. On the contrary, I propose, in the first place, to give a summary exposition of the principal phenomena of vegeta- tion, and then to support the statement by a detailed account of the more important proofs of all doubtful points. I am the more anxious that this should be understood, because I know how prone the world is to misconstruction : I therefore beg it to be remembered, that when particular opinions are here passed over in silence, it is because I do not think them sufficiently proved to be recorded consistently with the plan I have prescribed to myself. If we place a seed, — that of an apple, for instance, — in earth at the temperature of 32° Fahr., it will remain inactive till it finally decays. But if it is placed in moist earth above BOOK II. GENERAL CONSIDERATIONS. 211 the temperature of 32°, and screened from the action of light, its integument gradually imbibes moisture and swells, oxygen is absorbed, carbonic acid expelled, and the vital action of the embryo commences. It elongates downwards by the radicle, and upwards by the cotyledons; the former pene- trating the soil, the latter elevating themselves above it, acquiring a green colour by the deposition of carbon absorbed from the atmosphere in the light, and unfolding in the form of two opposite roundish leaves. This is the first stage of vege- tation : the young plant consists of little more than cellular tissue ; only an imperfect developement of vascular and fibrous tissue being discoverable, in the form of a sort of cylinder of bundles, lying just in the centre. The part within the cylin- der, at its upper end, is now the medulla, without it the bark ; while the cylinder itself is the preparation for the medullary sheath, and consists of vertical fibres passing through cellular tissue, which separates them horizontally in every direction. The young root is now absorbing from the earth its nutri- ment, which passes up to the summit of the plant by the cellular substance of the medulla, and is thence impelled into the cotyledons, where it is aerated and evaporated : such of it as is not fixed in the cotyledons passes down through the bark into the root. Forced onwards by the current of sap, which is continually impelled upwards from the root, the plumula next ascends in the form of a little twig, at the same time sending roots in the form of fibres downwards in the centre of the radicle, which become the earliest portion of wood that is deposited : these fibres, by their action, now compel the root to emit little rami- fications. Previously to the elongation of the plumula its apex has acquired the rudimentary state of a leaf: this continues to develope as the plumula elongates, until, when the first inter- nodium of the latter ceases to lengthen, the leaf has actually arrived at its complete formation. When fully grown it repeats in a much more perfect manner the functions previously per- formed by the cotyledons : it aerates the sap that it receives, and returns the superfluous portion of it downwards through p 2 212 PHYSIOLOGY. BOOK II. the bark to the root ; it also sends fibres down between the medullary sheath and the bark, thus forming the first stratum of wood in the new stem. During these operations, while the plumula is ascending, its leaf forming and acting, and the woody matter created by it descending, the cellular tissue of the stem is forming, and expanding horizontally to make room for the new matter forced into it ; so that developement is going on simultaneously both in a horizontal and perpen- dicular direction. This process may not inaptly be compared to that of weaving, the warp being the perpendicular, and the weft the horizontal, formation. In order to enable the leaf to perform its functions of aeration completely it is traversed by veins originating in the medulla, and has delicate evaporating pores (stomata), which communicate with a highly complex pneumatic system that extends to almost every part of the plant. After the production of its first leaf by the plumula, others are successively produced around the axis at its elongating point, all constructed alike, connected with the stem or axis in the same manner, and performing precisely the same func- tions as have been just described. At last the axis ceases to elongate; the old leaves gradually fall off; the new leaves, instead of expanding after their formation, retain their rudi- mentary condition, harden, and fold over one another, so as to be a protection to the delicate point of elongation ; or, in other words, become the scales of a bud. We have now a shoot with a woody axis, and a distinct pith and bark ; and of a more or less conical figure. At the axilla of every leaf a bud had been generated during the growth of the axis ; so that the shoot, when deprived of its leaves, is covered from end to end with little, symmetrically arranged, projecting points, which are the buds. The cause of the figure of the perfect shoot being conical is, that, as the wood originates from the base of the leaves, the lower end of the shoot, which has the greatest number of strata, because it has the greatest number of leaves above it, will be the thickest; and the upper end, which has had the fewest leaves to distend it by their deposit, will have the least diameter. Thus that part of the stem which has two leaves above it will have wood formed by two BOOK II. GENERAL CONSIDERATIONS. 213 successive deposits ; that which has nine leaves above it will have wood formed by nine successive deposits ; and so on : while the extreme vital point, as it can have no deposit of matter from above, will have no wood, the extremity being merely covered by the rudiments of leaves hereafter to be developed. If at this time a cross section be examined, it will be found that the interior is no longer imperfectly divided into two portions, namely, medulla and skin, as it was when first ex- amined in the same way, but that it has distinctly two, internal, perfect, concentric lines, the outer indicating a separation of the bark from wood ; and the inner, a separation of the wood from the medulla : the latter too, which in the first observ- ation was fleshy, and saturated with humidity, is become dis- tinctly cellular, and altogether or nearly dry. With the spring of the second year and the return of warm weather vegetation recommences. The uppermost, and perhaps some other, buds which were formed the previous year gradually unfold, and pump up sap from the stock remaining in store about them ; the place of the sap so removed is instantly supplied by that which is next it ; an impulse is thus given to the fluids from the summit to the roots ; new sap is absorbed from the earth, and sent upwards through the wood of last year ; and the phenomenon called the flow of the sap is fully completed, to continue with greater or less velocity till the return of winter. The axis of the buds elongates upwards, forming leaves and buds in the same way as the parent shoot : in like manner also each bud sends down its roots, in the form of fibres within the bark and above the wood of the shoot from which it sprang ; thus forming on the one hand a new layer of wood, and on the other a fresh deposit of bark. In order to facilitate this last operation, the old bark and wood are separated in the spring by the exudation from both of them of the glutinous, slimy substance called cambium ; which appears to be expressly intended, in the first instance, to facilitate the descent of the subcortical fibres of the growing buds ; and, in the second place, to generate the cellular tissue by which the horizontal dilatation of the axis is caused, and which maintains a communication between the p 3 214 PHYSIOLOGY. BOOK II. bark and the centre of the axis. These lines of communica- tion have, by the second year, become sufficiently developed to be readily discovered, and are in fact the medullary rays spoken of in the last book. It will be remembered that there was a time when that which is now bark constituted a homogeneous body with the medulla; and that it was after the leaves began to come into action that the separation which now exists between the bark and medulla took place. At the time when they were indissolubly united they both consisted of cellular tissue, with a few spiral vessels upon the line indicative of future separation. When a deposit of wood was formed from above between them they were not wholly divided the one from the other, but the deposit was effected in such a way as to leave a communication by means of cellular tissue between the bark and the medulla ; and, as this formation is at all times coaetaneous with that of the wood, the communication so effected between the medulla and bark is quite as perfect at the end of the third year as it is at the beginning of the first; and so it will continue to be to the end of the growth of the plant. The sap which has been sucked into circulation by the unfolding leaves is exposed, as in the previous year, to the effect of air and light ; is then returned through the petiole to the stem, and sent downwards through the bark, to be from it either conveyed to the root, or distributed horizon- tally by the medullary rays to the centre of the stem. At the end of the year the same phenomena occur as took place the first season : wood is gradually deposited by slower degrees, whence the last portion is denser than the first, and gives rise to the appearance called the annual zones : the new shoot or shoots are prepared for winter, and are again elongated cones, as was the first ; and this latter has acquired an increase in diameter proportioned to the quantity of new shoots which it produced, new shoots being to it now what young leaves were to it before. The third year all that took place the year before is repeated : sap is absorbed by the unfolding leaves ; and its loss is made good by new fluids introduced by the roots and transmitted through the alburnum or wood of the year before ; new wood and liber are deposited by matter sent BOOK II. GENERAL CONSIDERATIONS. 