BOTANICAL GAZETTE a cnieaen oaaels a NS os int eS ee Rep »)E4Lg THE | 32 BOTANICAL GAZETTE EDITORS: JOHN MERLE COULTER anp CHARLES REID BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS: {2C. ARTE bade University. Fritz NOLL, — me Bonn. “VoLNEY M. SPA CASIMIR DECANDOLLE, mire a. Univ enity ae Michigan. ]. Bs DELON ROLAND THAX Un scout of Padua. Har ADOLF ENGLE Uaiveriity of Berlin. LEON GUIGNARD, ae University, WILLIAM ‘LREL Missouri Botanica Garden. . MARSHALL WA L’Ecole de Pharmacie. University of Cambridge. RoBerT A. HA EUGEN. WARMING E Un nversity ae Wisconsin. Univ: ae of Copenhagen. JinzO MATSUMUR Im VEIT WITTROC perial Ture Tokyo. Ro oyal | Academy of Sciences, Stockholm VOLUME XXXII JULY —DECEMBER, 1901 WITH SEVENTEEN PLATES AND EIGHTY-TWO FIGURES IN THE TEXT CHICAGO, ILLINOIS PUBLISHED BY THE UNIVERSITY OF CHICAGO 1901 Mo.Bot. Garcc., 1902. ‘s ms : od pRTED BY FE ert ee The University of Chicago Press CHICAGO i : " 4 TABLE OF CONTENTS. GE PA On the origin and nature of the middle lamella - - - Charles E. Allen I Structural studies on southwestern Cactaceae (with nine figures) - - - - - - Carleton EL. Preston 35 Gametogenesis and fertilization in Albugo. Contributions from the Hull Botanical hecsespaty eS XXIX (with plates I-Iv) - Frank L. Stevens 77, 157, 238 The ecological relations of the vegetation of western Texas, Contributions from the Hull Botanical makin XX (with twenty-four text figures) - William L. Bray, 99, 195, 262 A study of the sporangia a gametophytes of Selaginella apus and agatts a Tupestris. Contributions from he Hull Bot iL Botanical Laboratory. XX x (with plates cee - - - - - - Florence M. Lyon 124, 170 The application of normal solutions to biological problems James B. Dandeno 229 Further notes on the physiology -E polymorphism in green algae. Contributions from the Hull Botanical Labo- ratory. XXXII - - Burton Edward Livingston 292 New or little known unicellular algae. II. Eremosphaera viridis and Excentrosphaera (with plates x—x11) George Thomas Moore 309 Development of the pollen in some Asclepiadaceae on- tributions from the Hull Botanical erie rat we XX XIII (with plate x1) - - iC, Prye 325 On the distribution of red color in vegetative parts in the New England flora - - - - : - - - F. Grace Smith 332 Some plant abnormalities (with thirty-six figures) - George Harrison Shull 343 The anatomy of the O d (with plates x1v—xvir) J. A. Faull 381 BRIEFER ARTICLES— Note on Basilima and Schizonotus of Rafinesque - Alfred Rehder 56 Potamogeton polygonifolius in Newfoundland, - - Arthur Bennett 58 A new Sphaeralcea - - - - - - - 7. D.A. Cockerell 60 The probabie function of calcium oxalate Pac saene in plants - - - Albert Schneider 142 Notes of travel. VII - - - - - - David G. Fairchild 218 Intramolecular respiration - - - - - T. C. Johnson 303 ee vi : CONTENTS [VOLUME XXXII Notes on re Canadensis Linn. and A. vul- garis Linn. - - J. Schneck 304 Meissner on evergreen needles (with one figure) Edwin Bingham Copeland 356 The instability of the Rochester nomenclature - - M.L. Fernald 359 Flower visits of oligotropic bees. III - - - Charles Robertson 367 Some erroneous references - . - - - £.W. Dd. Holway 421 Puccinia inanipes’~ - - - - . . - £.W.D. Holway 422 The position of pleurococcus and mosses on trees” - Henry Kraemer 422 Contribution to the knowledge of the physiology of karyokinesis (ith one figure) - Arthur Carr Lewis 423 OpEN LETTERS— Normal solutions : é is 2 x : Louis erie 437 James B. Dandeno 439 Basilima, Schizonotus, Sorbaria - - - John Hendley Barnhart 440 » CuRRENT LITERATURE— - - . - 61, 145, 218, 306, 369, 426 abe titles see index under author’s name and Re- Papers noticed in ‘“ Notes for Students” are said under author’s name and subjects NEws—_ - - et os = E - - - 74,154, 227, 307, 379, 443 DATES OF PUBLICATION. . 1, July 24; No. 2, August 20; No. 3, September 24; No. 4, October 21; No. 5, November 25; No. 6, December 26. ERRATA. P. 45, line 5 from below, for composed, read compressed. P, 50, line 14, for “In the absorptive roots the elements are wider, more regular ess thickened,” read “In the absorptive root the medullary rays are narrow and of thin-walled cells containing crystals; in the anchoring root, the elements are wider, without c P. 70, line 7 from below, for Laminaceae read Laminariaceae. P. 71, line 15 from below, insert parenthesis before Ceramothamnton. P. 76, line 13, for aaother read another. P, 78, line 9, for avtemisaefolia read artemisiaefolia. P. 120, line 9 from below, for vegeation read vegetation. P, 158, in legend of center figure of first row, insert one before functional. VOLUME XXXII] : CONTENTS ee | P. 166, line 20, for Monoblepharadineae read Monobiepharidineae. P. 212, legend of figure 12, for J. sabinioides read J. sabinoides. P. 237, line 1, for 1899 read 1889. P. 255, line 12, for philogenetically read phylogenetically. He P, 285, legend figure 23, for Tomillo read Tornillo. if P. 303, line 13, from below, for they read it. P. 306, heading, for LITERATRUE read LITERATURE. P. 325, line 4 of title, for XXXII read XXXIII. : P. 375, line 17, for byrological read bryological. On plates I, II, and III, at head, for XXVII read XXXII. } Vol: XXXII JULY, 1901 q x 3 ae THE ; oe tes ae mt ee ees oer ae J 7 s — sent 7 = ae ¥ as ~ vo eaatee cna «cal aes —_-—~ “wR ee | APPLES MAKE | ie ~~ 4 1 The soap made by PEARS, in Great Britain, is incom- parably the purest and best for the toilet and bath. All sorts of stores sell it, all sorts of people use it. All rights secured. 4 Botanical Gazette A Montbly Journal Embracing all Departments of Botanical Science Subscription per year, $4.00 Single Numbers, 40 Cents The subscription price must be paid in advance. No numbers are sent oe the expiration of the time paid for. No reduction is made to dealers or agen r FOREIGN AGENTS: a | Great Britain — WM. WESLEY & Son, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin St., Strand, London. 18 Shillings. SW. 46, Schénebergerstr. 17a. 18 Marks Vol. XXXII, No. 1 Issued July 24, 1903 4 : CONTENTS ON THE ORIGIN AND NATURE OF THE MIDDLE LAMELLA. Charles E. Allen - I STRUCTURAL STUDIES ON SOUTHWESTERN CACTACEAE (WITH NINE FIGURES). Carleton E. Preston - : s E ss ‘ : es 3 m : é “ 35 BRIEFER ARTICLES. NOTE ON BASILIMA AND SCHIZONOTUS OF RAFINESQUE. Alfred Rehder - - - 56 POTAMOGETON POLYGONIFOLIUS IN NEWFOUNDLAND. Arthur Bennett : : = 58 A SPHAERALCEA. 7. D. A. Cockerell - x : : 2 : : . m 60 Ci morse nf at noes TURE. k = 61 NDBOOK OF bitineceie Sickie: MINOR NOTICES i : : = ‘ i “i 62 NOTES FOR STUDENTS - - z 2 ‘ : = 2 63 NEWS - - - we ean : z 3 . “ . 74 Separates, if de: ing articles will be pris ted, of which 25 eink coe) will be furnished peo the actual cost of the remainde ) i or er (and rt . desired) to be paid for by the aut Pp es of “briefer articles ” (with without covers) will also be supplied at cost. The table below shows the oximate cost sep S canta of plain te a or text with line engravings. 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The journal will continue to appeal to teachers and parents, and each humber will contain practical plans for teaching in every grade from the kindergarten through the high and pedagogic schools. The magazine will appear regularly throughout the year, excepting in the months of August and September, and the subscription price has been reduced from $2.00 to $1.50 per year ; sample copies sent on applica- tion. Special rates to clubs. Agents wanted in all sections of the country. Address Lae DEEDES OF CHICAGO PRESS CHICAGO, ILLINOIS A Triumph of American Progress Established 1875.—The Pioneer of Industrial Life Insur- ance in the United States. Introducer of Many Reforms in Ordinary Life Insurance. Paid Over $50,000,000 to Policy=-Holders, on More Than Five Hundred Thousand Claims. Has in Force Over Four Million Policies, Equiva- lent to about Twelve Times the Population of Buffalo, Insuring Over $600,000,000. RECEIVED THE ONLY GOLD MEDAL GRANTED TO AN AMERICAN LIFE INSURANCE COMPANY AT THE PARIS EXPOSITION OF 1900. Writes one of the Most Liberal Polictes in the World. The Prudential Insurance Company of America JOHN F. DRYDEN, HOME OFFICE: President. Newark, N. J. VOLUME XXXII NUMBER 1 DOTANICAL (aA ZETEE JULY, roor ON THE ORIGIN AND NATURE OF THE MIDDLE LAMELLA. CHARLES BE. ALLEN: THERE has long been recognized in the cell walls of plant tissues, and particularly in the thickened walls of bast and xylem elements in woody plants, a central layer or plate, sharply dis- tinguishable by its optical and staining properties from other regions of the wall. To account for the origin of this layer, apparently homogeneous and equally closely related to each of the cells between which it occurs, various theories have been advanced, and the names “primary cell wall,’”’ “intercellular Substance,” and ‘‘ middle lamella” indicate the diversity of opin- ion that has prevailed as to its real nature. The theories of the origin of this ‘‘ middle lamella,” adopting as most convenient its commonest designation, may (1), generally Speaking, be grouped under three heads: (1) those which hold it to have been originally a common matrix in which the cells were imbedded; (2) the theory that the middle lamella is the Original cell wall, laid down in common by the two daughter cells in the process of cell division, and remaining distinct from the secondary thickening layers deposited upon its either face ; and (3) the view that it is a substance excreted by the daughter cells into a space left between them after their formation, or into a Space formed by their rounding up and drawing apart. The first theory, that the middle lamella is an original com- mon matrix of the cells, was shown to be groundless as soon as I 2 BOTANICAL GAZETTE [JULY any satisfactory investigation of the process of cell division could be made. The notion of the middle lamella as the primary cell wall until very recently was the most prevalent, and is the one still laid down in most text-books of botany. Vines (15) thus states this view : In the development of a tissue, whether by free cell formation or by cell division, septa are formed, that is, walls which are common to contiguous cells; these are very thin at first, and appear under the highest magnifying power as a simple plate. As the walls increase in thickness and acquire a more or less distinctly stratified structure, as seen in transverse section, the network of primary septa stands out from the thickening layers proper to each cell. The primary septum between any two cells is now distinguished as the middle lamella (sometimes also termed intercellular substance); it attains a considerable bulk at points where several septa meet at an angle. This theory was early stated by Strasburger, and until lately he has adhered to it in unmodified:form. In the first edition of the Botanisches Practicum (10), on p. 78, he describes a method of demonstrating the presence of this median layer in the endosperm of the seed of Ornithogalum umbellatum by treatment with sul- furic acid, the delicate network of the middle lamella being the last part of the cell walls to dissolve. This network, he says, corresponds to the original walls present before the process of thickening began. In another passage, on page 82, speaking of the tracheids of Pinus silvestris, he uses the term “ primary wall,”’ which, however, is not the same as the first wall laid down in cell division—the middle lamella— but includes the latter as a very delicate layer (TZhetlplatte). The middle lamella differs from the rest of the primary wall in being “cutinized.” To the original partition laid down in the process of cell division, Stras- burger (9) had in 1875 given the name ‘‘cell plate.” Treub (14), in 1878, studying the living cells of the pro- embryo of Orchis latifolia and the ovules of Epipactis palusirs, found, after its formation, a splitting of the cell plate, and the deposition of a cell wall between the two layers so formed. This is, | believe, the first suggestion that the median layer of the cell wall is not the plate first laid down in the process of cell Igor] THE MIDDLE LAMELLA 3 division, but is a substance secreted into a space between the daughter cells. The history of the cell plate theory has been carefully sum- marized by Timberlake (13), and reference may be made to his paper for the literature of the subject. The results of recent investigations go to show the essential correctness of Treub’s view. In 1898, Strasburger (12) announced that the cell plate splits to form a plasma membrane for each daughter cell. Dur- ing the splitting process, the rod-shaped elements visible in the cell plate, which represent the material furnished to the plate by the spindle fibers, are pulled out so as to become extremely thin in their middle portion. A middle layer then appears between the halves of the original layer. Whether or not the thread-like portions of the rods persist, either as part of this new layer or as protoplasmic connections between the plasma membranes of the daughter cells, he did not determine. So the question was left open whether the substance of the wall arises entirely by Secretion into the space formed by the splitting of the cell plate, or whether a portion of the original cell plate takes part in its formation; but Strasburger has wholly abandoned his old notion of the identity of the middle lamella with the original cell plate. Timberlake (13) finds in the cell divisions of the root tip of Allium Cepa, before the appearance of the cell plate elements, an accumulation of an orange-staining substance in the equatorial zone of the spindle. From the staining reactions of this sub- stance, he concludes that it is some form of carbohydrate, probably destined for the building of the wall between the daughter cells. But, while in general the material of this zone Stains like the cell wall, it is not stained, as is the wall, by either ruthenium red or iron hematoxylin. The cell plate is formed in the midst of this zone by the fusing together of. equatorial thickenings of the spindle fibers. The carbohydrate material disappears with the formation of the cell plate, but very soon after the splitting of the plate the young cell wall (middle lamella) appears in the cleft. The cell plate begins to split in its oldest part, that is, in the central portion of the spindle, so 4 “BOTANICAL GAZETTE [ JULY that a cell wall may be visible here before the cell plate by peripheral growth has reached the mother cell wall. Mangin (3, 4)5, 6, 7, 8), in a series of contributions which appeared from 1888 to 1893, investigated particularly the chemi- cal nature of the substance in question. As this side of the sub- ject and Mangin’s very interesting conclusions regarding it have been little discussed, a somewhat full résumé of his results may here be given. Dippel, quoted by Mangin (3), had already announced that the new partition deposited by the protoplasm has none of the properties of cellulose. Frémy found in the tissues of fruits and roots a substarice called by him pectose, which he was unable to separate from cellulose, and from which ate produced the pectic compounds found in the walls of fruits.: Maudet found in the pith of certain trees pectose and calcium pectate, form- ing a cement which holds the cells together. .Mangin (3) found plant cell walls to be generally formed by the association of two substances, cellulose and one which he provisionally called pectose. Ina large variety of adult tissues he found _pectose, in a pure state, forming the middle lamella or intercellular sub- stance, and associated with cellulose in the other layers of the cell wall. Dippel’s conclusion, that the middle lamella is free from cellulose, was thus confirmed. Some tissues ‘were found to be composed wholly of pectose. It plays the principal réle in what had been called cellulose fermentation, and Mangin con- _ sidered chemical modifications of cell walls, as lignification and cutinization, to be due to transformations of pectose, of which cellulose is