215 downwards by the buds ; cambium is exuded ; the horizontal developement of cellular tissue is repeated, but more exten- sively ; wood towards the end of the year is formed more slowly, and has a more compact character ; and another ring appears indicative of this year's increase. In precisely the same manner as in the second and third years of its existence will the plant continue to vegetate, till the period of its decay, each successive year being a repetition of the phenomena of that which preceded it. After a certain number of years the tree arrives at the age of puberty : the period at which this occurs is very uncer- tain, depending in some measure upon adventitious circum- stances, but more upon the idiosyncrasy, or peculiar constitu- tion of the individual. About the time when this alteration of habit is induced, by the influence of which the sap or blood of the plant is to be partially directed from its former courses into channels in which its force is to be applied to the pro- duction of new individuals rather than to the extension of itself; — about this time it will be remarked that certain of the young branches do not elongate, as had been heretofore the wont of others, but assume a short stunted appearance, probably not growing two inches in the time which had been previously sufficient to produce twenty inches of increase. Of these little stunted branches, called spurs, the terminal bud acquires a swollen appearance, and at length, instead of giving birth to new leaves, produces from its bosom a cluster of flower-buds, or alabastra, which had been enwrapped and protected from injury during the previous winter by several layers of imperfect leaves, now brought forth as bracteae. Sap is impelled into the calyx through the pedicel by gentle de- grees, is taken up by it, and exposed by the surface of its tube and segments to air and light ; but having very imperfect means of returning, all that cannot be consumed by the calyx is forced onwards into the circulation of the petals, stamens, and pistillum. The petals unfold themselves of a dazzling white tinged with pink, and expose the stamens ; at the same time the disk changes into a saccharine substance, which nourishes the stamens and pistillum, and gives them energy to perform their functions. p 4 216 PHYSIOLOGY. BOOK II. At a fitting time, the stigmatic surface of the pistillum being ready to receive the pollen, this is injected upon it from the anthers, which have remained in approximation to it for that particular purpose. When the pollen touches the stigma, the grains adhere firmly to it by means of its viscid surface, then emit a delicate membranous tube, which pierces into the stigmatic tissue, lengthens there, and conveys the vivifying matter contained in the pollen towards the ovula, which it finally enters by means of their foramen. This has no sooner oc- curred than the petals and stamens fade and fall away, their ephemeral but important functions being accomplished. All the sap which is afterwards impelled through the peduncle can only be disposed of to the calyx and ovarium, where it lodges : both these swell and form a young fruit, which con- tinues to grow as long as any new matter of growth is supplied from the parent plant. After a certain period the juices of the fruit cease to be increased by the addition of new matter, its surface performs the functions of leaves in exposing the juice to light and air ; finally the surface loses its green colour, assumes the rich, ruddy glow of maturity; the juices cease to be influenced by light ; the peduncle is no longer a passage for fluids, but dries up and becomes unequal to supporting the fruit, which at last falls to the earth. Here, if not destroyed by animals, it lies and decays : in the succeeding spring its seeds are stimulated into life, strike root in the mass of decayed matter that surrounds them, and spring forth as new plants to undergo all the vicissitudes of their parent. Such are the progressive phenomena in the vegetation, not only of the apple, but of all trees that are natives of northern climates, and of a large part of the herbage of the same coun- tries,— modified, of course, by peculiarities of constitution, as in annual and herbaceous plants, and in those the leaves of which are opposite and not alternate ; but all the more essential cir- cumstances of their growth are the same as those of the apple tree. If we reflect upon these phenomena, our minds can scarcely fail to be deeply impressed with admiration at the perfect sim- plicity and, at the same time, faultless skill with which all the BOOK II. GENERAL CONSIDERATIONS. 21? machinery is contrived upon which vegetable life depends. A few forms of tissue, interwoven horizontally and perpendicularly, constitute a stem ; the developement, by the first shoot that the seed produces, of buds which grow upon the same plan as the first shoot itself, and a constant succession of the same phe- nomenon, causes an increase in the length and breadth of the plant ; an expansion of the bark into a leaf, within which ramify veins proceeding from the seat of nutritive matter in the new shoot, the provision of air-passages in its sub- stance, and of evaporating pores on its surface, enables the crude fluid sent from the roots to be elaborated and digested until it becomes the peculiar secretion of the species; the contraction of a branch and its leaves forms a flower ; the disintegration of the internal tissue of a petal forms an anther; the folding inwards of a leaf is sufficient to constitute a pistil- lum ; and, finally, the gorging of the pistillum with fluid which it cannot part with causes the production of a fruit. In hot latitudes there exists another race of trees, of which palms are the representatives, and in the north there are many herbs, in which growth, by addition to the outside, is wholly departed from, the reverse taking place; that is to say, their diameter increasing by addition to the inside. As the seeds of such plants are formed with only one cotyledon, they are called monocotyledonous ; and their growth being from the inside, they are also named endogenous. In these plants the functions of the leaves, flowers, and fruit are in no- wise different from those of dicotyledons ; their peculiarity consisting only in the mode of forming their stems. When a monocotyledonous seed has vegetated it usually does not disentangle its cotyledon from the testa, but simply protrudes the radicle; the cotyledon swelling, and remaining firmly encased in the seminal integuments. The radicle shoots downwards to become root ; and afterwards a leaf is emitted from the side of the collum, which elongated at the same time as the radicle. This first leaf is succeeded by another facing it, and arising from its axilla ; the second produces a third facing it, and arising also from its axilla; and, in this manner, the production of leaves continues, until the plant, if caules- 218 PHYSIOLOGY. BOOK II. cent, is ready to produce its stem. Up to this period no stem having been formed, it has necessarily happened that the bases of the leaves hitherto produced have been all upon the same plane ; and as each has been produced from the bosom of the other without any such intervening space, as occurs in dicoty- ledonous plants, it would have been impossible for the matter of wood, if any had been formed, to be sent downwards around the circumference of the plant : it would, on the contrary, have been necessarily deposited in the centre. In point of fact, how- ever, no deposit of wood like that of dicotyledons takes place, either now or hereafter. The union of the bases of the leaves has formed a fleshy stock, cormus, or plate, which, if ex- amined, will be found to consist of a mass of cellular tissue, traversed by perpendicular bundles of vascular tissue and woody fibre, taking their origin in the veins of the leaves, of which they are manifest prolongations downwards ; and there is no trace of bark, medullary rays, or central pith : the whole body being a mass of pith, woody fibre, and vascular tissue mixed together. To understand this formation yet more clearly, consider for a moment the internal structure of the petiole of a dicotyledon : it is composed of a bundle or bundles of vascular tissue encased in woody fibre, surrounded on all sides with pith, or, which is the same thing, parenchyma. Now suppose a number of these petioles to be separated from their laminae, and to be tied in a bunch parallel with each other, and, by lateral pressure, to be squeezed so closely toge- ther that their surfaces touch each other accurately, except at the circumference of the bunch. If a transverse section of these be made, it will exhibit the same mixture of bundles of woody fibre and parenchyma, and the same absence of dis- tinction between medulla, wood, and bark, which has been noticed in the cormus, or plate, of monocotyledons. As soon as the plate has arrived at the necessary diameter it begins to elongate upwards, leaving at its base those leaves that were before at its circumference, and carrying upwards with it such as occupied its centre ; at the same time, new leaves continue to be generated at the centre, or, as it must now be called, at the apex of the shoot. As fresh leaves are developed, they thrust aside to the cir- BOOK II. GENERAL CONSIDERATIONS. 219 cumference those which preceded them, and a stem is by degrees produced. Since it has not been formed by additions made to its circumference by each successive leaf, it is not conical, as in dicotyledons; but, on the contrary, as its in- crease has been at the centre, which has no power to extend its limits, being strictly confined by the circumference which, when once formed, does not afterwards materially alter in dimensions, it is, of necessity, cylindrical : and this is one of the marks by which a monocotyledon is often to be known in the absence of other evidence. The centre being but little acted upon by lateral pressure, it remains loose in texture, and, until it becomes very old, does not vary much from the density acquired by it shortly after its formation ; but the tissue of the circumference being continually jammed together by the pressure outwards of the new matter formed in the centre, in course of time becomes a solid mass of woody matter, the cellular tissue once intermingled with it being almost obliterated, and appearing among the bundles it formerly surrounded, like the interstices around the minute pebbles of a mosaic gem. Such is the mode of growth of palms, and of a great pro- portion of arborescent monocotyledons. But there are others in which this is in some measure departed from. In the common asparagus the shoots produce a number of lateral buds, which all develope and influence its form, as the buds of dicotyledons ; so that the cylindrical figure of monocotyle- dons is exchanged for the conical ; its internal structure is strictly endogenous. In grasses a similar conical figure pre- vails, and for the same reason ; but they have this additional peculiarity, that their stem, in consequence of the great rapi- pidity of its growth, is fistular, with transverse phragmata at its nodi. It is not certain whether the subsequent internal growth of the stem is ever sufficient to fill up the central cavity ; but, from a specimen of a bamboo in my possession, I incline to think that the lower part of grass stems does some- times become filled up with solid matter. Upon one or other of the two plans now explained are all flowering plants developed ; but in flowerless plants it is dif- ferent. In arborescent ferns the stem consists of a cylinder of 220 PHYSIOLOGY. BOOK II. hard sinuous plates connected by parenchyma, and surround- ing a hollow axis, which sometimes becomes filled up with solid matter. It would seem, in these plants, as if the stem consisted of a mere adhesion of the petioles of the leaves in a single row ; but we are, at present, too little acquainted with them in a living state to form any fixed opinion upon the subject. In mosses and some Hepaticae the stem seems to consist of nothing but an axis formed of the united bases of the leaves ; and their growth may be considered analogous to that of an annual shoot of a dicotyledon without its wood. The re- mainder of flowerless plants are principally mere horizontal expansions of cellular tissue, analogous to nothing that is known in the other parts of the vegetable kingdom. 221 CHAPTER I. ELEMENTARY ORGANS. That of these the cellular tissue is the most important is apparent by its being the only one of the elementary organs that is uniformly present in plants ; and by its being the chief constituent of all those compound organs that are most essen- tial to the preservation of species. It transmits fluids in all directions. In most cellular plants no other tissue exists, and yet there a circulation of sap takes place ; it constitutes the whole of the medullary rays, convey- ing the elaborated juices from the bark towards the centre of the stem ; all the parenchyma in which the sap is diffused upon entering the leaf, and by which it is exposed to evapor- ation, light, and atmospheric action, consists of cellular tissue ; nearly all the bark in which the descending current of the sap takes place is also composed of it; and in endogenous plants, where no bark exists, there appears to be no other route that the descending sap can take than through the cel- lular substance in which the vascular system is imbedded. It is, therefore, readily permeable to fluid, although it has no visible pores. In all cases of wounds, or even of the developement of new parts, cellular tissue lis Jirst generated : for example, the granulations that form at the extremity of a cutting when imbedded in earth, or on the lips of incisions in the wood or bark ; the extremities of young roots ; scales, which are gene- rally the commencement of leaves ; pith, which is the first part created when the stem shoots up ; nascent stamens and pistilla ; ovula ; and, finally, many rudimentary parts ; — all these are at first, or constantly, formed of cellular tissue alone. It may be considered the Jlesh of vegetable bodies: the matter which surrounds and keeps in their place all the ramifi- 222 PHYSIOLOGY. BOOK II. cations or divisions of the vascular system is cellular tissue. In this the plates of wood of exogenous plants, the fibres of endogenous plants, the veins of leaves, and, indeed, the whole of the central system of all of them, are either imbedded or enclosed. The action of impregnation appears to take place exclusively through its agency. Pollen is only cellular tissue in a parti- cular state; when it bursts, the vivifying particles it contains are a still more minute state of the same tissue : the coats of the anther are composed entirely of it ; and the tissue of the stigma, through which impregnation is conveyed to the ovula, is merely a modification of the cellular. The ovula themselves, with their sacs, at the time they- receive the vivi- fying influence, are a semitransparent congeries of cellules. It is, finally, the tissue in "which alone amylaceous or sac- charine secretions are deposited. These occur chiefly in tubers, as in the potato and arrow-root; in rhizomata, as in the ginger; in soft stems, such as those of the sago-palm and sugar-cane ; in albumen, as that of corn ; in pith, as in the Cassava; in the disk of the flower, as in Amygdalus; and, finally, in the bark, as in all exogenous plants ; and cellular tissue is the principal, or exclusive, constituent of these. Woody fibre is apparently destined merely for the con- veyance of fluid upwards or downwards, from one end of a body to another, and for giving firmness and elasticity to every part. That it is intended for the conveyance of fluid in particular channels seems to be proved, 1. from its constituting the principal part of all wood, particularly of that which is formed in stems the last in each year, and in which fluid first ascends in the ensuing season ; 2. from its presence in the veins of leaves where a rapid circulation is known to take place, form- ing in those plants both the adducent and reducent channels of the sap ; and, 3. from its passing downwards from the leaves into the bark, thus forming a passage through which the peculiar secretions may, when elaborated, arrive at the stations where they are finally to be deposited. Mr. Knight is clearly of opinion that they convey fluid either upwards or downwards ; in which I fully concur with him : the power of CHAP. I. ELEMENTARY ORGANS. 223 cuttings to grow when inverted seems, indeed, a conclusive proof of this. Dr. Dutrochet, however, endeavours to prove that they merely serve for a downward conveyance. With regard to its giving Jirnmess and elasticity to every pa?i, we need only consider its surprising tenacity, as evinced in hemp, flax, and the like; and its constantly surrounding and protecting the ramifications of the vascular system, which has no firmness or tenacity itself. To this evidence might be added, the admirable manner in which it is combined to answer such an end. It consists, as has been seen, of ex- tremely slender tubes, each of which is indeed possessed of but a slight degree of strength ; but being of different lengths, tapering to each extremity, and overlapping each other in various degrees, these are consolidated into a mass that considerable force is insufficient to break. Any one, who will examine a single thread of the finest flax with a microscope that magnifies 180 times, will find, that that which to the eye appears a single thread, is in reality composed of many distinct fibres. The real nature of the functions of the vascular system has been the subject of great difference of opinion; and may, indeed, be said to be so still. Spiral vessels have been most commonly supposed to be destined for the conveyance of air ; and it seems difficult to conceive how any one accustomed to anatomical observations, and who has remarked their dark appearance when lying in water, can doubt that fact. Nevertheless, many others, and among them Dr. Dutrochet, assert that they serve for the transmission of fluids upwards from the roots. This observer states, that if the end of a branch be immersed in coloured fluid, it will ascend in both the spiral vessels and ducts ; but that in the former it will only rise up to the level of the fluid in which the branch is immersed, while, throuo-h the latter, it will travel into the extremities of the branches. It has, however, been asked with much justice, how the opinion that spiral vessels are the sap-vessels is to be recon- ciled with the fact of their non-existence in multitudes of plants in which the sap circulates freely. To which might have been, or perhaps has been, added the questions, why they do not exist in the wood, where a movement of sap chiefly 224 PHYSIOLOGY. BOOK II. takes place in exogenous trees ? and also, how it happens that their existence is almost constantly connected with the presence of sexes, if they are only sap-vessels ? And further, it has always been remarked, that if a transverse section of a vine, for instance, or any other plant, be put under water, bubbles of air rise through the water from the mouths of the spiral vessels. But then, it has been urged, that coloured fluids manifestly rise in the spiral vessels ; a statement that has been admitted, when the spiral vessels are wounded at the part plunged in the colouring fluid, but denied in other circumstances. Indeed, to any observer acquainted with the difficulty of microscopic investigations, the obscurity that practically surrounds a question of this sort must be apparent enough. The subject has, however, been investigated with much care by Dr. L. W. Theodore Bischoff, who instituted some very delicate and ingenious experiments, for the purpose of deter- mining the real contents and office of the spiral vessels. It is impossible to find room here for a detailed account of his experiments, for which the reader is referred to his thesis, De vera Vasorum Plantarum Spiralium Structura et Functiotie Com- mentatio: Bonnae, 1829. It must be sufficient to state, that, by accurate chemical tests, by the most careful purification of the water employed from all presence of air, and by separating bundles of the spiral vessels of the gourd {Cucurbita Pepo), and of some other plants, from the accompanying cellular substance, he came to the following conclusions, which, if not exactly, are probably substantially, correct : " That plants, like all other living bodies, require, for the support of their vital functions, a free communication with air ; and that it is more especially oxygen, which, when absorbed by the roots from the soil, renders the crude fluid fit for the nourishment and support of a plant, just as blood is rendered fit for that of animals. But, for this purpose, it is not sufficient that the external surface should be surrounded by the atmosphere ; other aeriferous organs are provided, in the form of spiral vessels, which are placed internally, and convey air contain- ing an unusual proportion of oxygen, which is obtained through the root, by their own vital force, from the earth and water. CHAP. I. ELEMENTARY ORGANS. 225 In a hundred parts of this air twenty-seven to thirty parts are of oxygen, which is in part lost during the day by the surface of plants under the direct influence of the solar rays." With such evidence of the aeriferous functions of the spiral vessels it is difficult to contend ; and, indeed, it seems pro- bable that this question is settled as far as spiral vessels, pro- perly so called, are concerned. But there are many vessels abounding in the wood, to which they give a porous appear- ance when cut across, and which I have called ducts, that, although perhaps mere modifications of the spiral vessel, are, nevertheless, so far distinct as to convey air at one period of their existence, and fluid at another. In the vine, for instance, the true spiral vessels of the medullary sheath and of the herb- aceous parts, always filled with air, must be carefully distin- guished from the ducts of the wood ; which are, undoubtedly, filled during the principal flow of the sap with that fluid, although they finally become dry and empty. And it may be further remarked, that the dotted ducts of such plants as Phytocrene gigantea, or water-vine, so well represented by Mr. Griffith in Dr. Wallich's Plantce Asiaticce Rario?rs, are apparently the principal conduits in that curious plant, as they are in Gramineae, and other monocotyledons, of the fluid absorbed from the earth. So that, while true spiral vessels may be admitted as un- doubted vehicles of air, ducts of all kinds, and especially dotted ducts, cannot be doubted to be the passages through which fluid is conveyed when great rapidity is required. I have already stated that, although all these vessels are, in the pre- sent state of our anatomical knowledge, considered as equally belonging to the vascular system, yet that the dotted will rather be referred eventually to the cellular, and then their lymphatic office will be unquestioned. In regard to the functions of air-cells and lacunae, it may be sufficient to remark, that in all cases in which they form a part of the vital system, as in water plants, they are cavities regularly built up of cellular tissue, and uniform in figure in the same species ; while, on the other hand, where they are not essential to vitality, as in the pith of the walnut, the rice- 226 PHYSIOLOGY. BOOK II. paper plant, the stems of Umbelliferae, and the like, they are ragged, irregular distensions of the tissue. In the former case they are intended to enable plants to float in water ; in the latter, they are caused by the growth of one part more rapidly than another. 