incapable. At this time (1888) he repeatedly speaks of the pectose layer as the first membrane formed in cell division, the “fundamental! layer of the cellular membrane.” This seems to indicate an acceptance of the view then! held by Strasburger as to the identity ofthe mss soni with the cell plate. But’ in’ 1890, Mangin (5) suggested the Psuiihraseacsss sot restoring to this layer the name ‘‘intercellular’substance,” as expressing better than that of ‘‘middle lamella” its ‘origin and Igor] THE MIDDLE LAMELLA 5 mode of formation. He found it to consist, not of pectose proper, but of a cement of insoluble pectates. To demonstrate the presence of pectic acid, tissues are macerated in a mixture of alcohol and hydrochloric acid, and then treated with a weakly alkaline solution. The tissues dissociate into their constituent cells and fibers, the intercellular substance passing into solution. The solution is shown by analysis to contain pectic acid, which appropriate staining now shows to be absent from the cell walls. In this process, the insoluble pectates are first changed into pectic acid, which, in the alkaline solution, forms a soluble alka- line salt. If sections first treated with acid alcohol are stained with a pectic stain, as phenosafranin or methylene blue, the pectic acid present stains more deeply than the pectic com- pounds associated with cellulose in the wall layers lying between the intercellular substance and the cell cavity. This indicates that it is not pectic acid and its derivatives, but some of the neutrgl pectic substances, as pectin or pectose, which are in combination with cellulose in the later deposited layers. The intercellular substance thus deeply stained forms a thin layer on the whole surface of contact of adult cells; where the cells draw apart, it produces a thick cushion; when the cells separate so as to form intercellular spaces, these spaces are bounded by a pectic layer. This frame of pectic acid is some- times thickened irregularly, so as to form knobs, points, and sculptures of various forms ornamenting the frame itself or pro- jecting into the intercellular spaces. Sometimes the spaces are partially or completely filled with a jelly-like mass, a soluble transformation product of pectic acid.» In the meristem the intercellular layer is not disclosed by staining, but the chemical teactions of cellulose and pectic compounds are given by the cellmembranes. That a thin layer of intercellular substance ts present is shown by the dissociation of the cells as in older tis- sues after treatment with acid alcohol and an alkaline solution. Mangia explains the variations in thickness and form of the intercellular cement by its partial transformation in the course of the development of the tissues into soluble pectates; this 6 BOTANICAL GAZETTE [JULY makes possible the splitting of the membrane, the formation of intercellular spaces, and an exudation of material to form the knobs, points, and other structures already referred to. A further proof of the presence of pectic compounds in the wall is furnished by treating tissues with a cellulose solvent; the gen- eral outlines of the cells remain, the middle lamella, and often a large part, or in some cases all, of the other wall layers preserving their form. The framework thus left takes up pectic acid stains. and is indifferent to cellulose stains. The best distinctive stain for pectic substances is ruthenium red, the ammoniacal sesqui- chlorid of ruthenium. Our knowledge of the value of this sub- tance is due to Mangin (8), who first described its remarkable properties as a staining reagent for vegetable tissues. Mangin’s attention, it is seen, has been directed to the chemical composition and transformations of the “ intercellular substance,” and he has not connected his valuable results obtained in that field with a study of its origin. It was in part to supplement his work in this important regard that the present investigation was undertaken. Strasburger (12) and Dippel (2) accept Mangin’s conclusions as to the widespread presence and the importance of pectic sub- stances in plant tissues, especially in the middle lamella and the thin layer immediately surrounding the intercellular spaces. Dippel’s account of the history of the middle lamella in the wood and bast of higher plants is briefly as follows (pp. 570 ff.). The radial walls of dividing cambium cells are separated from one another by a lax, weakly refractive Zwischenmasse or Zwischensubstanz. This substance decreases in amount as we pass outward from the cambium layer into the differentiated wood and bast tissues, until it finally disappears, excepting at the angles formed by the junction of three or four cells. Remains of the cambium mother cell walls may occasionally be recog- nized in the Zwischensubstanz, which Dippel believes to be at least partially produced by their disorganization. Chemically, the Zwischensubstanz may consist of a union of pectose and callose, which is soluble under the conditions of cell wall development, Igor] THE MIDDLE LAMELLA 7 and so, for the most part, is absorbed into the cambial daughter walls. The cambial cell walls (p. 575), that is the cell walls of the cambium mother* and daughter cells, are composed of pectic acid, which, at least after the transformation of these walls into an “intercellular substance,” exists largely in the form of’ calcium pectate. As the cambial daughter cells are transformed into bast and wood tissue, the primary cell walls, consisting of pectose and cellulose, are deposited next to the now apparently simple cambial walls. These latter now undergo a transformation, forming an ‘intercellular substance,” or K7¢t- substanz, which, through a loss of water and the extension in length of the radial walls, becomes so thin as to be in most cases invisible except after special treatment. It is, besides, fused into apparently intimate connection with the primary cell walls. This combination of the intercellular substance and the primary cell walls forms the middle lamella of mature tissues, against which in the process of development secondary thickenings are deposited. The continued existence of the “intercellular substance” as a middle plate ( Theilplatte) of the middle lamella Dippel demonstrates by the action of the wall layers upon polarized light, as well as by treatment with various macerating and staining reagents, notably ruthenium red. Dippel’s peculiar view, developed in connection with his studies on the algae, that cell division is.accompanied by the formation of a new cell wall entirely enclosing each daughter cell, has doubtless influenced his account of the development of the middle lamella in the higher plants. For if this were the case in the predominantly radial growth of cambium cells, the result would be to leave, between the radial walls of the newly- formed daughter cells, the old mother cell walls, which, if they became disorganized, would form a Zwischensubstanz such as Dippel describes. The tenability of this view of the origin of the Zwischensubstanz will be discussed hereafter. *Here there seems to be a contradiction, for Dippel has already spoken (p. 575) of the cambium mother cell walls as being disorganized toform the Zwischensubstanz of the daughter cell walls. 8 BOTANICAL GAZETTE [JULY MATERIAL AND METHODS. Woody tissues of the following species were studied: Preris aquilina L., Pinus silvestris L., Nerium Oleander L., Kosa sp., Tilia Americana 1.., and [lex opaca Ait. Of these, Mangin had investigated tissues of Pinus and Ilex. Free-hand sections were cut, usually both transverse and longitudinal. In many cases, before staining, they were treated for twenty-four hours,as suggested by Mangin (5), with a mixture of one part of concentrated hydrochloric acid and four or five of absolute alcohol, then washed in distilled water. The stains characteristic of pectic acid, especially ruthenium red, are taken up, as a rule, more freely after this treatment, and almost invari- ably the coloration is more clear-cut and distinctive. For staining, Griibler’s preparations were used. Unless other- wise stated, they were in neutral saturated solution. When an alkaline solution was used, it was prepared by adding to the neutral solution about % per cent. of concentrated ammonia solution. Acidulated solutions were prepared by adding a fe like proportion of acetic acid. In some cases where saturated ae solutions, ¢. g., of methylene blue and methyl violet B, were too strong for good results, they were diluted to such a strength that a few seconds’ exposure gave a differential stain. Aniline- | water-safranin was prepared as directed by Zimmermann (16). | An aqueous solution of ruthenium red of a strength of about one five-thousandth was used. An increase in strength above this did not seem to make any noticeable difference in the depth of the stain or in the time of exposure required. This substance is reduced by the action of light in the presence of water, hence the solution must be kept in the dark. It was tried in neutral, alkaline, and acidulated solutions, the alkaline and acidulated solutions being prepared as described for the other coloring matters, substituting hydrochloric for acetic acid. In all cases, the alkaline solution was found to give the deepest and most distinctive colorings, and it is this preparation of ruthenium red that is referred to, hereafter. But a few minutes’ exposure to the stain is required. Igor} THE MIDDLE LAMELLA 9 In general, sections after staining were mounted in water and studied immediately. Ruthenium red preparations can be dehy- drated by absolute alcohol, passed through clove oil, and mounted in Canada balsam without affecting the stain. But the color gradually fades out from sections so treated, some tissues, notably the xylem and bast, losing their color almost entirely within a few weeks, while others, as the parenchyma and pith, retain it much longer. Sections stained with ruthenium red and mounted in glycerin jelly lose their color almost entirely in the course of a few hours. Mangin (4, 6, 7, 8) has classified the coloring matters used in staining plant tissues, and has described their reactions upon the various substances found in cell walls. Of the great num- ber of stains named by him, the following were employed in this investigation: | Orseille, which colors cellulose in neutral or slightly acidu- lated solution, but does not affect callose. Aniline-water-safranin (phenosafranin), methylene blue, Bis- marck brown (Vesuvian brown), fuchsin, and methyl violet B (violet de Paris), which do not color callose or cellulose, but do color pectic acid in neutral or acidulated solution. They also color equally well nitrogenous substances (lignin, cutin, etc.), but these may be distinguished by their retention of the color after treatment with alcohol, glycerin, or acids, which treatment decolorizes pectic acid. According to Strasburger (11), safranin colors the protoplasmic contents of the cell, the lignified wall, cork, and cutinized membranes cherry-red, the pectic substances Orange-yellow. Methylene blue colors the protoplasmic con- tents and lignified walls blue, the pectic substances violet-blue. Acid brown, nigrosin, and ponceau, which do not color pec- tic compounds, but strongly color nitrogenous substances. By mixing one of this series with one of the preceding, Mangin obtained distinctive double stains. Ruthenium red, which Mangin found the most satisfactory distinctive stain for pectic compounds. It differs from the other pectic stains, as safranin and methylene blue, by the fact that 10 BOTANICAL GAZETTE [JULY pectic compounds stained by it are not decolorized by glycerin or alcohol. Ruthenium red, he finds, also stains gums and mucilages formed by the decomposition of pectic substances, but does not affect decomposition products of cellulose or cal- lose. Lignified tissues, not stained by ruthenium red when fresh or preserved in alcohol, take it up after treatment with alkalies or Javelle water; but their affinity for ruthenium red is always less than for certain basic organic stains, such as methyl- ene blue, so that by combining its action with that of one of the latter class fine double colorations may be obtained. Cuti- nized membranes, according to Mangin, are stained in many cases, but not the cuticle. Ruthenium red also stains in vary- ing degrees the protoplasmic cell contents. PTERIS AQUILINA. The rhizome was studied-in cross section. In unstained sec- tions the middle lamella of the sclerenchyma walls is plainly distinguishable from the inner layers by its greater density and refractive power. It appears as a dark yellowish-brown layer constituting a considerable proportion of the total thickness of the wall, enlarged at the angles where three or four cells abut into a triangular or quadrangular form, often enclosing at such places a similarly shaped intercellular space. In the walls of the stone cells, while the absolute thickness of the middle lamella is approximately the same as in the » sclerenchyma, its thickness relatively to that of the whole wall is of course considerably less. Otherwise its appearance is much the same as in the sclerenchyma. The canals appear as trans- verse lines seeming to pass quite through the wall, including the middle lamella. On treatment with methylene blue, the sclerenchyma wall is colored green, except the middle lamella, which, if stained at all, yet appears yellowish-brown in contrast with the other lay- ers. The walls of the fundamental parenchyma, as well as the protoplasmic cell contents, are stained blue. The parenchyma walls are apparently continuous with, and of the same thickness Igor] THE MIDDLE LAMELLA 11 as, the middle lamellae of the sclerenchyma. The green appear- ance of the sclerenchyma walls is probably not a differential col- oring, but is due to a combination of their original yellowish- brown with the blue stain. In sections treated with ruthenium red, the sclerenchyma and stone cell walls do not take up the stain to any appreciable extent. The walls of the fundamental parenchyma are clearly brought out, and are again plainly continuous with the middle lamellae of the sclerenchyma and stone cells. The indifference of the middle lamella in these thickened walls to methylene blue and ruthenium red, although this layer is directly continuous with the thin, readily-staining parenchyma wall, is in marked contrast to its character in the tissues of the spermatophytes Studied; it may be due to a change in the chemical composition of the middle lamella which occurred simultaneously with the deposition of secondary thickening layers. The fundamental parenchyma walls show no evidence of secondary thickening. They are stained by ruthenium red deeply and quite uniformly, Save that at the angles the wall is thicker, the triangular or quad- rangular area so formed is less deeply stained, and contains apparently a less dense substance than that forming the remain- er of the wall. This less deeply stained area corresponds to the thickened corners and the intercellular spaces of the scleren- chyma. The protoplasmic contents of the parenchyma cells are stained unevenly, the nuclei very deeply. In the vascular bundles, the parenchyma walls are deeply and uniformly stained by ruthenium red. These walls appear con- tinuous with the stained middle lamellae of the large vessels and Sieve tubes. In the walls of the vessels the middle lamella is relatively thin; the rest of the wall is uncolored. The middle lamella of the sieve tubes occupies a larger proportion of the wall thickness, the interior unstained layers being very thin. At corners formed by three or four cells the middle lamella appears more dense.and somewhat, though not greatly, enlarged. The staining reaction of the middle lamella in these elements is in contrast to its conduct in the sclerenchyma and stone cells, 12 BOTANICAL GAZETTE [JULY and agrees with results generally noted in tissues of spermato- phytes. PINUS SILVESTRIS. Sections of young