227 CHAPTER II. OF THE ROOT. It is the business of the root to absorb nutriment from the soil, and to transmit it upwards into the stem and leaves; and also to fix the plant firmly in the earth. Although moisture is, no doubt, absorbed by the leaves of all and the stems of many plants, yet it is certain that the greater part of the food of plants is taken up by the roots; which, hence, are not in- correctly considered vegetable mouths. But it is not by the whole surface of the root that the absorption of nutriment takes place ; it is the spongioles almost exclusively to which that office is confided : and hence their immense importance in vegetable economy, the absolute necessity of preserving them in transplantation, and the certain death that often follows their destruction. This has been proved, in the following manner, by the celebrated Senebier: — he took a radish, and placed it in such a position that the extremity only of the root was plunged in water : it remained fresh several days. He then bent back the root, so that its extremity was curved up to the leaves : he plunged the bent part in water, and the plant withered soon ; but it recovered its former freshness upon relaxing the curvation, and again plunging the extremity of the root into the water. This explains why forest trees, with very dense umbrageous heads, do not perish of drought in hot summers or dry situa- tions, when the earth often becomes mere dust for a consider- able distance from their trunk, in consequence of their foliage turning off the rain: the fact is obviously that the roots near the stem are inactive, and have little or nothing to do as pre- servatives of life except by acting as conduits, while the func- tions of absorption go on through the spongioles, which, being at the extremities of the roots, are placed beyond the influence of the shadow, and extend wherever moisture is to be found. This property prevents a plant from exhausting the earth in fi 2 228 PHYSIOLOGY. BOOK II. which it grows ; for, as the roots are always spreading further and further from the main stem, they are continually entering new soil, the nutritious properties of which are unexhausted. It is generally believed that roots increase only by their extremities, and that, once formed, they never undergo any subsequent elongation. This was first noticed by Du Hamel, who passed fine silver threads through young roots at differ- ent distances, marking on a glass vessel corresponding points with some varnish : all the threads, except those that were within two or three lines of the extremity, always continued to answer to the dots of varnish on the glass vessel, although the root itself increased considerably in length. Variations in this experiment, which has also been repeated in another way by Mr. Knight, produced the same result. It is possible that this peculiarity may be universal in exogenous plants; but it certainly is not constant in endogenous plants ; and I doubt very much whether it is not confined to roots with a woody structure. From the following experiments it will be seen that in Orchideae the root elongates independently of its extremity. On the 5th of August I tied threads tightly round the root of a Vanilla, so that it was divided into three spaces, of which one was 7 inches long, another 4 inches, and the third, which wras the free growing extremity, 1 inch and f. On the 19th of September the first space measured *J\ inches, the second 4f inches, and the third or growing extremity 2-J inches. A root of Aerides cornutum was, on the 5th of August, divided by ligatures into spaces, of which the first measured 1 foot 3 inches, the second 1\ inches, the third 3^ inches, and the fourth, or growing end, 1 inch and -J. On the 19th September the first space measured 1 foot 3^ inches, the second 2 J inches, the third §\ inches, and the fourth 4f inches. Occasionally roots appear destined to act as reservoirs of nutriment, on which those of the succeeding year may feed when first developed, as is the case in the Orchis, the Dahlia, and others. But it must be remarked, that the popular no- tion extends this circumstance far beyond its real limits, by including among roots bulbs, tubers, and other forms of stem in a state of anamorphosis. CHAP. II. ROOT. 229 By some botanists, and among them by M. De Candolle, it has been thought that roots are developed from special organs, which are to them what leaf-buds are to branches; and this function has been assigned to those little glandular swellings so common on the willow, called lenticular glands by Guettard, and lenticelles by De Candolle. Accordingto Mr. Knight, the energies of a variety artificially produced exist longer in the system of the root than in that of the stem; so that it is more advisable to propagate old varieties of fruit trees from cuttings of the root than from those of the stem. The roots not only absorb fluid from the soil, but they return a portion of their peculiar secretions back again into it; as has been found by Brugmans, who ascertained that some plants exude an acid fluid from their spongioles ; and also by Mr. Macare, who has proved that to excrete super- abundant matter from the roots is a general property of the vegetable kingdom. This, taken together with the fact that plants cannot digest their own secretions, explains why soil is so deteriorated by one species having long grown in it, that it will not support other individuals of the same species, until the fecal matter deposited in it shall have been decomposed. This is the solution of the necessity of the rotation of crops. 8 3 230 PHYSIOLOGY. BOOK. II. CHAPTER III. OF THE SAP. For the sustenance of plants a fluid is necessary which is absorbed by the roots from the earth, then sent upwards into the stem, afterwards impelled into the leaves, whence it descends through the liber, transferring itself to the inmost parts of the wood. This fluid, which constitutes the blood of plants, is called the sap. When first introduced into the sys- tem, and even when altered in some degree, by having dis- solved the various substances it encounters in its passage, it is true sap ; afterwards, when its nature has been more changed by elaboration in the leaves, it becomes what is called the proper juice. If the sap be examined in its most simple state, it will be found to consist of water, mucilage, and sugar. As the two last can scarcely have been absorbed directly from the earth, it is inferred that as soon as the fluids, taken up by the roots, enter the system they suffer some chemical decomposition, the result of which is the production of mucilage and sugar. In addition to the supply of sap which is obtained by the roots, a certain quantity is, no doubt, also absorbed from the atmosphere by the leaves ; as is evident from succulent plants, which will continue to grow and acquire weight long after their roots are severed from the earth. This absorption on the part of the leaves chiefly takes place at night, or in cloudy weather ; while perspiration, on the other hand, goes on in the daytime in bright weather. With regard to the chemical nature and changes of the sap, I cannot do better than give the statement of Link, with some necessary alterations. The food of plants must be composed of oxygen, hydrogen, carbon, and azote. Water, consisting of oxygen and hydrogen alone, is not sufficient. Many ex- periments, indeed, have been instituted to prove that pure water is a sufficient food, especially by Van Helmont, Eller, chap. in. sap. 231 Bonnet, Du Hamel, and others ; but it is probable, as Wal- lerius has inferred, that the water out of which plants are formed already contains the necessary chemical principles. To this it is objected, that plants grown in water alone never arrive at perfection or mature their seeds. But this is not strictly true : they do perfect their seeds ; but it is not sur- prising that crude water should be insufficient for purposes which are fully answered by water properly mixed and tempered. That the extractive matter contained in earth was the real food of plants, was long ago stated by Woodward and Kylbel; and most physiologists have adopted this opinion. But it has been estimated by Theodore de Saussure that a plant when dried does not derive more than a twentieth part of its weight from extractive matter and carbonic acid dissolved in water : now, supposing this calculation not to be very accurate, yet it is probable that it is not far from the truth ; and it at least serves to show that extractive matter and carbonic acid are not alone sufficient for the nutriment of plants. Nevertheless, if neither extractive matter nor carbonic acid can be considered to constitute exclusively the food of plants, it is at least quite certain that they not only cannot exist with- out the latter, but that it forms by far the greater part of their food. It is well known that roots cannot perform their func- tions unless within the reach of the atmosphere. This arises from the necessity for their feeding upon carbonic acid, which, after having been formed by the oxygen of the atmosphere combining with the carbon in the soil, is then received into the system of the plant, to be impelled upwards, dissolved in the sap till it reaches the leaves, where it is decomposed by light, the oxygen liberated, and the carbon fixed. It has also been ascertained that, feed plants as you will, they will neither grow nor live whether you offer them oxygen, hydrogen, azote, or any other gaseous or fluid principle, unless carbonic acid is present. Those principles are called foreign to plants which cannot be referred to either hydrogen, oxygen, carbon, or azote: such, for example, are carbonate of soda, sulphate of soda, nitrate of soda, the carbonates of potash, lime, and magnesia, 232 PHYSIOLOGY. BOOK II. phosphate of lime, chlorides of soda and potash, and the oxides of aluminum, silicium, iron, and manganese ; and even, occasionally, phosphorus. De Saussure has demon- strated that the chemical principles which are present in soil are also to be found in the ashes of a plant that has grown in it; but he admits they undergo certain chemical changes, in consequence of the organic powers of vegetation. Dr. John, also, has ascertained that several salts, when taken up by the roots, undergo peculiar changes; as, for instance, potash into soda. To these, many well known observations may be added ; plants growing in saline places contain so much salt, that it is perceptible to the taste. Potash also is yielded by the ashes of plants growing in salt soil ; so that there is no doubt that this substance is produced by the detraction by organic power of the chlorine of chloride of potash. There are, however, some experiments which, if they could be depended on, would materially weaken these hypotheses. Schrader grew barley and rye in well washed flower of sul- phur, moistened with distilled water : they were afterwards analysed, and found to contain silex, lime, and magnesia, as well as oxides of iron and manganese. The same plants produced in earth did not yield a greater weight of ashes than those grown in sulphur; and these experiments are confirmed by those of