twigs and branches cut in the fall were used ; these, therefore, were past the season of rapid cam bial division, and the cambium cells were in a condition of compara- tive rest. Sections of old and seasoned pine wood were also used for comparison with the young tissues. In cross sections through the cambium and neighboring tissues of young wood, treated with acid alcohol as above described, the cambium walls are stained throughout their thick- ness by methylene blue a deep and apparently uniform blue. The result is similar in sections not previously treated with acid alcohol. Similar cross sections, both in the fresh condition and after treatment with acid alcohol, were stained with ruthenium red. This stain is also taken up freely by the cambium walls. The radial walls are noticeably thicker than the tangential. In the middle of the radial walls there is often a less deeply staining layer, recognizable as Dippel’s Zwischensubstanz. The rest of the walls are deeply stained, except that in many of the older cambium cells, whose walls are thicker than those of the young- est cells present, there is, next the cell interior, a light line indi- cating a later deposited, unstained, or less deeply stained wall layer. The corners where radial and tangential walls join are especially deeply stained. In many cases, a continuation of the radial cambium wall into the middle lamella of the xylem and bast can easily be traced. A difference in depth of stain between radial and tangential walls similar to that given by methylene blue is found in the longitudinal sections treated with acid alcohol and exposed for a few minutes to the action of Bismarck brown. In cross sections treated with acid alcohol and Stained one hour with fuchsin, the cambium walls are uncolored; the proto- plasmic contents of some of the cambium cells are stained red. The difference between the affinity of these walls for Bismarck Igor | THE MIDDLE LAMELLA 13 brown and fuchsin is worthy of note; both these stains belong to the class in which a basic coloring matter is united to an acid; according to Mangin, all the stains of this class stain pectic compounds and do not color cellulose or callose. In unstained cross sections through the xylem, the middle lamella is to be distinguished from the rest of the wall by its different refractive power, at some levels of focusing appearing brighter, at other levels darker, than adjacent layers. The same is the case in cross sections through old pine wood, except that here the middle lamella appears, if anything, more dense than in the young wood. In cross sections through young wood treated with acid alcohol and methylene blue, the whole xylem is colored green. The middle lamella is distinguishable only by its greater density, as also in sections not previously treated with acid alcohol. Xylem walls in cross sections treated with acid alcohol stain more deeply with ruthenium red than in sections not so treated. The middle lamella stains much more deeply than the inner layers, the boundary between the lightly and deeply stained por- tions being sharply marked. The difference in depth of stain is much more apparent in acid alcohol sections than in the others. The tori of the pits in acid alcohol sections are stained deep red; this occasionally occurs also in sections not treated with acid alcohol. In some cases in acid alcohol sections, the torus appears plainly to be a continuation of the middle lamella. In some cases also the middle lamella appears continuous with the interior layer of the pit cavity, which stains deeply. The middle lamella is more plainly differentiated in xylem of the present year’s growth. In some preparations the oldest cells of the present year’s growth (the spring wood) have the whole wall quite deeply stained. These facts may indicate that it is in the younger xylem walls that pectic substances exist in purest form. Where splitting of the walls occurs in preparing the section, it is sometimes on the line of demarcation between the middle lamella and the less stained layers; very often, how- ever, the split is approximately through the center of the middle 14 BOTANICAL GAZETTE (JULY lamella. Intercellular spaces occur in the xylem where three or four cells abut; they appear as triangular or quadrangular breaks in the uniformly stained middle lamella. Comparing the results obtained from methylene blue and ruthenium red, we find that by both the whole xylem wall is stained, the middle lamella being differentiated by the red and not by the blue. Since methylene blue stains nitrogenous sub- stances equally well with pectic acid, and does not stain cellulose or callose, while ruthenium red stains pectic derivatives and not nitrogenous substances, it appears that the secondary layers of the xylem, as well as the thin layer of the cambium wall which does not take up ruthenium red, contain a mixture of nitro- genous and pectic compounds. The use of orseille, a cellulose stain, gave no satisfactory results, showing that celulose, if pres- ent at all in these walls, is of minor importance. The staining power of the tori of the pits indicates that they represent in the main the same layer as the middle lamella; this does not, how- ever, preclude the possibility of their also possessing a thin, subsequently deposited layer. The deeply stained layer lining the pits is an example, of which others will be mentioned later, of the possible deposition of a pectic layer very late in the his- tory of the growth of the cell wall. The relations of the middle lamella, tori, intercellular spaces, and thickening layers appear about the same in older pine wood stained with ruthenium red as in the young tissues already described. In cross sections previously treated with acid alcohol and exposed for one hour to fuchsin, which belongs to the same class as methylene blue, the xylem walls are stained throughout. Other tissues, except the contents of the cambium and collen- chyma cells, are unstained. Cross sections of older pine wood were left in ponceau solu- tion for three or four days. The xylem is quite uniformly stained vermillion, the middle lamella generally appearing darker than the inner layers of the wall, but the distinction of coloration is not strongly marked, and the difference may be wholly due to the difference in density. : ; 1901] THE MIDDLE LAMELLA 15 The xylem walls of young wood are not stained after a half- hour’s treatment with acidulated nigrosin solution. This fact, by itself, would mean the absence of nitrogenous substances ; but, as against that furnished by methylene blue, and by ponceau, which belongs to the same class as nigrosin, such negative evi- dence is of little value. In longitudinal sections of young tissues treated with acid alcohol and stained with ruthenium red, the tori of the pits are stained deep red. The appearance of the tori and of the red- staining bars between the pits, which bars are referred by Dippel (2) to the remains of the Zzischensubstanz, is well shown by him in his figure 397. In many cases, probably in partly developed pits, the stain is only a deep spot at the center. In some cases, an unstained center is surrounded by a stained ring. The pits are not so noticeable where they connect with the cells of the medullary rays, but where the ring appearance just described is found in older wood, the tori of pits so situated are completely stained. Both the walls and the cell contents, especially the nuclei, of the medullary rays are deeply stained by methylene blue, both in cross sections previously treated with acid alcohol and in those not so treated. This is true of both young and old wood. Similar results are effected by ruthenium red. In cross sections of old wood stained with ruthenium red and then exposed for a few seconds to methylene blue, the red stain of the xylem is replaced by the blue, while, if the action of the blue has not been too long continued, the medullary rays remain red. The greater affinity of the xylem walls for the blue than for the red is due probably to their containing nitrogenous and pectic compounds, both of which take up methylene blue, while the. former, unless Specially treated, have no affinity for ruthenium red. N cross sections treated for one half hour with acidulated nigrosin solution, the medullary rays and their cell-contents take up the stain freely. In cross sections of older pine wood treated for three or four days with ponceau, the medullary rays are unstained. 16 BOTANICAL GAZETTE [JULY The readiness with which the walls of the medullary rays take up methylene blue and ruthenium red, and also nigrosin, indi- cates the presence in them of both pectic and nitrogenous sub- stances. Their preference for ruthenium red indicates a greater proportion of pectic constituents than in the xylem walls. But against this view is the negative evidence of the ponceau, though this, perhaps, is not of much weight. The cells about the resin pits and their contents, if any, are always deeply stained by either methylene blue or ruthenium red. This indicates, according to Mangin, that the resin and other substances formed in the breaking down of these cells are wholly or chiefly decomposition products of pectic acid and its derivatives. The staining of the bast is in general similar to that of the xylem. In both acid alcohol and fresh sections, the whole wall takes up more or less freely either methylene blue or ruthenium red, the middle lamella being much more deeply stained than the inner layers. The middle lamella, of course, is thicker rela- tively to the total thickness of the wall, but is of about the same absolute thickness as in the xylem. The middle lamella of the collenchyma walls is generally stained more deeply than the other wall layers by methylene blue or ruthenium red, especially in sections treated with acid alcohol. Here, except at the corners, the unstained portion of the wall is very thin, and often the whole wall appears quite uniformly stained. The stain appears more diffuse than in the xylem, and the distinction between the more and less deeply stained layers is not so sharp. The middle lamella is enlarged at the angles of the cells, and sometimes encloses a triangular or quadrangular intercellular space. The appearance of the collenchyma is very much the same in longitudinal as in cross sections. The collenchyma walls become red when treated for thirty minutes with acidulated orseille solution, indicating the presence of some cellulose, but no distinctive stain for different portions of the wall is observable. The walls, except the outer ones, of the epidermal cells in Igor | THE MIDDLE LAMELLA 17 either acid alcohol or fresh sections are deeply stained by ruthe- nium red throughout their whole thickness, but the cuticle remains unstained. This accords with Mangin’s results on cutinized membranes. NERIUM OLEANDER. Cross sections were cut through growing stems of various ages. The whole of the cambium wall is deeply and quite uniformly stained by ruthenium red either after or without previ- ous treatment with acid alcohol. Similar results are attained by a few seconds’ exposure to dilute methylene blue solution. The cambium walls are not stained by an exposure of three or four minutes to acid brown. The middle lamella of the xylem is deeply stained by methylene blue or ruthenium red, the other layers of the wall being colored very slightly if at all. The xylem walls are colored blue by methyl violet B. They are Stained quite uniformly brown in a few minutes by acid brown. They are stained uniformly red in forty-five minutes by aniline- water-safranin. After thirty minutes’ exposure to this safranin solution the xylem walls are colored bright red, the middle lamellae between the large tubes being decidedly darker than the thickening layers. After forty minutes’ exposure to ponceau, the thickening layers of the xylem are stained somewhat, though not evenly, the middle lamella remaining uncolored. Sections were exposed for twenty-four hours to neutral, acidulated, and alkaline solutions of orseille. The neutral and alkaline solutions produced the same coloration of the various tissues as the acidulated, but to a less degree. Therefore only the action of the acidulated solution will be described. The ‘coloration of the xylem and medullary rays isa purplish-red, of all the other tissues affected, brown. The walls of the xylem have a general purplish tinge, but no distinctive staining is apparent in the different layers. The contents of the medullary ray cells are deeply colored, and the walls are stained similarly to those of the xylem. An exposure of two and one half hours to ponceau produces a reddish tinge in the xylem walls, and no further stain. 18 BOTANICAL GAZETTE [JULY The contents of the medullary ray cells are stained by acid brown. Both the walls and the cell contents are deeply stained by methylene blue as well as by ruthenium red. The results as to the tissues just described agree closely with those obtained for Pinus. The cambium walls and the middle lamellae of the xylem and medullary ray walls are of pectic nature; the remaining layers of the xylem and medullary ray walls are composed of a mixture of pectic and nitrogenous sub- stances, the pectic constituents being perhaps predominant in the medullary rays; and there is evidence of the presence of a small proportion of cellulose in these thickening layers. The bast fibers, when treated with ruthenium red, show a peculiar and most beautiful differential stain. It is most clearly brought out after treatment with acid alcohol, though it appears in sections not so treated. The middle lamella, which is rela- tively very thin, is deeply stained. It is sharply distinguished from the layer next within it on either side, which is colorless or very slightly stained. Next, passing toward the interior of the cell, comes a layer with a little color, then one a little more deeply stained, and so on, the depth of stain increasing until the last thickening layer, reaching almost to the center of the cell, is colored to about the same degree as the middle lamella. It should be noted that the stain shown in these walls is rather a purplish-red than the bright red commonly found in ruthenium red preparations. This behavior with ruthenium red is markedly different from that shown by Dippel (2) in his figure 139, in which all the wall of the bast fibers except the middle lamella is shown as unstained. On page 219, he mentions, as an instance of pectose-free cell walls, the bast fibers of Merium Oleander. This is plainly an error. The bast fibers are not stained at all by exposure for a few seconds to methylene blue. By methyl violet B (three or four minutes), the middle lamella of the bast and the other wall lay- ers except those next the middle lamella are stained violet. Acid brown has an effect exactly opposite to that of ruthenium red; that is, the middle lamella is not stained; the next layer is 1901] THE MIDDLE LAMELLA tg quite deeply colored, the next is lighter, and the colors grow less and less intensé until the innermost layers show no coloration at all. After a ten minutes’ exposure to acid brown, sections were dehydrated, passed through clove oil, and mounted in Canada balsam, without affecting the staining, except that the color of the bast fibers was somewhat faded. The contents of the bast fibers are deeply colored by acid brown. The action of orseille (acidulated solution, twenty-four hours exposure) is exactly like that of acid brown. The effect of aniline-water-safranin, on the other hand, is similar to that of ruthenium red. Exposure to ponceau for various periods up to two and one half hours does not stain the bast. By treatment for a few minutes with methyl violet B, the middle lamella of the bast and all the other layers except that next the middle lamella are stained violet. It appears that in the bast, as generally elsewhere, the mid- dle lamella is of pectic nature; that the first of the secondary layers is nearly or quite free from pectic substances, and