Braconnot, as recorded in the Annales de Chimie, vol. lxi. p. 187. White mustard was grown in well washed litharge and flower of sulphur, moistened with distilled water ; and its ashes yielded oxide of silicium and alumi- num, carbonate of lime, and oxide of iron. Both these ob- servers conclude that these foreign principles were produced by the organic power of vegetation; but Gehlen suggests that, as many principles are dissolved in oxygen and dis- persed through the atmosphere, they may be communicated to plants by that medium. As it seems, however, certain that chemical principles do undergo changes from organic powers, it is not improbable that all the foreign principles of plants were originally suspended in water ; but that plants have no capability of doing more than decompose such com- pound principles as they may absorb. To reconcile the experiments of Saussure and John with CHAP. III. SAP. 233 those of Schrader and Braconnot, it is suggested by Professor Link that the power and necessity of taking up matter from the soil varies in different plants : that some depend wholly upon it for their formation, others less, and some not at all. Thus, Salsolas grow but in saline soilj and in soil destitute of salt become languid and weak; other plants will only vegetate in calcareous earth : Trifolium pratense prefers gypsum, and succulent plants scarcely require soil at all. That some plants have the power of secreting one kind of accessory principle, and others another kind from the same food, is clear from the fact, that, if wheat and peas be grown in the same water, earth, or medium, the former will uniformly deposit silex in their cuticle, and the latter never. The course which is taken by the sap after entering a plant is the next subject of consideration. The opinion of the old botanists was, that it ascended from the roots between the bark and the wood : but this has been long disproved by modern investigators, and especially by the experiments of Mr. Knight. If a trunk is cut through in the spring, at the time the sap is rising, this fluid will be found to exude more or less from all parts of the surface of the section, except the hardest heart-wood, but most copiously from the alburnum. If a branch is cut half through at the same season, it will be found that, while the lower face of the wound bleeds copiously, scarcely any fluid exudes from the upper face; from which, and other facts, it has been fully ascertained that the sap rises through the wood, and chiefly through the alburnum. Ob- servations of the same nature have also proved that it descends through the liber. But the sap is also diffused laterally through the cellular tissue, and this with great rapidity; as will be apparent upon placing a branch in a coloured in- fusion, which will ascend and descend in the manner just stated, and will also disperse itself laterally in all directions round the principal channels of its upward and downward route. Corti, in 1774, Fontana, L. C. Treviranus, and especially Professor Amici, have made some most curious observations upon the movement of the sap in Chara. If a portion of Chara flexilis, or of any of the transparent species, or of any 234; PHYSIOLOGY. BOOK II. crustaceous kind, the opaque cuticle of which is first scraped away, be examined, a current of sap will be distinctly seen in each cellule setting from joint to joint, flowing down one side and returning up the other, without any membrane intervening to separate the opposing curi'ents. Each cellule has a move- ment of its own, independent of that of the cellules above and below it. Sometimes the movement stops, and then goes on again after a brief interval If a cellule is divided into two by a ligature passed round it, a separate movement is seen in each of the divisions. This motion is rendered distinctly obvious by the numerous minute green granules which float in the transparent fluid, and which follow the course of the currents. Another sort of motion, which it is probable is common to all descriptions of plants, has been seen by Link and others, particularly by Schultz, in the Chelidonium, Ficus Carica, and other plants. It is described as an exceedingly rapid motion of the fluid, which rushes out of one set of vessels, apparently tubes of woody fibre, into another, in a constant, uninterrupted stream. This has been denied, it is true, by Dutrochet; but he does not appear to have succeeded in seeing that which is nevertheless visible enough, if proper precautions are taken. Mirbel and Cassini both confirm the statements of Schultz ; and to their testimony I may be per- mitted to add my own. In Alisma plantago and the trans- parent stipules of Ficus elastica I have distinctly seen powerful currents, such as are described, rushing along the tubes, like a stream of water down an inclined channel. That there must be an exceedingly rapid flow of sap in many plants, is evident from the great loss they often expe- rience by perspiration, — all which must be made good by fluid absorbed by the roots. A young vine-leaf, in a hot day, per- spires so copiously, that, if a glass be placed next its under surface, it is presently covered with dew, which, in half an hour, runs down in streams. Hales computed the perspir- ation of plants to be seventeen times more than that of the human body. He found a sun-flower lose one pound four ounces, and a cabbage one pound three ounces a day by perspiration. Guettard asserts that the young shoots of CHAP. II. SAP. 235 Cornus mascula lose twice their own weight a day. This perspiration is regulated by the number of the stomata : hence evergreens, in which they are small, and less numerous than in deciduous or herbaceous plants, perspire much less. With this function are connected all the phenomena that attend transplantation. If a growing plant is removed from one situation to another in the summer, it will die ; because its spongioles will be so much destroyed as to be incapable of absorbing fluid from the soil as fast as it is given off by the leaves ; and hence the system will be emptied of fluid. But if a plant is growing in a pot, it may be transplanted at any season of the year; because its spongioles, being uninjured, will be able to counterbalance the loss caused by perspiration, as well after transplantation as before, if not better. With regard to the vessels through which this universal dif- fusion of the sap takes place, it has already been stated that its upward course is always through the woody fibre, and probably also through the ducts; and that it passes down- wards through the woody fibre. But there can be no reason- able doubt that it is also dispersed through the whole system by means of some permeable quality of the membranes of the cellular tissue, which is invisible to our eyes, even aided by the most powerful glasses. It has also been suggested that the sap finds its way upwards, downwards, and laterally through the intercellular passages which exist at the points of union of every individual elementary organ. That such a channel of communicating the sap is employed by Nature to a certain extent I do not doubt, especially in those plants in which the intercellular passages are very large ; but whether this is an universal law, or has only a partial operation, is quite unknown, and is not perhaps susceptible of absolute proof. Link seems disposed to deny any conveyance of fluids through the intercellular passages. The accumulation of sap in plants appears to be attended with very beneficial consequences, and to be deserving of the especial attention of gardeners. It is well known how weak and imperfect is the inflorescence of the turnip tribe, forced to flower before their fleshy root is formed ; and how vigorous it is after that reservoir of accumulated sap is completed. Mr. 236 PHYSIOLOGY. BOOK II. Knight, in a valuable paper upon this subject, remarks that the fruit of melons, which sets upon the plant when very young, uniformly falls off; while, on the contrary, if not allowed to set until the stem is well formed, and much sap accumulated for its support, it swells rapidly, and ripens without experiencing any deficiency of food in the course of its growth. In like manner, if a fruit tree is by any circum- stance prevented bearing its crop one year, the sap that would have been expended accumulates, and powerfully contributes to the abundance and perfection of the fruit of the succeeding year. And again, the plan recommended by Mr. Knight, of always planting large tubers of the potatoe, is another proof of the importance of plenty of accumulated sap to the vigorous growth of all plants. The cause of the motion of the sap is a subject which has long excited great curiosity, and has given rise to numberless conjectures. It was for a long time believed that there was a sort of circulation of the sap of plants, to and from a common point, analogous to that of the blood of animals : but this was disproved by Hales, and is not now believed. This excellent observer, whose " Statics " are an eternal monument of his industry and skill, thought that the motion of the sap, the rapidity of which he had found to be greatly influenced by weather, depended upon the contraction and expansion of the air, which exists in great quantities in the interior of plants. Others have ascribed the motion to capillary attraction. Mr. Knight was once of opinion that it depended upon a hygrometrical property of the plates of silver grain (me- dullary rays), which traverse the stem in all directions. A number of other theorists have called to their aid a supposed irritability of the vessels ; but no contraction of the vessels has ever yet been noticed, and certainly does not take place in Chara, where the motion has been most distinctly observed. Du Petit Thouars suggests that it arises thus : — in the spring, as soon as vegetation commences, the extremities of the branches and the buds begin to swell : the instant this hap- pens a certain quantity of sap is attracted out of the circum- jacent tissue for the supply of those buds ; the tissue, which is thus emptied of its sap, is rilled instantly by that beneath or CHAP. III. SAP. 237 about it: this is in its turn replenished by the next; and thus the whole mass of fluid is set in motion, from the extremities of the branches down to the roots. Du Petit Thouars is therefore of opinion that the expansion of leaves is not the effect of the motion of the sap, but, on the contrary, is the cause of it ; and that the sap begins to move at the extremities of the branches before it stirs at the roots. That this is really the fact, is well known to foresters and all persons accus- tomed to the felling or examination of timber in the spring. Some good observations upon this were communicated to Mr. Loudon's Gardener's Magazine, by Mr. Thomson, gardener at Welbeck; who, however, drew a wrong inference from them. Amici is of opinion that the motion in Chara depends upon galvanic action ; and Dutrochet has since formed a theory of all the motions of fluids in plants depending upon the same agency. He found that small bladders of animal and vegetable membrane, being filled with a fluid of greater density than water, securely fastened, and then thrown into water, acquired weight ; he also remarked, that if the experiment was reversed, by filling with water and immersing them in a denser fluid, the contrary took place, and that the bladders lost weight : he took a small bladder, and filled it with milk, or gum arabic dissolved in water ; to the mouth of this bladder he adapted a tube, and then plunged the bladder in water: in a short time the milk rose in the tube, whence he inferred that water had been attracted through the sides of the bladder. This experi- ment was also reversed, by filling the bladder with water, and plunging it in milk : the fluid then fell in the tube, whence he inferred that water had been attracted through the coat of the bladder into the milk. From these, and other experiments, M. Dutrochet arrived at the inference that, if two fluids of unequal density are separated by an animal or vegetable membrane, the denser will attract the less dense through the membrane that divides them : and this property he calls endosmose, when the attraction is from the outside to the inside; and exosmose, when it operates from the inside to the outside. In pursuing this investigation he remarked, that if an empty bladder is immersed in water, and the negative pole 238 PHYSIOLOGY. BOOK II. of a galvanic battery introduced into it, while the positive pole is applied to the water on the outside, a passage of fluid takes place through the membrane, as had previously hap- pened when the bladder contained a fluid denser than water ; by reversing the experiment, the reverse was found to take place: from all which Dutrochet deduces the following theory, that when two fluids of unequal density are separated by an intervening membrane, the more dense is negatively electri- fied, and the less dense positively electrified ; in consequence of which two electric currents of unequal power set through the membrane, carrying fluid with them ; that which sets from the positive pole, or less dense fluid, to the negative pole, or more dense fluid, being much the most powerful : and that the fluids of plants being more dense than those which sur- round them, a similar action takes place between them and the water in the soil, by means of which the latter is con- tinually impelled into their system. Philosophers do not seem disposed to admit the legitimacy of M. Dutrochet's con- clusion, that this transmission takes place by means of galvanic agency ; but that the phenomenon is correctly described by the ingenious author, and that it is constantly operating in plants, is beyond all dispute. It is by endosmose that vapour is absorbed from the atmosphere, and water from the earth ; that sap is attracted into fruits by virtue of their greater density ; and probably that buds are enabled to empty the tissue that surrounds them when they begin to grow : it will, perhaps, be found the most ready explanation of most of the phenomena connected with the movement of fluids. 239 CHAPTER IV. OF THE PITH, WOOD, AND BARK. Various are the notions that from time to time have been entertained about the pith. The functions of brain, lunors, stomach, nerves, spinal marrow, have by turns been ascribed to it. Some have thought it the seat of fecundity, and have believed that fruit trees deprived of pith became sterile; others supposed that it was the origin of all growth ; and another class of writers, we cannot say observers, have declared that it was the channel of the ascent of sap. It is, however, no part of the plan of this work to refute this and similar ex- ploded speculations. It is probable that its real and only use is to serve in the infancy of a plant for the reception of the sap, upon which the young and tender vessels that surround it are to feed when they are first formed ; a time when they have no other means of support. M. Dutrochet considers it to act not only as a reservoir of nutriment for the young leaves, but also to be the place in which the globules, which he calls nervous corpuscles, are formed out of the elaborated sap. {JO Agent Imme'diat, &c, p. 44, &c.) The medullary sheath seems to perform a far more im- portant part in the economy of plants ; it diverges from the medulla whenever a leaf is produced, and, passing through the petiole, ramifies among the cellular tissue of the lamina, where it appears as veins : hence veins are always composed of bundles of woody fibre and spiral vessels. So situated, the veins are in the most favourable position that can be imagined for absorbing the fluid that, in the first instance, is conducted to the young pith, and that is subsequently impelled upwards through the woody fibre. So essential is the medullary sheath to vegetation in the early age of a branch, that, as is well known, although the pith and the bark, and even the young wood, may be destroyed, without the life of a young 240 PHYSIOLOGY. BOOK II. shoot being much affected ; yet, if the medullary sheath be cut through, the pith, bark, or wood being left, the part above the wound will perish. The bark acts as a protection to the young and tender wood, guarding it from cold and external accidents. It is also the medium in which the proper juices of the plant in their descent from the leaves are finally elaborated, and brought to the state which is peculiar to the species. It is from the bark that they are horizontally communicated to the medullary rays, by them to be deposited in the tissue of the wood. Hence, the character of timber is almost wholly dependent upon the influence of the bark, as is apparent from a vertical section of a grafted tree, through the line of union of the stock and scion. This line will be found so exactly drawn, that the limits of the two are determined in the oldest spe- cimens as accurately as if they were fixed by rule and line : the woody tissue will be found uninterruptedly continuous through the one into the other, and the bark of the two indissolubly united ; but the medullary rays emanating from the bark of each will be seen to remain as different as they were, while the stock and scion were distinct individuals. As the bark, when young, is green like the leaves, and as the latter are manifestly a mere dilatation of the former, it is highly probable, as Mr. Knight believes, that the bark exer- cises an influence upon the fluids deposited in it wholly analogous to that exercised by the leaves, which will be here- after explained. Hence it has been named, with much truth, the universal leaf of a vegetable. The business of the medullary rays is, no doubt, exclu- sively to maintain a communication between the bark, in which the secretions receive their final elaboration, and the centre of the trunk, in which they are at last deposited. This is apparent from tangental sections of dicotyledonous wood manifesting an evident exudation of liquid matter from the wounded me- dullary rays, although no such exudation is elsewhere visible. In endogenous plants, in which there appears no necessity for maintaining a communication between the centre and circum- ference, there are no medullary rays. These rays also serve to bind firmly together the whole of the internal and external CHAP. IV. PITH, WOOD, AND BARK. 2il parts of a stem, and they give the peculiar character by which the wood of neighbouring species may be distinguished. If plants had no medullary rays, their wood would probably be, in nearly allied species, undistinguishable ; for we are scarcely aware of any appreciable difference in the appearance of fibrous or vascular tissue. But the medullary rays differing in abundance, in size, and in other respects, impress charac- ters upon the wood which are extremely marked. Thus, in the cultivated cherry, the plates of the medullary rays are very thin, the adhesions of them to the bark are very slight, and hence a section of the wood of that plant has a pale, smooth, homogeneous appearance ; but in the wild cherry the medullary plates are much thicker, they adhere to the bark by deep broad spaces, and are arranged with great irre- gularity, so that a section of the wood of that variety has a deeper colour, and a twisted, knotty, very uneven appearance. As the medullary rays develope only horizontally, when two trees in which they are different are grafted or budded to- gether, the wood of the stock will continue to preserve its own peculiarity of grain, notwithstanding its being formed by the woody matter sent down by the scion ; for it is the hori- zontal developement that gives its character to the grain, and not the perpendicular fibres which are incased in it. The wood is at once the support of all the deciduous organs of respiration, digestion, and impregnation, the deposit of the secretions peculiar to individual species, and also the reservoir from which newly forming parts derive their sustenance until they can establish a communication with the soil. Regarding the precise manner in which it is created, there has been great diversity of opinion. Linnaeus thought it was produced by the pith ; Grew, that the liber and wood were deposited at the same time in a single mass which afterwards divided in two, the one half adhering to the centre, the other to the circumference ; Malpighi conceived that the wood of one year was produced by an alteration of the liber of the previous season. Duhamel believed that it was deposited by the se- cretion already spoken of as existing between the bark and wood, and called cambium : he was of opinion that this cam- bium was formed in the bark, and became converted into both 242 PHYSIOLOGY. BOOK II. cellular tissue and woody fibre; and he demonstrated the^ fallacy of those theories according to which new wood is produced by the wood of a preceding year. He removed a portion of bark from a plum tree ; he replaced this with a similar portion of a peach tree, having a bud upon it. In a short time a union took place between the two. After wait- ing a sufficient time to allow for the formation of new wood, he examined the point of junction, and found that a thin layer of wood had been formed by the peach bud, but none by the wood of the plum, to which it had been tightly applied. Hence he concluded that alburnum derives its origin from the bark, and not from the wood. A variety of similar experi- ments was instituted with the same object in view, and they were followed by similar results. Among others, a plate of silver was inserted between the bark and the wood of a tree at the beginning of the growing season. It was said, that if new wood was formed by old wood it would be subsequently found pushed outwards, and continuing to occupy the same situation ; but that if new wood was deposited by the bark, the silver plate would in time be found buried beneath new layers of wood. In course of time the plate was examined, and was found enclosed in wood. Hence the question as to the origin of the wood seemed settled ; and there is no doubt that the experiments of Du- hamel are perfectly accurate and satisfactory as far as they go. It soon, however, appeared that, although they certainly proved that new wood is not produced by old wood, it was not equally clear that it originated from the bark. Accord- ingly a new set of experiments was instituted by Mr. Knight, for the purpose of throwing a still clearer light upon the pro- duction of the wood. Having