is com- posed of a mixture of cellulose and nitrogenous substances, and that the subsequent layers gradually increase in pectic content at the expense of the cellulose and nitrogenous constituents, until the last layer is perhaps again purely pectic. But, from the purplish-red color given by ruthenium red and the indiffer- ence shown toward methylene blue, it may be inferred that the pectic compounds found in these walls are not exactly the same as those common to the middle lamellae of other tissues. These inner layers, like the pectic lining of the canals in the xylem of Pinus, show that the ability of the cell to secrete pectic acid is not limited to a single period of its development. Acid alcohol sections treated for a few seconds with methyl- ene blue show a deep blue color in the middle lamella of the collenchyma walls, much deeper than in the remaining portions of the wall, which, however, are also somewhat stained. The boundary between the middle lamella and the next adjoining layers is not so sharply defined as in the xylem and bast. At 20 BOTANICAL GAZETTE [JULY the corners of the cells there is an angular, less deeply stained area enclosed by the middle lamella, just as in the fundamental parenchyma of Pteris. The effect of ruthenium red on the collenchyma is similar to that of methylene blue. The collenchyma walls are not stained by acid brown (three or four minutes), but the cell contents are deeply colored. The contents are also stained brown by acidulated orseille solution, and the walls show a brownish tinge, but no distinctive stain for the different layers. The cell contents are stained by ponceau (forty minutes), but not the walls. Acid alcohol sections treated for thirty minutes with aniline- water-safranin show a deeper stain in the middle lamella of the collenchyma than in the other wall layers. The walls of the cork cells are not colored by methylene blue nor by acidulated orseille solution. Orseille also does not stain the cuticle. In acid alcohol sections the middle lamella of the cork cells is stained more deeply by aniline-water-safranin (thirty minutes) than is the rest of the wall. The pith cell walls take up ruthenium red freely, the middle lamella, which occupies the greater portion of the thickness of _the wall, being deeply stained, especially at the corners. Some- times the whole wall appears quite uniformly stained. The intercellular spaces are sometimes angular, but very often rounded, elliptical, or circular. The contents of pith cells are stained by acid brown, the walls unstained. The action of orseille is similar. The failure of these two stains to affect the pith walls indicates that those walls are composed chiefly of pectic compounds, probably in purest form in the middle lamella. ROSA SP. In general, the results were the same as for Nerium. Cross sections were treated with acid alcohol and stained with ruthenium red. The cambium walls stain quite uniformly 1901} THE MIDDLE LAMELLA 21 throughout. The xylem shows a distinctive stain for the middle lamella, which is enlarged and especially deeply stained at the angles. The medullary ray walls and cell contents stain deeply. The middle lamella and successive thickening layers of the bast fibers stain exactly as described for Nerium, including the purplish tinge. The stain of the middle lamella is deepest at the angles. The lining layer of the canals running from fiber to fiber is deeply stained where it traverses unstained or less deeply stained layers. This recalls the deeply stained layer of the pits of pine tracheids, and, with the deep stain of the last - deposited wall layer in these tissues of the rose, shows these cells to possess the power very late in their history of depositing pectic wall material. The pith cells show a very distinctive stain for the middle lamella. In the collenchyma, the walls are stained throughout, the middle lamella much more deeply than the other layers, but the boundary between more and less deeply stained portions is not so clearly defined as in the xylem. The epidermal walls are Stained about the same as the collenchyma. The cuticle is unstained. Cross sections of a slightly older stem than that just described were stained with ruthenium red, then exposed for a few seconds to methylene blue. The xylem walls are stained purple by the combination, the middle lamella most deeply. The bast fibers take up both stains, the relative depth of stain of the various layers being about the same as with the red alone. The epidermal walls take up the blue more freely than the red. The collenchyma cells remain red. Cross sections of a very young growing shoot were treated with acid alcohol and stained with ruthenium red. The cambium walls stained deeply throughout. Very thin tangential red lines are visible, representing the earliest deposit of wall material. Such very thin tangential walls are what one would expect to find in material gathered in the season of active cambial growth. Young xylem elements have the walls entirely stained, 22 BOTANICAL GAZETTE (JULY showing a long-continued period of pectic deposition. In older fibers a middle lamella is deeply stained, the remaining layers very slightly or not at all. The bast fibers are stained throughout; no thickening layers are yet distinguishable. The middle lamella is already differen- tiated in the collenchyma, the other layers remaining uncolored. The middle lamella is especially deeply stained in the corners, and angular intercellular spaces are in some cases inclosed by it. Similar sections not treated with acid alcohol yield similar results. TILIA AMERICANA. Cross sections through rather young growing shoots were treated with acid alcohol and ruthenium red. _ The cambium walls stain deeply, in the youngest cells throughout their entire thickness. The radial walls are more deeply stained than the tangential, the corners especially deeply. Passing from the youngest cambium cells toward the bast or toward the xylem, the beginnings of the secondary thickening may be seen in a very thin unstained or only slightly stained layer of the wall within the stained portion. This is similar to the behavior of the cambium walls of pine, except that in Tilia, as generally, there is no evidence of a Zwischensubstanz, which, so far as I have observed, is found only in the pine cambium walls, The continuation of the deeply stained layer can be traced in the middle lamella of the much thickened xylem and bast elements. The middle lamella of the xylem stains deeply, especially at the angles, where it is enlarged. The rest of the wall is slightly tinged with red. The contents of the medullary ray cells are stained. Their walls are more uniformly stained than those of the xylem ele- ments, but the middle lamella is still plainly distinguishable. The bast has a deeply stained middle lamella, the other layers of the very much thickened wall being unstained or only slightly stained. In this respect, as in most others, the staining reactions of Tilia resemble those of pine. Pk 1901] THE MIDDLE LAMELLA 23 The walls of the parenchyma cells between the bast bundles show a stain throughout, the middle lamella appearing thin and deeply stained, and somewhat enlarged at the angles. In the collenchyma the whole wall and the cell contents are stained. Here, too, the middle lamella is thin and deeply stained, often including at the angles a less deeply stained area. The walls of the cork cells stain quite uniformly throughout, but not very deeply. The color is purplish, different from the bright red typical of the middle lamella in other tissues. No middle lamella is here distinguishable. If anything, the inner- most wall layer, next the lumen, is most deeply stainéd. Here, as in some of the tissues of Pteris, there is evidence that the middle lamella has undergone a chemical change, some or all of its pectic content having been replaced by other substances. The cuticle is unstained. ILEX OPACA. The tissues of the holly resemble those of the pine in their reactions to ruthenium red, the only stain I have used upon them. In cross sections, either with or without previous treatment with acid-alcohol, the cambium walls are stained throughout by ruthenium red, and their connection may be traced with the mid- dle lamellae of the xylem and bast. The middle lamella of the xylem is thickened at the corners and stains deeply; the rest of the wall stains but slightly. The middle lamella of the pith, which takes a deeper stain than the rest of the wall, is thickest at the angles, where it fre- quently encloses intercellular spaces. These spaces are some- times triangular or quadrangular, but usually with more or less rounded angles, and often of an elliptical or circular form, The middle lamella of the bast stains deeply, the rest of the wall slightly. The same is true in the collenchyma walls, where frequent angular intercellular spaces occur. The walls of the cork cells show no stain. 24 ‘ BOTANICAL GAZETTE [JULY DISCUSSION, In almost every tissue studied in which there was evidence of the deposition of wall layers at different periods, the staining reactions of the middle lamella showed it to differ in chemical composition from the adjoining layers; my results confirm Man- gin’s in every respect upon this point. Its distinctive character was brought out most clearly by ruthenium red. The reasons for concluding that, in general, this peculiarly staining layer is com- posed of pectic acid and its derivatives, have already been given at length. It does not follow, however, that its chemical com- position is unchangeable. On the contrary, it is clear that, dur- ing cell growth and development, changes occur in the chemical constitution as well as in the form and mass of the middle lamella. In pointing out the existence of a Zwischensubstanz between the radial walls of the cambial cells, and in distinguishing this layer from the ‘intercellular substance,” either as characterized by Mangin or by himself, Dippel has contributed essentially to our clear conception of the conditions in the cambium of the pine; but his notion that the Zweschensubstanz is derived from the walls of the cambium mother cells has little evidence in its favor. I have never seen in this mass the fragments of these walls of which he speaks (2, p. 575). Besides, were Dippel’s view cor- rect, the tangential walls would be expected to contain at least as much of this substance as the radial walls; but, in fact, noth- ing of the sort is to be seen between the tangential walls, which, however, as shown in Dippel’s own figures, are no thicker than are the radial walls without the Zwischensubstanz. On this point of the origin of the Zwschensubstanz Dippel is not clear, for he distinctly says (p. 575) that it is derived from the degenerated cambium mother cell walls, and again, on the same page, he speaks of the latter as going to make up the intercellular sub- stance. It seems to me more probable that this Zwischensubstanz represents pectic acid which has exuded through the cambium cell walls into an intercellular cleft formed by the splitting of the radial walls, and that it is analogous to the lax substance 1901] THE MIDDLE LAMELLA 25 sometimes found at the angles of cells before the appearance of empty intercellular spaces. Its ultimate fate is doubtless to be absorbed into the adjacent cell walls, so that it does not appear as a visible layer in the middle lamella of mature tissues except in the case of the areas represented by the red-staining bars of the pine tracheids (see Dippel’s figure 397), and except also, perhaps, on the surface of intercellular spaces. The intercellu- lar spaces of the pine wood, then, are to be interpreted as remnants of the radial clefts between the cambium cells, rather than as newly formed in the adult tissue. But this does not pre- clude the probability of changes in the form and extent of the Spaces due to the rounding up of the adjacent cells. It would be interesting to know whether this Zzwischensubstanz is of wide occurrence. I have not observed it elsewhere than in the pine, and Mangin does not specifically note its existence. A fact which is important as showing the plastic nature of the middle lamella at an early period of its history is its varia- tion in thickness in different portions of older walls. This is shown commonly by an enlargement at the angles anda decrease in proportional thickness at the sides of the cells. Such varia- tion is in marked contrast in the spermatophytes studied to the practical uniformity in thickness of the completely stained cam- bium walls. These are continuous with the middle lamella of the differentiated tissues, and, were it not for this change in form, would appear identical with the middle lamella. In some of the tissues examined, the substance of the middle lamella appears more dense at the enlarged angles, and seems to take up the characteristic stains more freely at those places. This appearance, however, may be accounted for by the optical effects produced by the greater thickness at the angles. But in other Cases, as in the fundamental parenchyma ot Pteris and the collenchyma of Nerium, these enlarged angles enclose a less deeply staining substance, or are becoming empty. These may be considered as stages in the development of the intercellular Spaces, which are located, like the areas of less deeply stained substance just mentioned, at the angles formed by the junction 26 BOTANICAL GAZETTE (JULY of three or four cells, are enclosed within the apparently split middle lamella, and are commonly angular in form. Since there i§ no trace, except in the pine, of a Zwzschensub- stanz whose absorption would leave intercellular spaces, such spaces in the tissues of other plants can be accounted for only by the rounding up and drawing apart at their corners of adjoin- ing cells. This induces a splitting of the middle lamella. The cleft so formed may be temporarily filled by a lax, fluid or semi-fluid substance, a solution, perhaps, of pectic acid or one of its trans- formation products, but this in time is absorbed into the walls, leaving an empty space. The surfaces of the wall exposed to the so-formed intercellular space may become further modified chemically, and even so softened as to flow enough to form a rounded instead of an angular space; but there is no evidence that such chemical changes ever occur except where the wall is exposed in an intercellular space, and it is more likely that the plasticity here displayed by the pectic acid has characterized it from the time of its deposition, but has not been shown so plainly because of its confinement within comparatively rigid limits. On the basis of the later investigations respecting the rela- tion of the cell wall to the cell plate, we have seen that the middle layer appearing after the splitting of the cell plate is to be considered as formed by deposition from the split halves of the original plate; the middle lamella of mature tissues would include, then, in, addition to possible later deposits, both the layers deposited on the inner surfaces of the daughter plasma membranes. This being the case, we might expect to find in the history of the development of the middle lamella evidences that it consists of two layers. This is just what is found in the case of the intercellular spaces, which are very evidently caused by a split through the center of the middle lamella. If the middle lamella were not of a double nature, we should hardly expect it always to split through the middle, but, in view of its marked difference from the adjoining layers, we should expect that sometimes the whole layer would be pulled to one side or the a ee ea tm hee Igor | THE MIDDLE LAMELLA 27 other as the cells round up and draw apart at their corners. As noted above, in sections split in cutting, there were also frequent cases where the middle lamella split rather than tore away from the other layers. The splitting in all these cases seems to indi- cate a weakness of cohesion in the plane between the two layers first