removed a ring of bark from above and below a portion of the bark furnished with a leaf, Mr. Knight remarked that no increase took place in the wood above the leaf, while a sensible augmentation was observable in the wood below the leaf. It was also found that if the upper part of a branch is deprived of leaves, the branch will die down to the point where leaves have been left, and below that will flourish. Hence an inference is drawn that the wood is not formed out of the bark as a mere deposit from it, CHAP. IV. PITH, WOOD, AND BARK. 24-3 but that it is produced from matter elaborated in the leaves and sent downwards, — either through the vessels of the inner bark, along with the matter for forming the liber by which it is subsequently parted with ; or that it and the liber are transmitted distinct from one another, the one adhering to the alburnum, the other to the bark. I know of no proof of the former supposition : of the latter there is every reason to believe the truth. Mr. Knight is of opinion that two distinct sets of vessels are sent down, one belonging to the liber, the other to the alburnum ; and if a branch of any young tree, the wood of which is formed quickly, be examined wdien it is first bursting into leaf, these two sets may be distinctly seen and traced. Take, for instance, a branch of lilac in the be- ginning of April and strip off its bark : the new wood will be distinctly seen to have passed downwards from the base of each leaf, diverging from its perpendicular course, so as to avoid the bundle of vessels passing into the leaf beneath it; and if the junction of a new branch with that of the previous year be examined, it will be found that all the fibres of wood already seen proceeding from the base of the leaves, having arrived at this point, have not stopped there, but have passed rapidly downwards, adding to the branch an even layer of fibrous matter or young wood ; and turning off at every pro- jection which impedes them, just as the water of a steady but rapid current would be diverted from its course by obstacles in its stream. Now, if the new wood were a mere deposit of the bark, the. latter, as it is applied to every part of the old wood, would deposit the new wood equally over the whole surface of the latter, and the deviation of the fibres from ob- stacles in their downward course could not occur. This, therefore, in my mind, places the question as to the origin of the wood beyond all further doubt. Mirbel, who formerly advocated the doctrine of wood being deposited by bark, has, with the candour of a man of real science, fairly ad- mitted the opinion to be no longer tenable ; and he has sug- gested in its room that wood and bark are independent formations, — which is no doubt true, — but, he adds, created out of cambium, in which it is impossible to concur : for this reason. All the writers hitherto mentioned or adverted H 2 244 PHYSIOLOGY. BOOK II. to have considered the formation of wood only with reference to exogenous trees, and to such only of them as are the com- mon forest plants of Europe. Had they taken into account exotic trees or any endogenous plants, they would have seen that none of their theories could possibly apply to the form- ation of wood in that tribe. In many exogenous plants of tropical countries wood is not deposited in regular circles all round the axis, but only on one side of the stem, or along certain lines upon it : were it a deposit from the bark, or a metamorphosis of cambium, it would necessarily be deposited with some kind of uniformity. In endogenous trees there is no cambium, and yet wood is formed in abundance; and the new wood is created in the centre, and not in the circum- ference : so that bark can have, in such cases, nothing what- ever to do with the creation of wood. No doubt aware of most of the difficulties in the way of the common theories of the formation of wood, M. Du Petit Thouars, an ingenious French physiologist, who had possessed opportunities of examining the growth of vegetation in tro- pical countries, constructed a theory, which, although in many points similar to the one proposed, but not proved, by his countryman, De la Hire, is nevertheless, from the facts and illustrations skilfully brought by the French philosopher to his aid, to be considered legitimately as his own. The attention of Du Petit Thouars appears to have been first especially called to the real origin of wood by having re- marked, in the Isle of France, that the branches which are emitted by the truncheons of Dracaena (with which hedges are formed in that colony) root between the rind and old wood, forming rays of which the axis of the new shoot is the centre. These rays surround the old stem ; the lower ones at once elongate greatly towards the earth, and the upper ones gradually acquire the same direction ; so that at last, as they become disentangled from each other, the whole of them pass downwards to the soil. Reflecting upon this curious fact, and upon a multitude of others which I have no space to detail, he arrived at the conclusion, that it is not merely in the property of increasing the species that buds agree with seeds, but that they emit roots in like manner; and that the CHAP. IV. PITH, WOOD, AND BARK. 245 wood and liber are both formed by the downward descent of bud- roots, at first nourished by the moisture of the cambium, and finally imbedded in the cellular tissue which is the result of the organisation of that secretion. That first tendency of the embryo, when it has disengaged itself from the seed, to send roots downwards and a stem and leaves upwards, and to form buds in the axilla? of the latter, is in like manner possessed by the buds themselves ; so that plants increase in size by an endless repetition of the same phenomenon. Hence a plant is formed of multitudes of buds or fixed embryos, each of which has an independent life and action: by its elongation upwards forming new branches and con- tinuing itself, and by its elongation downwards forming wood and bark ; which is therefore, in Du Petit Thouars's opinion, a mass of roots. A great deal of opposition has been offered to this view, especially among this writer's own countrymen ; but it is re- markable that many of his antagonists have been from a class of naturalists of whom it may be said, that they are better known in consequence of the celebrity of the object of their attack than for any reputation of their own. To this, how- ever, there are some exceptions, as, for instance, MM. Mirbel and Desfontaines, two of the most learned botanists of France. This theory, nevertheless, seems the only one that is adapted at once to the explanation of the real cause of the many anomal- ous forms of exogenous stems which must be familiar to the re- collection of all botanists, and that, at the same time is equally applicable to the exogenous and endogenous modes of growth ; a condition which, it will be readily admitted, is indispensable to any theory of the formation of wood that may be proposed. It also offers the simplest explanation of the phenomena that are constantly occurring in the operations of gardening. It has recently been a subject of discussion in the Academy of Sciences at Paris; when M. Poiteau supported the theory, and MM. Mirbel, Cassini, and Desfontaines opposed it. The arguments used by the latter were two, both of which are un- doubted fallacies. The first was, that if a large ring of bark be taken from the stem of a sycamore, and be replaced by a similar ring entirely destitute of buds from a red maple, the R 3 246 PHYSIOLOGY. BOOK II. new bark will graft itself with the sycamore, and in time red maple wood will be formed beneath it. They said this lig- neous production could not be derived from the buds of the red maple, because the ring of bark was devoid of any ; nor could it proceed from the buds of the sycamore, because they would produce sycamore wood. But it is obvious that, in this experiment, the character of the red maple wood was derived from its medullary rays, which first formed an adhesion with those of the sycamore, and afterwards an independent hori- zontal formation, through which the fibres of the sycamore descended without altering its character. The other case was, that if a large ring of bark be taken from the trunk of a vigorous elm or other tree without being replaced with any thing, new beds of wood will be found in the lower as well as upper part of the trunk ; while no ligneous production wilt appear on the ring of wood left exposed by the removal of the bark. Now this is so directly at variance with the observ- ations of others, that it is impossible to receive it as an ob- jection until its truth shall have been demonstrated. It is well known that if the least continuous portion of liber be left upon the surface of a wound of this kind, that portion is alone sufficient to establish the communication between the upper and lower lips of the wound ; but, without some such slight channel of union, it is directly contrary to experience that the part of a trunk below an annular incision should in- crease by the addition of new layers of wood until the lips of the wound are united, unless buds exist upon the trunk below the ring. The secretion called cambium, in the opinion of those who believe wood and bark to be independent simultaneous form- ations from the surface of the old wood and bark, is the matter which finally becomes organised as such : in Du Petit Thouars's theory it is a matter of organisation only as far as regards the origin of the cellular tissue of the medullary rays, and of the bark ; while the superfluity of its moisture is a pro- vision made by nature for the nutriment of the young fibres that descend through it. 247 CHAPTER V. OF THE LEAVES. Leaves are at once organs of respiration, digestion, and nutrition. They elaborate the crude sap impelled into them from the stem, parting with its water, adding to it carbon, and exposing the whole to the action of air ; and while they supply the necessary food to the young fibres that pass downwards from them, and from the buds in the form of alburnum and liber, they also furnish nutriment to all the parts immediately above and beneath them. There are many experiments to show that such is the purpose of the leaves. If a number of rings of bark are separated by spaces without bark, those which have leaves upon them will live much longer than those which are destitute of leaves. If leaves are stripped off a plant before the fruit has commenced ripening, the fruit will fall off and not ripen. If a branch is deprived of leaves for a whole sum- mer, it will either die or not increase in size perceptibly. The presence of cotyledons, or seminal leaves, at a time when no other leaves have been formed for nourishing the young plant, is considered a further proof of the nutritive purposes of leaves : if the cotyledons are cut off, the seed will either not vegetate at all, or slowly and with great difficulty ; and if they are injured by old age, or any other circumstance, they pro- duce a languor of habit which only ceases with the life of the plant, if it be an annual. This is the reason why gardeners prefer old melon and cucumber seeds to new ones : in the former the nutritive power of the cotyledons is impaired, the young plant grows slowly, a languid circulation is induced from the beginning ; by which excessive luxuriance is checked, and fruit formed rather than leaves or branches. Various are the secretions of plants that take place through the leaves : in those of monocotyledonous tropical plants in our hot-houses, nothing is more common than to see drops of water forming upon them from the effect of perspiration ; in r 4 248 PHYSIOLOGY. BOOK II. Limnocharis Plumieri there is a large pore terminating the veins of the apex of the leaf, from which water is constantly distilled. The pitchers of Nepenthes, which are only a par- ticular kind of leaves, secrete water enough to fill half their cavity. But, besides this more subtle fluid, secretions of a grosser quality take place in plants. The honey dew, which is so often attributed to insects, is one instance of the perspir- ation of a viscid, saccharine substance ; the manna of the ash is another; and the gum ladanum that exudes from the Cistus ladaniferus is a third instance of this kind of per- spiration. It is believed that absorption takes place indifferently by either the upper or under surface of the leaf, but that some plants absorb more powerfully by one surface than by the other. Bonnet found that while the leaves of Arum, the kidney-bean, the lilac, the cabbage, and others, retained their verdure equally long whichever side was deprived of the power of absorption, the Plantago, some Verbascums, the mar- vel of Peru, and others, lost their life soonest where the upper surface was prevented from absorbing ; and that in a number of trees and shrubs the leaves were killed very quickly by preventing absorption by the lower surface. From this there is only one safe conclusion to be drawn ; that the absorbing surface of leaves varies in different species, and depends upon their peculiar organisation. Leaves usually are so placed upon the stem that their upper urface is turned towards the heavens, their lower towards the earth ; but this position varies occasionally. In some plants they are imbricated, so as to be almost parallel with the stem ; in others they are deflexed till the lower surface becomes almost parallel with the stem, and the upper surface is far re- moved from opposition to the heavens. A few plants, more- over, invert the usual position of the leaves by twisting the petiole half round, so that either the two margins become opposed to earth and sky, or the lower surface becomes up- permost : this is especially the case with plants bearing phyl- lodia, or spurious leaves. At night a phenomenon occurs in plants which is called their sleep : it consists in the leaves folding up and drooping, CHAP. V. LEAVES. 249 as those of the sensitive plant when touched. This scarcely happens perceptibly except in compound leaves, in which the leaflets are articulated with the petiole, and the petiole with the stem : it is supposed to be caused by the absence of light, and will be farther spoken of under the head of irritability. After the leaves have performed their functions, they fall off: this happens at extremely unequal periods in different species. In some they all wither and fall off by the end of a single season ; in others, as the beech and hornbeam, they wither in the autumn, but do not fall off till the succeeding spring ; and, in a third class, they neither wither nor fall off the first season, but retain their verdure during the winter, and till long after the commencement of another year's growth : these are our evergreens. Mirbel distinguishes leaves into three kinds, as characterised by their periods of falling: — 1. Fugacious or caducous, which fall shortly after their appearance ; as in Cactus opuntia. 2. Deciduous or annual, which fall off in the autumn; as Pyrus malus. 3. Persistent, evergreen, or perennial, which remain perfect upon the plant beyond a single season ; as Ilex aquifolium, Primus pseudo-cerasus, &c. With regard to the cause of the fall of the leaf a number of explanations have been given, which may be found in JVilldenoio's Principles of Botany, p. 336. Sir James Smith was of opinion with Vrolik, that it is evidently a sloughing or casting off diseased or worn out parts, and in this I agree with them ; but neither of these authors afford any ex- planation of the cause of this sloughing ; nor do I think that it has been satisfactorily accounted for by any one, except Du Petit Thouars. If you will watch the progress of a ti-ee, — of the elder for example, — says this writer, you will perceive that the lowest leaves upon the branches fall long before those at the extremities. The cause of this may be, perhaps, explained upon the following principle. In the first instance, the base of every leaf reposes upon the medulla of the branch to the sheath of which it is attached. But, as the branch increases in diameter by the acquisition of new 250 PHYSIOLOGY. BOOK II. wood, the space between the base of the leaf and the medulla becomes sensibly augmented. It has, therefore, been ne- cessary that the fibres by which the leaf is connected with the medulla should lengthen, in order to admit the deposition of wood between the bark and the medulla. Now how does this elongation take place ? As the bundles of fibres which run from the medulla into the leaf-stalk are at first composed only of spiral vessels, it is easy to conceive that they may be susceptible of elongation by unrolling. And in this seems to lie the mystery of the fall of the leaf; for the moment will come when the spiral vessels are entirely unrolled, and in- capable of any further elongation : they will, therefore, by the force of vegetation, be stretched until they snap, when the necessary communication between the branch and the leaf is destroyed, and the latter falls off. It is highly probable that this interruption of the necessary communication between the leaves and their support is the principal cause of the fall of the leaf in all cases ; and it seems to be strongly supported by the following phenomena : 1. the early fall of the lower leaves of plants during the period of vegetation ; 2. the fall of the leaves of evergreen trees, and of those whose foliage withers and persists during winter, at the period when new wood is formed by the operation of new shoots in the spring ; 3. the ultimate fall of those leaves which are subject to no fixed periods of de- foliation ; 4. the persistence of the leaves of stunted trees, which have not formed wood enough during a single season to cause a rupture of the conducting vessels of the petioles, as the beech and hornbeam. The weil known indication by which gardeners judge of their probable success in trans- planting a tree or other plant in leaf, may also be considered a further proof of the justness of Du Petit Thouars's theory. On such occasions, if the leaves wither and hang upon the branches, the omen is unfavourable ; but if they are cast off, it is a certain indication of success. Here the action of the spiral vessels may be understood to be impaired by the in- terruption of the regular transmission of oxygenated air through them, caused by the act of transplantation : as soon as the energies of the plant are renewed, a sudden increase of CHAP. V. LEAVES. 251 diameter supervenes, the communication is cut off between the leaves and the stem by the too rapid extension of the spiral vessels, and the leaves fall off. Respiration takes place by the power the leaves possess of inspiring and expiring oxygen and decomposing carbonic acid. They have been found to vitiate the atmosphere at nio-ht by inhaling oxygen abundantly, and exhaling a small quantity of carbonic acid ; and to restore the air to its purity in the sun's rays, by decomposing their carbonic acid and partino- with their oxygen. It was long since remarked by Priestley, that if leaves are immersed in water and placed in the sun, they part with oxygen. This fact has been subsequently demonstrated by a great number of curious experiments, to be found in the works of Ingenhouz, Saussure, Senebier, and others. Saussure found that plants in cloudy weather, or at night, inhaled the oxygen of the surrounding atmosphere, but exhaled carbonic acid if they continued to remain in obscurity. But, as soon as they were exposed to the rays of the sun, they respired the oxygen they had previously inhaled, in about the same quantity as they received it, and with great rapidity. Dr. Gilly found that grass leaves exposed to the sun in a jar for four hours produced the following effect : — At the beginning of the Experiment there were in the Jar : — Of nitrogen - - 10.507 Of carbonic acid - - -5.7 Of oxygen ... 2:793 19.000 At the close of the Experiment there were : — Of nitrogen - 10.507 Of carbonic acid - - .37 Of oxygen - - 7.79 18.667 Heyne tells us that the leaves of Bryophyllum calycinum in India, are acid in the morning, tasteless at noon, and bitter in the evening; Link himself found that they readily stained litmus paper red in the morning, but scarcely produced any such effect at noon. The same phenomenon is said also to occur in other plants, as Cacalia ficoides, Sempervivum arboreum, &c. This stain in the litmus paper could not have arisen from the presence of carbonic acid, as that gas will not alter blue paper, but it must have been caused by the oxygen inhaled at night. It has also been found that this last power 252 PHYSIOLOGY. BOOK II. is retained even at noon, if the plant is not exposed to the sun. A similar explanation may be given of a phenomenon remarked by Pajot de Charmes, who found that the flowers of Cichorium intybus were daily changed from blue to white, according to the action of light. It is also well known that fruit is more acid in the morning than in the evening. As the decomposition of carbonic acid gas is thus evidently an important part of the act of respiration, it might be supposed that to supply a plant with a greater abundance of carbonic acid than the atmosphere will usually yield would be attended with beneficial consequences. To ascertain this point several experiments have been instituted ; the most important of which are those of Saussure, who found that, in the sun, an atmosphere of pure carbonic acid gas, or even air, containing as much as sixty per cent., was destructive of vegetable life ; that fifty per cent, was highly prejudicial ; and that the doses became gradually less prejudicial as they were diminished. From eight to nine per cent, of carbonic acid gas was found more favourable to growth than common air. This, however, was only in the sun : any addition, however small, to the quantity of carbonic acid naturally found in the air was pre- judicial to plants placed in the shade. Nitrogen, per se, is incapable of affording any support to the developement of plants, as was proved by Saussure, who found that, five days after immersion in pure nitrogen, the buds of poplars and willows were in a state of decay. This is remarkable, considering how large a proportion of the air we breathe consists of nitrogen. While oxygen and carbon are thus essential to vegetation when not administered in excess, almost all other gases are more or less deleterious. Drs. Turner and Christison found that so small a quantity as to,