deposited by the plasma membranes; and confirmation is thus given of the view advanced by Strasburger and confirmed by Timberlake that the cell plate splits before the new cell wall is laid down, the latter thus having a double nature from the start. We should not overlook the possibility that the very first layer which appears between the daughter plasma membranes is so thin as not to form a noticeable fraction of the thickness of the mature lamella; and that the splitting at a later stage in the middle lamella’s history is due to a decomposition of this first thin layer, or that the split is really on one side or the other of it. I have, however, seen no evidence in support of either of these hypotheses. It cannot be supposed that the middle lamella consists only of the material first deposited from the young plasma mem- branes. Its thickness in adult cells, and its varying thickness in different tissues of the same plant, ¢.g., the large vessels and the sclerenchyma of Pteris, at once negative such an assump- tion. The older view, that it was the cell plate directly meta- morphosed into cell wall material, is just as effectively negatived - by the same considerations, which should have had more weight with the earlier observers. The middle lamella is rarely difficult of differentiation in adult thick-walled tissues simply by its greater density and less apparent stratification. Moreover, its growth in thickness can often be traced, for it stains continuously from the youngest wall (the thin tangential red lines in the cam- bium of the rose) to the middle lamella of mature tissues. We must conclude that the middle lamella consists of the layers first deposited by the plasma membranes plus a certain amount of material subsequently deposited in contact with these layers, which is generally rich in pectic compounds as compared with still later deposited strata. The middle lamella may vary 28 BOTANICAL GAZETTE [JULY considerably in thickness in different tissues, according as a greater or less amount of material rich in pectic substance has been deposited inthe cell. That it is not the only part of the cell wall, however, that contains pectic compounds is shown from the taking up by the other layers of the wall, though less freely than by the middle lamella, of pectic acid stains; it is shown also in the pectic layers lining the pits of the pine and the canals in the bast fibers of the rose, and in the innermost thickening layers of the bast of Nerium and rose. Although the middle lamella usually retains its pectic nature, it undergoes a change by which, in the course of cell develop- ment, it loses the power of adapting itself to the varying form of the adjoining cells and becomes fixed and inflexible. The evi. dence seems satisfactory that this change, as Mangin suggests, is one from pectic acid to insoluble pectates, chiefly the calcium salt. Suchachange is indicated by a less deep staining in older tissues, unless they are first treated with acid alcohol. A further transformation appears to take place in the cork cells of Tilia, by which the power of distinctive staining is entirely lost, and the middle lamella cannot be distinguished from the other layers of the suberized wall. The pectic acid here seems to have been partly replaced by suberin. The coloration of the cork walls is purplish, indicating a possible similarity in chemical nature to ’ the later layers of the bast elements in Nerium and rose. In the sclerenchyma and stone cells of Pteris, also, though a middle lamella is plainly present, its staining reactions do not distinguish it from the rest of the wall; yet it can be traced as a continua- tion of the thin, characteristically-staining walls of the funda- mental parenchyma. It is possible that the middle lamella in this case was from the start non-pectic ; but it seems more probable that, as secondary thickening proceeded, the pectic acid was replaced or masked by material similar to that deposited to form the other layers. Further light could be thrown upon this ques- tion by tracing the development of the wall from the meristem to the mature tissues. The difference in depth of stain between radial and tangential walls in Tilia may indicate that the tangential 1901 | THE MIDDLE LAMELLA 29 walls in this case are not composed of pure pectic acid, or that it exists in them in a somewhat different form. Such a dif- ference in chemical nature might easily occur, since the tangen- tial walls are being laid down anew, while the radial walls are older, and at most are simply added to as the cells increase in number and size. There is no evidence of the existence here of a mass corresponding to Dippel’s Zwéschensubstanz. The power of the cell to secrete cell wall materials of very different chemical composition at different periods in its history is much greater even than is indicated by Mangin’s work. That the compounds stained by ruthenium red are not derived from the decomposition of a previously deposited cellulose wall is proved by the complete staining of the young cambium wall. Still, that there is a possibility of a closer relation than Mangin supposes between cellulose and other cell wall materials on the one hand, and pectic substances on the other, perhaps even involving transformations of material from one class into the other, is to be inferred from the action of ruthenium red upon the walls of the cells surrounding the resin pores in pine. But that the power of secreting pectic compounds is not limited to a single period in the history of the cell is shown by the changes already discussed in the mass of the middle lamella. Since the very youngest and thinnest cambium walls are stained by ruthenium red, and at a later stage the thicker walls are still completely Stained, the power of depositing pectic acid must last for a con- siderable time. That it continues in some cases even beyond the cambium stage is indicated by the variations in the thickness of the middle lamella in older tissues. Pectic substances, though perhaps not in the form of pectic acid, are also secreted at later Stages in cell wall development, usually in combination with other materials, as cellulose, callose, and nitrogenous substances. In most of the thickened walls studied, evidence of the presence of pectic substances was found in the secondary thickening layers. The general rule, that pectic compounds are deposited in the cell wall early in the life of the cell, and that layers depos- ited later are predominantly non-pectic, is seen to have exceptions 30 BOTANICAL GAZETTE [JULY in the case of the innermost strata of the bast fibers of Nerium and rose, and in the lining of the canals of the rose and of the pits of the pine. In these cases we have the deposition first of a pectic layer, then of non-pectic or mixed layers, and then again of a pectic layer. The purplish tinge imparted to the bast thickening layers just mentioned by ruthenium red indicates that these layers consist of some pectic substance or substances, other than pectic acid, at least in the form in which it exists in the cambium walls and the middle lamellae of other tissues. Whether the period during which the cell generally deposits pectic acid, and thus forms the middle lamella, marks any spe- cial stage in its development which can be sharply distinguished from subsequent periods in its history is not certain. The evi- dence at present available seems to indicate that the pectic layer continues to increase in thickness about as long as the cambial cell is increasing in size. It is possible that the attain- ment by the cell of its adult size marks the limit of the growth of the middle lamella. In this case we might say that pectic acid is deposited so long as the metabolic processes of the cell result in a f/us which is expressed in cell growth, but that later, when a metabolic equilibrium has been established, or when the excess of food is stored instead of being used for growth, or when the protoplast degenerates, a predominance of other cell- wall materials is deposited. The evidence for such a view, however, is far from complete. It must be remembered, too, that in the cambium cells the distinction between middle lamella and inner non-pectic layers is very early present, and that the middle lamella continues to increase in thickness after it is sepa- rated from the protoplast by the non-pectic strata deposited later. The middle lamella is not to be considered as consisting merely of the wall layers early deposited by the protoplast, without undergoing any later modification. It is quite possible that the pectic acid which characterizes it may in part be sec- ondarily formed or deposited by infiltration through non-pectic layers which separate it from the protoplast. Some such hypothesis as this is necessary to account for its increase in Igor | THE MIDDLE LAMELLA 31 thickness during the later stages of cambial growth. Of inter- est upon this point is the fact noted by Mangin that, although in the meristem of some of the plant tissues studied by him no middle pectic layer could be detected by staining, yet, upon a treatment known in other cases to dissolve pectic acid, the cells are dissociated. My study of very young rose shoots, though not carried back far enough to dispute Mangin’s results, yet shows that, at a point where the tissues have become very slightly differentiated from the meristem, the cambium walls show strongly by their affinity for ruthenium red their pectic nature. CONCLUSIONS. The facts that have been cited seem to me to show conclu- sively that the middle lamella is not merely the partition wall first laid down, either as a single or a double layer, by the plasma membranes. Nor is it, on the other hand, merely an intercellular substance of cement, a means for-binding the cells together, as Mangin holds. Individual cells are not separated from one another, either in their formation or their later develop- ment, and no reason appears why an intercellular cement should be secreted. There is, however, abundant reason why, in woody tissues especially, there should be during cell growth a plastic region in the cell wall, which should be in a measure adaptable to the changing size and form of the protoplast itself, and to the firm, resistent layers whose form must correspond to that of the protoplast at the time of their deposition. The middle lamella is, therefore, a wall layer with a complicated history, undergoing after its first appearance changes in form, increase, and probably at times decrease in mass, and changes in chemical composition; its history, too, is not identical in different tissues. There can be no doubt that, in the higher plants studied, the young cam- bium walls are included in, and form the basis of, the middle lamella of the older tissues. These may at first include, at least in the pine, an amorphous Zwischensubstanz, which occupies a cleft formed by the splitting of the young radial walls. Quite early in the history of the cambium cells, a non-pectic layer 32 BOTANICAL GAZETTE [JULY appears within the first-formed pectic stratum; but the latter, now properly a middle lamella, continues to increase in thick- ness even after the appearance of the non-pectic layer. The substance of the middle lamella, now, is not rigid, but is more or less plastic, or even, as Dippel believes, soluble under certain conditions of development. Its solubility may lead at times to its partial absorption by the rest of the wall. More often, prob- ably, its mass is increased by further secretion of pectic material from the adjoining cells into the spaces formed by their round- ing up. The plastic nature of the whole layer allows the modi- fication of its shape, also, as the cells round up and as more resistent layers are deposited against it, leading to its being massed at the corners and pressed out to a thinner layer at the sides. The pressure may be in certain cases so great at the corners as to increase the density at those places; but in gen- eral, as the cells round up and draw away from the corners, the pressure there is relaxed, and, if new material is not deposited, the substance becomes less dense, perhaps swelling by the absorption of water. Or it may be that already the middle lamella has split, and the cleft has been filled by the exudation into it of a pectic fluid, which is later to be reabsorbed. At any rate, sooner or later the tension incident to the rounding-up process brings about a split in the middle lamella, and an inter- cellular space is formed. In case a Zwischensubstanz was pres- ent, a splitting at this time is unnecessary, and the intercellular space results from the absorption of this substance. If the sub- stance of the middle lamella is sufficiently rigid, the intercellular space remains angular; but if the material still retains some plasticity, or if it undergoes a chemical change upon its expo- sure at the newly formed surface, the angles may be rounded, or may disappear entirely, leaving a circular or elliptical space. The chemical’ changes effected in the middle lamella after the cell has attained its final form consist in the conversion of pectic acid into insoluble pectates, or even, as in the cork cells of Tilia, its replacement by entirely different substances. It seems to me plain that the term ‘ middle lamella,” Igor | THE MIDDLE LAMELLA 33 indicating merely the position of the layer in question, is prefer- able to either ‘‘cement”’ or “intercellular substance.’ The latter term, arising from a totally false conception as to the origin of cells, should certainly be dropped. There is also no evidence that the cell wall layers which, taken together, form the middle lamella have any significance as a cement for binding the cells together, and soit is inappropriate to apply to them a name sug- gesting any such function. The fact of the origin of the middle lamella as a double layer deposited on the surfaces of two plasma membranes, and the fact that in the formation of intercellular spaces it always splits in a median plane, must always be borne in mind. This constant plane of cleavage may be called the primary cleavage plane, or, perhaps, the primary cleft when it is spoken of with reference to the space between the daughter plasma membranes in which the cell wall material is deposited. I would distinguish cambial walls as the walls of celis yet capable of division, and primary cell walls as those layers added during the growth of the cambium cell into a full-sized wood or bast element. Secondary thickening would include the strata deposited in the subsequent history of such cells. This second- ary thickening may be subdivided further where evidence exists that its deposition has been interrupted and subsequently resumed; we should then have tertiary thickening, and so on. For the adult condition of thickened cell walls, the following terminology might be proposed as most exact from the stand- point of our present knowledge : The boundary plane between adjacent cells, commonly invisi- ble except where indicated by intercellular spaces, would be, following the suggestion already made, the primary cleavage plane. Including this on both sides, we should have the appa- rently homogeneous middle lamella, including the cambial walls and more or less of the primary cell walls, according to the pro- portional pectic acid content of the latter. Then would come those layers of the primary wall, if there be such, which are pre- dominantly non-pectic; and lastly the layers of secondary and 34 BOTANICAL GAZETTE [JULY perhaps tertiary thickening formed in the metamorphosis of the adult cell into its permanent condition as the element of a tissue. The investigations here discussed were undertaken at the suggestion of Professor R. A. Harper, and have been carried on under his direction. All the results accomplished are due in the highest degree to his suggestions and assistance. UNIVERSITY OF WISCONSIN. LITERATURE CITED. ~ . Bessey, C. E.: Botany, seventh edition. New York, 1892 2. DIpPEL, L.: Das Mikroskop und seine Anwendung. Zweite umgearb, Braunschweig, 1898. 3- MANGIN, L.: Sur la constitution de la membrane des végétaux. Compt, Rend. 1072144. 1888. 4. : Sur la présence des composés pectiques dans les végétaux. Compt, Rend. 109:579, 1889. 5: : Sur la substance intercellulaire. Compt. Rend. 110: 295, 1890. 6. : Sur les réactifs colorants des substances fondamentales de la SEEN Compt. Rend. 111: 120. 1890 7. : Observations sur la membrane exitiviadues Compt. Rend. 113: ee 1891 = Sur eee du rouge de ruthénium en anatomie végétale. mpt. Rend. 116: 653. 1893. 9. Pepe E.: Ueber Zellbildung und Zelltheilung. Jena, 1875. : Das Botanische Practicum. Jena, 1884 11. -—_——: Das Botanische Practicum. Dritte umgearb, Aufl. Jena, 1896. : Die pflanzlichen Zellhaute. Jahrb. f. wiss. Bot. 31:511. 18098. 13. Tieiienalen: H. G.; The development and ee of the cell plate in higher plants. Bot. Gaz. 30: 73, 154. 14. TREUB, M.: Quelques recherches sur le oe de noyau dans la division des cellules végétales. Amsterdam, 1878. 15. VINES, S. H.: Student’s Text-book of Botany. London, 1896. 16. ZIMMERMANN, A.: Botanical } Microtechnique. ‘Amer. ed. 1893. STRUCTURAL STUDIES ON SOUTHWESTERN CACTACEAE. CARLETON E. PRESTON. (WITH NINE FIGURES) Tuat the Cactaceae, a family in every respect appearing to be of recent origin, are unique in the extent of their adaptive modifications to arid conditions is a commonplace. Not only does this adaptation affect the external parts as gross structures, but in still greater measure does it work changes upon the minute anatomy by modifying the histological elements. These internal changes, keeping pace with and supplementing the external, raise this family to the first rank among the desert plants. A critical study of the minute anatomy from the stand- point of ecology, therefore, should be most fruitful. Moreover, since the family is one of late origin, in which definite groups have so far not crystallized out, but between which nearly all connecting links are present, a general comparative examination of internal structure should afford at least some further clue as to the lines upon which the various groups have developed or are still developing, adding to or confirming the evidence already obtained from the study of external characters. Viewed from the standpoint of the systematist, this paper, being a study of the general histology of eight southwestern forms, can by itself be of little value, serving only as an addition to the number of species already critically examined, Although in collecting the material for study it was my intention to choose such forms as could be considered typical, I found, as work went on, that too much variation occurs to admit of their use in this way except upon the broadest lines. In order to do complete and Satisfactory systematic work on the basis of structural char- acters, evidently one would have to compare, not only all the genera within the family, but at least the larger subdivisions and c*90%| 35 36 BOTANICAL GAZETTE [JULY the doubtful species in each genus. The single genus Cereus contains a multiplicity of forms, the structures of which must be very diverse. How are these related, and what are their several connections with the other genera? Does the gradual change in internal structure agree with the external and strengthen the validity of the various systematic groups as now recognized ? Schumann? has pointed out that the original ancestor of the Cac- taceae was probably something like Peireskia. Ganong? has followed up and confirmed the relation between this genus and Opuntia, chiefly on the basis of external characters. What are the steps by which the corresponding internal structure has changed? Schumann’ has hinted rather strongly at the con- necting links between Opuntia and Cereus; anatomical studies may give more definite information upon this point. This paper seeks to impress the importance of careful struc- tural study to help out the ordinary work on external characters, to point out the general trend of evolution exhibited, and to interpret certain anatomical conditions upon a physiological basis. As regards environmental adaptation, next to diminution of surface and other modifications to lessen transpiration, the stor- age of water is most important. The turgor exhibited in many of the more succulent forms is surprising. Especially is this the case in the younger portions of the arborescent species, and throughout the interior of those which are low and cespitose, where for a long time turgor alone is sufficient to sustain all the weight placed upon it—-in other words, to preserve the rigidity of the plant. Hence there is no necessity for a strong vascular system ; in fact, its room is more valuable than its presence, and we find in general that the vascular system is of late develop- ment, and in low forms almost never reaches a high state of perfection. The component elements also are oftenest of a kind generally considered primitive, showing an apparent retro- gression, but one which in many cases is complicated by the 1 Gesammtbeschreibung der Kakteen. 1897. ?Beitrage zur Kenntniss der Morphologie und Biologie der Kakteen. Fiora 79: 49-86. 1894. 901] SOUTHWESTERN CACTACEAE 37 addition of adaptive modifications. This is true particularly in the first form examined, Cereus Fendleri, of cespitose habit, and one of the most succulent. Upon cutting a shoot transversely one will notice that the bundles recede into the tissue for some distance, both in the severed portion and in that remaining on the plant, while the parenchymatous tissue still remains turgid. It is thus easily demonstrable that the bundles are normally under great tension. The mechanical arrangement by which such elas- ticity is rendered possible will be found in the structure of the xylem. Another point of interest, probably of value upon sys- tematic grounds, is the behavior of the bundles in the matter of secondary medullary ray formation, development of inter-, fascicular cambium, and anastomosis. Before considering the various forms separately, there are certain points of structure which can best be mentioned in gen- eral. Schumann, both in his monograph and in Engler and Prantl’s Pflanzenfamilien, gives an idea of the ordinary structure of the group. A condensed account is also given by Ganong? which I take the liberty to quote asa basis for the following discussion, As to the tissues, it is enough here to say that the characteristic xero- philous appearances are a strong cuticle, thick epidermis, perfect cork, sunken stomata, collenchymatous hypoderma, deep palisade layers, great development of pith and cortex, which consist of large, round, splendidly- pitted water-storing cells, often containing mucilage; a fibrovascular system, in general simple in its make-up, lacking annual rings, composed as to its xylem of strongly-ringed and spiraled tracheids, which are often collected into gland-like masses, the whole system conforming closely to the external form and following its morphological changes. Regarding the character of sunken stomata and their relation to hypoderma slight comment is necessary. The stomata are not sunken in the sense of having deep-lying guard cells; on the contrary, they are well upon the surface, as Schumann notes. Whatever vestibule there may be is of a different nature. With the increase in thickness of hypoderma an interval is produced 3Recent problems in the anatomy, morphology, and biology of the Cactaceae. Bor. Gaz. 20: 130. 1895 38 BOTANICAL GAZETTE [juLY between the assimilative tissue and the guard cells. Through this collenchymatous tissue run canals connecting the two; these canals must at all times be filled with water vapor, and probably act in a much more effective manner than would an equally large external vestibule. The hypoderma is of varying thickness, of one cell layer only in some species of Mamillaria, of seven to ten in certain other forms, as in Ceréus giganteus. There appears to be no difference in the character of the cells in the various groups; the walls are irregularly thickened, the thin portions allowing osmotic transfer of materials between the cells, and from them to the epidermis. Portions of hypoderma are shown in figs. I—}. With respect to assimilative tissue, Ganong speaks of deep palisade layers. Schumann, on the other hand, says that the cells are not in palisade form, but spherical. In point of fact, there seems to be considerable variation in this regard. In no form which I have examined have I found an assimilative tissue of absolutely spherical unmodified cells. There has always been an elongation perpendicular to the surface, sometimes slight, as in Opuntia leptocaulis, in other cases very well-marked, as in Echinocactus Wishzeni. There is also a general compacting of the tissue in this region. These elongated cells extend inward in the form of more or less definite chains or filaments, grading rather slowly into the ordinary parenchyma tissue, which in many cases contains chlorophyll in some quantity, in others is color- less. In the latter instance the chlorophyll, as noted with the naked eye, seems confined to a well-marked rind. It is perhaps only fair to suggest that environment may have a considerable influence upon the development of the palisade form in this family, as well as in those which possess functional leaves. The classical works of Stahl, Pick, and other investi- gators leave little question but that the elongation of the assimi- lative tissue, at least in some degree, is in direct response to the stimulus of light. The habitat of the plants here described and contrasted is one marked for its great light intensity, which, as I have found from a rather comprehensive examination of foliar Igor] structures, leaves its imprint upon the arrangement and de- velopment of assimilative tis- sue in almost all cases. Schu- mann’s statement that the assimilative cells are spherical may be true of the plants when a general average is taken, in- cluding epiphytic types, and others growing under entirely different light conditions. Pos- sibly also the effects of arti- ficial cultivation may have some place in his results. The medulla in many cases is of thick-walled and pitted cells, as Ganong describes, but this is by no means universal, often varying within the limits of a genus as now established. The cells are often exceedingly thin-walled and have no need of pits to maintain connection between them. The presence in this tissue of mucilage, crys- tals, etc., is likewise variable with the species, and needs separate discussion. Not only do the crystals vary in distribu- tion to a wide extent, but also in relative frequency, some Species being far richer in this deposit than others of the same habitat, a fact which, it seems SOUTHWESTERN CACTACEAE Fic. 1.—Epidermis and hypoderma of Opuntia arbuscula; opening of stoma does not show, owing to slightly oblique charac- ter of section. Fic. 2.—Epidermis and hypoderma of Cereus Fendleri in section, showing stoma. Fic. 3.—Hypoderma of Cereus Fendleri, seen from inside, showing character of walls and opening of stoma canal (s). to me, can only be referred back to the varied selective power of roots for chemical solutions. 40 BOTANICAL GAZETTE [JULY The particular value of Ginong’s terse and concise descrip- tion of the fibrovascular system lies in the last clause, ‘‘the whole system conforming closely to the external form, and fol- lowing its morphological changes.” Upon this fact depends the usefulness of internal structure as evidence of systematic value, a point which this writer thus recognizes, and one which I feel that I cannot too strongly insist upon. Schumann gives a slightly fuller account of the vascular system, noting the pres- ence of annular and spiraled ducts, especially in young bundles, and their subsequent replacement by the various tracheids. This is also the statement of the fact that a regression toward more prim- itive conditions is taking place, by the substitution of elements of a less highly developed character. In this particular espe- cially considerable variation is shown, offering, together with the methods of bundle-branching and anastomosis, a very fruitful field for research from a systematic point of view. Annual or periodic rings are not always absent, as will be shown; in fact, in one form I have examined they are peculiarly marked. The work presented in this paper was done on fresh material collected in the vicinity of Tucson, Ariz., during December 1900, and examined immediately at that place. Opportunity was thus given to study the plants in their natural habitat, a very valuable advantage in dealing with physiological phenomena. CrREUS FENDLERI Engelm.—The hypoderma is compara- tively thin, of about three layers of cells; the cuticle is thick as usual; there are no crystals in the epidermis, only an occasional one occurring in the deeper cells of the hypoderma. The assim- ilative cells are large and fairly elongated, those nearest the sur- face containing the most chlorophyll, those toward the center grading into a very thin-walled parenchyma also slightly chloro- phyllous, and filled with an intensely stringy and slimy mucilage, which, by taking stains readily and by preventing dehydration, renders any delicate manipulation of sections upon fresh tissues out of the question. It is by this means that the plant is able to hold water so persistently. In the parenchyma are imbedded the bundles, slightly separated from each other in most cases, 1901] SOUTHWESTERN CACTACEAE 41 but with some anastomosing. The lateral branches, which apparently start for the tubercles, break up and form a network through the tissue. In this species the bundles appear to start ina definite circle in cross section, in contrast to the undulate line in which they occur in certain other forms. There seems to be no interfascicular cambium, the bundles in their exceedingly — pelea 1 ee sess slow growth apparently push- Re q ) ing their way outward into the “succulent tissue, compressing it about them, and taking up the room it formerly occupied, in- stead of the growth of the medullary tissue keeping pace with the increase of the bundles. The development of phloem is exceedingly slight, consisting wholly of sieve tubes and com- panion cells, with no trace of thick-walled elements. Of the xylem more needs to be said. A bundle is repre- sented in fig. 4. The elements which compose it are but two, ie fees a sects “dt sie ¢ spiraled ducts and tracheids pundle in Cereus Fendleri. with a peculiar flattened spiral, the latter being by far the more abundant. These alternate in time of development, showing what appear to be periodic changes. The very earliest portions of the xylem are composed of the ordinary spiraled ducts, irregular in size and position, and there- after a small layer of these cells appears at what is probably the beginning of each season’s growth. The flat spiraled tracheids, an element already mentioned by Ganong and Schumann, form in definite rows, each apparently the product of a single cambium cell, which, after cutting off the few irregular segments to form the spiraled ducts, apparently undergoes no more division by radial re 5 OI = iG Py LY Ss LOTS] OO)p (% SY Serie SAIN 6 COOLERS TOKO a oy) "> Ca) le Oe foloIG ee ASO 42 BOTANICAL GAZETTE [JULY walls during the season. All'the broadening of the bundles, there- fore, occurs at the beginning of a period of growth. Likewise, when a branching is to occur, the resulting medullary ray appears at the beginning of the season, as shown in the drawing. It seems likely that the flattening of the spiral in the tra- cheids, which compose most of the xylem, is what gives the elasticity so prominent in this bundle. Such a flattening inwards would allow a far greater stretching of the spiral than would be possible in the commoner round type. A fact which strengthens this assumption is that the walls of the tracheids in a preparation will be ; seen to be wrinkled between the spirals (jig. 5). ee aie living condition those were in all proba- tracheid of flattened spiral type from bility perfectly straight, and the turns of the xylem of stem bundle spiral at a greater distance apart. ee Another point which can be deduced safely from an examination of the figure is with respect to periodicity of growth. From the comparison with the amounts of earlier years, it will be seen that the xylem for the season was about half formed at the time of collection of material. As this took place in the latter part of December, it seems probable that the season of growth in the southwest is limited by the sum- mer drouth and intense heat, rather than by the winter cold. The branching of bundles and formation of secondary medullary rays in the stem is not of common occurrence in the Cactaceae, so far as my investigation goes. In most of the forms to be described a different state of affairs will be noted. The bundles of the root, however, branch in the ordinary manner in all the species I have examined. The absorptive root* of Cereus Fendleri has a cork four to six cell layers in thickness. Successive layers of this kind are formed, with parenchymatous intervals, so that the bark is being constantly shed in flaky fragments, and is with equal rapidity replaced. The cork cambium lies a short distance outside the 4 Bor. GAZ. 30: 348. 1g00. Igor] SOUTHWESTERN CACTACEAE 43 secondary phloem of the bundles. This phloem is composed almost wholly of sieve tubes which, when examined January 1, had no callus visible. Lateral plates occur in abundance, in addition to those at the ends of the cells. The xylem has but two elements, at least*in the secondary portions. The more prominent, and by far the more abundant of these, is the tracheid, with ellipsoidal pits in more or less regular lines, an element which, from the shape of the pits, hardly coincides with the ordinary scalariform vessel, as seen in Pteris, for instance, but which, for convenience, I shall here designate by that term. Though the separate cells are compar- atively short, there is often a continuous passage through several (a transition stage toward the true trachea), the walls between the adjacent cells being partially dissolved, and represented by nothing more than athick and distinct ring. The other element is the obliquely pitted fiber tracheid, which apparently supplants the ordinary wood fiber in the majority of the Cactaceae.s The medullary tissue is but little compressed in the rays, giving a rather loose structure to the root as a whole. Crystals here, as also throughout the entire plant, are very few in comparison to their number in all the other species examined, occurring in the various parts of the parenchyma tissue, always in the aggregate form. The anchoring root differs from the absorptive chiefly in the relative numbers of fibrous and vascular elements, as stated ina previous paper,® in which the phrase ‘‘ wood cells’’ was used by mistake for “ fiber tracheids.”’ There is also a slight difference in the amount of sieve tissue developed in the phloem, the greater amount belonging naturally to the absorptive root. In Species where the two systems are not clearly differentiated, the elements are more nearly equal as regards amount of space Occupied by each. As a rule, the only difference between the two roots is one of proportion of elements, not of their variety. 5Fora drawing of this element see Strasburger’s 7 vxt-book of Botany, translated by Porter, fg. 143 ft. ° Bor. Gaz. 30: 348. 1900. 44 BOTANICAL. GAZETTE [JULY The absorptive root, therefore, will be the only one taken under consideration in discussing the remaining forms, except in one or two special cases. The difference between the two is hardly as well marked in Cereus Fendleri as in Opuntia fulgida, to be mentioned later. Ecurnocacrus WisiizEnt Engelm.—The epidermis and hypo- derma are without crystals, the latter of six or seven layers of cells. The assimilative tissue is compact, with cells much elongated perpendicular to the surface, even in the sinuses between the ridges. The young parts do not develop chloro- phyll in the sinuses until the ridges of tubercles, at first close- pressed together; become separated. The chlorophyll in this form is confined to a definite rind of palisade-like cells, which on the inside changes somewhat abruptly to a colorless paren- chyma, rather thick-walled and fitted with a splendid system of pits and openings. Within this tissue are occasional crystals. Mucilage does not seem to be abundant in this species, but its place is taken by a watery solution, especially prominent in parts still growing, which upon long exposure to the air causes the tissue to assume a pinkish tinge. The medullary tissue about the bundles contains a fairly large quantity of starch, greatest in amount and widest in distribution in the young portions. The bundles, as seen in cross section in a young stage, are not in a perfect circle, but rather in a zigzag line (fig. 6, dia- gram), with alternate points and depressions. There seems to be no branching or secondary medullary ray formation, and but little increase in the width of the original bundles. The zigzag line does not appear to be entirely the result of bundles passing out of the tubercles, for nearly all of the bundles in section appear to be cut directly across when all are cut in the same plane. They anastomose to a considerable extent, and, taking as centers the bundle at the base of each inner depression, join together with it to form a complex wood structure. The direc- tion of growth of the side bundles is slightly inclined toward the central one, bringing the xylem of the whole group into Igor] SOUTHWESTERN CACTACEAE 45 close contact at an early stage. The phloem portions, however, remain distinct for each bundle. By this slight divergence, considerable medullary connection is left between the pith and the thick layer of storage tissue outside the poorly developed bundle cylinder, a far greater connection than could occur with the usual distribution, thickening, and branching of bundles seen in dicotyledonous stems. This peculiar grouping will be noted fh ' am ri eri U 1 ‘ ‘ ‘ > ae ai ae ee oe ANE aa Sere ie ~ ~ \ \ a9 eC ; ‘ ! Ce / ! J J ' 1 i] i] ‘ ' ' 4 ; , 1 ) , ' ! ! a ~ wer eee ee eee : y Fic. 6.—Diagram of position of bundles in Echinocactus Wislizeni and others. in a far greater degree of perfection in Cereus giganteus, next to be described. Here, as there, there seems to be no anastomosis of these groups of bundles, the incipient wooden ribs. In Opuntia, on the contrary, where this anastomosis of bundles likewise occurs to some degree, a still further reticulation is found, in that the groups as units anastomose. As regards more minute structure in £. Wisiizent, the phloem is composed mainly of sieve vessels, the outer of which, together with the adjacent parenchyma, are composed and rendered sclerenchymatous, forming a sheath, here separate for each indi- vidual bundle. The xylem, at least in later portions, is com- posed entirely of spiraled tracheids of uniform size, but not in definite rows, the spiral also being of the ordinary type. A 46 BOTANICAL GAZETTE [JULY certain amount of elasticity of bundle occurs here, but nothing like that in C. Fendlert. The enormous amount of thick-walled parenchymatous storage tissues seems to have more effect in giving firmness to the internal structure than does turgor. The bundles have almost no supporting function, and are never well developed, appearing, in the widest part of a fair sized specimen, only 1 in thickness of xylem. The branches passing out to the tubercles or ridges divide through the tissues as in C. Fendlert. The absorptive root contains numerous small crystals in the rays, more in proportion than appear in the stem. The chief xylem element is the scalariform tracheid ; fiber tracheids also occur. In the anchoring root, as before, the proportions are reversed. The phloem is mainly composed of sieve tubes, but has a slight sclerenchymatous compressed sheath, a character which will be noted as fairly constant in both stem and root. through the majority of forms. CEREUS GIGANTEUS Engelm.—The epidermis and hypoderma are both without crystals, the latter being exceedingly thick, of twelve to fourteen layers of cells. The assimilative tissue is of the usual elongated cells, the filamentous arrangement of which is extremely well marked, to such a degree even that the fila- ments in a preparation can be separated easily. Chlorophyll extends deep into the tissue, and the elongated character of cell is found for nearly 20™™. The medullary tissue is thick-walled and pitted, containing starch in the immediate vicinity of the bundles. Mucilage, though present in quantity sufficient to hinder the staining of sections, is not very thick. There is, however, especially in the younger parts, a watery solution apparently identical with that seen in £. Wislizeni, but much stronger, causing the tissue upon exposure to turn almost immediately from white to pinkish, and then to a dark purple. In bundle formation, the method described for EZ. Wislizent above is carried to greater perfection. Here not only do the bundles join their xylem portions, but all those of a single rib are surrounded by one sheath, formed gradually. In a young stage it may be seen outside each separate bundle, later two or IgOI | SOUTHWESTERN CACTACEAE 47 more bundles are embraced by a single sheath, and this process apparently continues until all the bundles of the rib are thus bound together. This sheath is not the ordinary one of com- pressed sclerenchyma so commonly found next the sieve tubes. This outer one seems to be a special structure, composed of the obliquely pitted fiber tracheid so often mentioned, which here inclines more than ordinarily to the tracheid form, with wide lumen and with the extremities of the cells scarcely pointed. The individual cells are much larger than those of the same nature found in the xylem. The ribs alternate with the ridges of tubercles, botii undergoing more or less branching. A full- grown rib, according to measurements taken from dead skeletons, seldom exceeds 3™ in thickness. The phloem has the usual development of sieve tubes; the xylem is made up almost wholly of scalariform tracheids, an element so far noted only in the root. Besides these, there also occur fiber tracheids, undoubtedly in increasing proportion as the size of the rib and the corresponding function as a support increases. The absorptive root differs from the others described in having in the phloem an abundance of fiber tracheids, forming a wedge pointing outward, starting just outside the sieve tubes of each bundle. These cells are also numerous in the xylem, together with the scalariform tracheids. Crystals are common in the cortex as well as in that of the stem, and are large and closely aggregate. They do not seem to be entirely homoge- neous, but take some foreign substance asa nucleus. At times they appear to grow by concretion, showing when viewed in optical section a series of concentric circles. The necessity for a fibrous strengthening in the absorptive root of this large form can best be understood by examining the method of growth. In starting as a seedling, the plant begins life with a base slightly sunken in the ground, but far narrower than the heavy column which it is to support. Therefore, as this column develops, the whole becomes very top-heavy, and would certainly fall were slightly greater surface exposed to the wind. The anchoring root, at this time well developed, holds 48 BOTANICAL GAZETTE [JULY it firmly to the ground, but only to a slight degree prevents a swaying movement, easily induced, even in plants 1.5™ high, by comparatively little force. Hence to the large lateral roots, developed from the sunken base at a later period, is given the function of steadying the whole plant, in addition to their pri- mary work of absorption. MAMILLARIA GRAHAMI Engelm.—The epidermis is free from crystals, but the one-layered hypoderma contains them in almost all its cells. The assimilative tissue is of small cells elongated, as usual, perpendicular to the surface. The colorless medullary portion is of thick-walled cells pitted as in &. Wishzeni and C. giganteus, and grades into the assimilative tissue. The lateral bundles can here be traced out into the tubercles. The stem bundles appear to start in a more or less wavy line, and to suffer fusion to some extent. There is comparatively little branching, but secondary medullary rays are occasionally found, and there is also a growth of the bundles in width, so that finally a rather compact cylinder is formed. As regards branching and internal structure of bundle, this species approaches C. Fendleri nearer than any of the others. The xylem is composed of alterna- ting portions of spiraled ducts and tracheids of the flattened- spiral type, the latter being the more prominent. The phloem has a slight compressed sheath of sclerenchyma. Sieve tubes appear rather poorly developed, a_ thick-walled, somewhat elongated parenchyma taking up most of the region of the phloem. In this small species there is apparently no distinction of anchoring and absorptive root systems. The bundle cylinder is compact, the pith practically absent, and the medullary rays reduced generally to a single cell layer. The phloem is slight and of sieve tubes. Outside the phloem, filling the small space between it and the cork cambium, is a parenchyma tissue of large rather thick-walled cells. The xylem consists chiefly of spiraled tracheids, with some amount of extra reticulation, and of smaller ringed ducts, the rings flattened inward, sometimes also bound together by extra spirals or reticulations. Crystals occur but 1901] SOUTHWESTERN CACTACEAE 49 seldom in the root, but are present in some quantity in the cortex and medulla of the stem. OPUNTIA PHAEACANTHA Engelm.—The hypoderma is rather thick, of six to eight layers, the outer one compactly filled with crystals, which occur also in great numbers throughout the assimilative and medullary tissues, clustering more or less about the bundles, but especially prominent in and about large duct- like spaces just exterior to certain of the bundles in both stem and root (Schumann’s Arystalschldéuche). Mucilage is fairly abun- dant. The assimilative cells are somewhat elongated, perhaps less so than in other forms so far recorded. Chlorophyll is deepest in the outer layers, but present as well in almost all parts of the deeper thin-walled parenchyma. The stem bundles form a loose reticulated network, branching early, especially on the flat surfaces of the stem, forming very wide secondary medul- lary rays. The phloem has the usual development with a scle- renchyma sheath. The xylem in early parts consists almost entirely of smaller annular ducts with flattened rings, but later forms spiraled tracheids and a large amount of obliquely pitted fiber tracheids. In the root especially are seen the Krystalschlauche in a high state of development, with a diameter of almost 1™™. The crystals seem to be deposited mainly along the edges of the duct, which may serve also to some extent for the transfer of mucilage. The phloem of the bundle has a slight sheath; the xylem is composed of large and small scalariform tracheids and the usual fiber tracheids. OpunTIA FULGIDA Engelm.— The epidermis contains a deposit of crystals, a small quantity of which also occurs in the relatively thin hypoderma. The assimilative tissue is of cylindrical cells in lines, grading in the interior into large thin -walled paren- chyma, also chlorophyllous. Mucilage is exceedingly abundant, and the ducts for its accommodation and transfer are both large and numerous. Large crystals, loosely aggregated, occur in small numbers throughout the tissue. The bundles here remain very small through several joints, Mo.Bot. Garden, 1902. 5° BOTANICAL GAZETTE [JULY later becoming fused into a reticulated skeleton, the separate bundles of a single strand also anastomosing. The wavy line formation is seen to a slight extent in these Cylindropuntias also; there is apparently no branching, and little increase in width of bundle. The phloem presents no unusual characters ; the xylem is composed of annular ducts, with flat rings at regular intervals; later on there occur also spiral tracheids, with steps grading to the true trachea. In an earlier article the roots of this species have been referred to as anchoring and absorp- tive. Portions of the xylem of each are figured here to show the difference in texture (figs. 7,8). In the absorptive roots the elements are wider, more regular, and less thickened. The phloem differs but slightly in the extent of development of sieve tubes; the xylem dif- ers as usual in proportions of elements, con- sisting in both of scalariform tracheids and fiber tracheids. From the prominence of the latter, both here and elsewhere, it would appear that the main function, if not the only one, which this element performs in the Cactaceae tionof portionof is one of support. xylem from absorp- OPUNTIA ARBUSCULA Engelm.—The hypo- — of Opuntia derma is fairly thick, with a compact deposit OnE atte ier of crystals in the outermost layer. The assimi- tionof portion of lative tissue is compact, the cells being small xylem from anchoring and greatly elongated inregions near the sur- pa of Opuntia ful- face. The chloréphyll/extends through the : thin-walled parenchyma as well. Crystals are very abundant throughout, varying considerably in size and com- pactness of aggregation. The stem is very turgid and mucilagi- nous, kept rigid by turgor for some time, with late bundle development. The-bundles finally form the same kind of reticu- lated skeleton seen in O. fulgida. The phloem is as usual; the 1901 | SOUTHWESTERN CACTACEAE BL xylem shows an occasional ringed duct, apparently an early ele- ment, but is principally composed of scalariform and fiber tracheids. Starch is present in considerable quantity through the stem. The cortical parenchyma of the absorptive root is thickened in many places for storage pur- poses. The cells are filled with a thick mucilage, which aids in holding the water. A consider- able deposit of starch, as well ip : us — AIF AL I as of the regular calcium oxa C Oger late crystals, occurs in the me- ss LOAF S F ah “A SBLBO, S28 dullary rays, but not at all in (ae eeope the cortex, an apparent differ- entiation of regions for vari- ous reserve substances. The phloem is of ordinary charac- ter, with a slightly developed Fic. sheath ; the xylem is composed °! Omntia arbuscula. mainly of scalariform tracheids of various sizes, but contains a few fiber tracheids as well. A rather peculiar phenomenon is presented by the convergence of the cortical tissue toward the center of the phloem. Whether this was to hasten transfer of materials between cortex and phloem in young stages, before the sheath was thickened, in a way analogous to the clustering of palisade about the leaf bundles, which so commonly occurs, is a question which at once suggests itself, but upon which I am able to throw no light at present (fig. 9). OpunTIA LEPTOcAULIs DC.—The epidermis contains crystals in cells specially enlarged to receive them. The hypoderma is of four or five layers, and the assimilative tissue is elongated very slightly. The parenchyma of pith and cortex is nearly colorless, thin-walled, and contains considerable starch, evidently of the translocation variety. This species is not so succulent as the majority, and has an earlier development of the bundles. A point in which it differs from all the rest examined is in its Boe ) DBRS (0) os LU @ 0 San ess eeeyo we y { Sd Yoyto S\0) io me g yf 3 KD) XO] EC ) IOC ae Le By iy ~ a G. 9.—Portion of b 52 BOTANICAL GAZETTE [JULY possession of a well-marked interfascicular cambium in the stem, allowing the medullary tissue to increase in proportion to the growth of the bundle, instead of having the bundle compress and supersede it, as is apparently the condition in the others. That even here this cambium is not so active as that of the bundle seems to be demonstrated by the fact that outside the phloem the scleren- chyma sheath still appears compressed, though to a less degree. The xylem is composed mainly of fiber tracheids, intermingled with which are scattering spiraled ducts. The ordinary Opuntia reticulation occurs together with the finer anastomosis of the bundles of a single strand. Between these closely connected unbranched bundles the medullary rays develop very thick walls, apparently lignified, and are so filled with a deposit of starch that in section they appear much more opaque than does the rather close-grained wood on each side of them. Mucilage occurs, but not so abundantly as in the majority of cases, and crystals of rather small size are scattered through all the paren- chyma tissue. The root does not materially differ from those already described. The cortical parenchyma is slightly thickened, and the xylem is composed of the scalariform and fibrous types of tracheids. SUMMARY AND CONCLUSION.—Several inferences may be drawn from the foregoing descriptions. In the first place, it will be seen that the roots, as regards branching and elements concerned, apparently undergo but slight variation, being com- posed, in almost all cases,of fiber tracheids and scalariform tracheids in the xylem, and of sieve tubes only in the phloem. To this general rule, as regards xylem portions, Mamillaria Gra- hami is the only exception among the forms here compared, and even in this the tracheid appears in greater prominence than the true duct. This exception, however, is enough to suggest that a more comprehensive examination will reveal more of a similar nature, and also perhaps disclose some links in the evolu- tionary chain between this and and the ordinary type. In the stem, on the other hand, there is a great range of 1901 | SOUTHWESTERN CACTACEAE es: structural deviations, which seem to take place along definite lines and by definite steps. This variation extends to bundle branching and further reticulation, to the extent of succulence, character of parenchyma of pith and cortex, development of mucilage, and even to the kinds of elements composing the xylem of the bundle. In the phloem alone there is little change. The practically universal substitution of fiber tracheids for wood fibers, and the equally common appearance of tracheid elements in place of true ducts, except in early growth, points certainly to a general regression. The fiber tracheid, moreover, is variable in itself. In most cases it is quite sharply pointed and thick-walled, with a lumen, though broader than that of an ordinary wood fiber, still rather narrow. In a few instances, however, this same element loses its pointed character, widens its lumen still more, and except for its thick walls and oblique pits, scattered and exceedingly small, closely resembles an ordinary tracheid. This is very well shown in Cereus giganteus, a rather anomalous example, however, since here the fiber tracheid supplants a bast fiber instead of a wood fiber, and does not undergo an equal enlargement in the xylem portions of the bundle. Crystal distribution and various storage peculiarities should be looked upon more as specific differences, I believe; likewise the thickness of hypoderma. A word or two may be said of Opuntia leptocaulis, The early development of the bundles, the smaller degree of succulence, the persistent character of the leaves, at least in some varieties, as noted by Coulter,’ the very slight elongation of the assimi- lative tissue, and above all the persistence of the interfascicular cambium, all tend to place this form in structure much nearer the ordinary dicotyledonous stem than any of the others exam- ined. It seems safe to conclude that this species comes lower in the evolutionary line of adaptive modification than the rest of the Opuntias here mentioned, and may be considered nearer the primitive form. 7Contrib. U. S. Nat. Herb. 3: 456. 54 BOTANICAL GAZETTE [JULY Perhaps the most striking results of any to be found here from the systematic point of view will be obtained by a compar- ison of the first three forms. Cereus Fendleri and Echinocactus Wislizent are entirely dissimilar as regards character of medullary tissue, mucilaginous contents, firmness of interior structure, and especially in respect to arrangement and branching of bundles. The third form, Cereus giganteus, corresponds in all the above mentioned peculiarities, not with the other Cereus, as would be expected, but with the Echinocactus. So far as I can tell from plates,® this closer resemblance to Echinocactus is carried out equally well in external characters, especially in those of the fruit, but to some degree at least in those of the spines and areolae (the last comparisons taken from living material). I have too little systematic knowledge of this group to contest the position of the species in classification, and can simply point to the fact that the two Cerei here mentioned belong at least to two different evolutionary branches within the genus, from one of which branches Echinocactus seems to have sprung. The genus Cereus seems to me an especially fruitful field for the study of internal anatomy on a comparative basis. It is rather evident, from a hasty glance at the great variety of forms, that many different lines of development begin in this vicinity, lines which, though at present somewhat unraveled from external character, could at least be well subjected to con- firmatory tests upon structural grounds. This would be of especial value in searching for the origin of the various other genera, the supposed offspring of this group. It must not be thought, however, that I wish to give too great importance in classification to the internal anatomy, pre- ferring it to the external characters. Its principal use is con- firmatory. Too many internal variations spring up independently in several forms to make them alone sufficiently reliable. There must be some gradual internal metamorphosis, however, to cor- respond with the external. In what deep-seated organs this is shown, and what parts are more easily influenced by environmental ® Engelmann’s Cact. Mex. Boundary. Igo1 | SOUTHWESTERN CACTACEAE 55 conditions, can best be determined by careful study. At present Iam inclined to believe that the most permanent and satisfactory evolutionary characters will be found in the fibrovascular sys- tem. When its metamorphosis is fully understood a far clearer insight can be gained, not only into the interrelationships of these peculiar and at present much confused groups of plants, but also into the steps by which they became gradually accus- tomed to their present rigorous habitat. HARVARD UNIVERSITY. Later investigation shows that the statement (p. 42, line 17 ff.) regarding periodicity of growth is incorrect. The smaller amount of growth noted was due to the drouth of the summer of Igoo. In spite of their storage facilities, these plants are apparently affected by temporary unfavorable conditions to a degree almost surprising. BRIEFER ARTICLES. NOTE ON BASILIMA AND SCHIZONOTUS OF RAFINESQUE. As Basilima of Rafinesque is still considered by many botanists to be the valid generic name for the genus based on Sfiraca sorbifolia Linn., it will probably not be uninteresting to point out that it has no claim to the priority attributed to it. Until 1891 the genus has been best known under the name Sorbaria, proposed first by Séringe (DC. Prodr. 2: 545. 1825) as a section of Spiraea, and given generic rank by A. Braun (Ascherson’s Flor. Brandenb. 177. 1864). In 1891, however, E. L. Greene (Flor. Francisc. 57) and O. Kuntze (Rev. Gen. Plant. 1: 215) almost simultaneously took up the Rafinesquean name Basilima for Sorbaria, and claimed priority for it on the ground that Rafinesque in his Mew Flora of North America (3:75. 1836) quotes “Raf. 1815” as authority for his Basilima, but neither of them seems to have seen or even known the original place of publication. If they had seen it, they would have been aware that Basilima of 1815 is a nomen nudum pure and simple, and therefore not entitled to priority. Rafinesque unfortunately makes no mention of the place of publication, but after some searching I was fortunate enough to find the name in his Analyse de la mature ou tableau de Luniverse et des corps organisés, a very rare book of 244 duodecimo pages, published in Palermo in 1815. It aims to give a short review of the whole range of natural history, and contains on page 173, under the order Rhodan- thia, the following passage reproduced here verbatim et literatim : 5. Famille Spireadia. Les Spiréades. Etamines héterines, perigone double, l'interne corollé, fruits capsules dehiscentes interieurement. I. Spirea L. 2. Rhodalix R.sp.do. 3. Drymopogon R. sp. do. 4. Basilima R. sp. do. 5. Filipendula T. 6. Gillenia Mench. From this enumeration it is impossible to gain any idea which species of Spiraea Rafinesque intended to include under his genera Rhodalix, Drymopogon, and Basilima, and he could therefore not claim priority for it, as he does, and quote as synonyms of his Basilima two names published before 1836. The older of these two names is 56 [JULY | | | ies a j . | Igor | BRIEFER ARTICLES 57 the Sorbaria of Séringe mentioned above, the second the Schizonotus of Lindley, published in 1830 in his Zu¢rod. Wat. Syst. Bot., p. 81, and (under no. 703) in Wallich’s Catalogue, which appeared about the same time. In both places, however, the name is mentioned only as a Synonym, and it seems doubtful whether this ought to be considered as a real publication of a generic name. The passages in both pub- lications read as follows: Wallich, Cat., p. 21, “no. 703. Spiraea Lindleyana Wall. Schizonotus Lindl. (gen. nov. Spir. sorbifoliam amplectens);” Lindl. Introd. Nat. Syst. Bot. ed. 1, p. 51, “ Rosaceae. . The fruit of Spiraea sorbifolia (Schizonotus m.) is capsular.” is ths second edition of his work, however, which appeared in 1836, Lindley enumerates (p. 145) Schizonotus as a genus, and characterizes it (p. 441) by mentioning Spiraea sorbifolia as the type. In the same year, Rafinesque published in Mew Flor. V. Am. (3:75-76) the genus Basiliina, with descriptions of two species, B. sorbifolia and B. Pygmaea. As it seems impossible to ascertain which of the two works was really published first, the genus Basilima ought to be preferred, since it had specific names attached to it, while Schizonotus had none and received none until 1840, when Lindley (Bot. Reg. 26: misc. 71) proposed the name Schizonotus tomentosus for Spiraea Lindleyana Wall. Those botanists who object to Sorbaria because it was published Only as a sectional name must therefore choose between Schizo- notus and Basilima. It would be very unfortunate, however, to revive the name Schizonotus, since it was applied afterwards and has been in use for two other genera, first to a genus with Spiraea discolor Pursh as the type by Rafinesque, who entirely misunderstood a note of Lindley in Bot. Reg. 16. 1830 in the text to A/. 7365, and says in Mew flor. N. Am. 3:75 under Schizonotus “I have adopted this genus and name on the suggestion of Lindley, who proposed to unite to it the next [Basilima], but the habit is too different.” But Lindley did not Say this. He only mentions there on the same page these two species of Spiraea, together with some others, and the name Schizonotus. The second application of the name Schizonotus was by A. Gray (Proc. Amer. Acad. 12:66. 1876) to a genus of Asclepiadaceae, which was subsequently changed by E. L. Greene to So/anoa; while on Spiraea @iscolor the name Holodiscus was bestowed first by C. Koch, who pro- posed it in his Dendrologie (1: 309. 1869) as a subgenus of Spiraea, which was afterwards given generic rank by Maximowicz in his excel- lent monograph of the Spiraeaceae (Acta Hort. Petr. 6: 253- 1879), 58 BOTANICAL GAZETTE [JULY without either of the authors being aware of the existence of a Schzzo- notus Raf. In /ndex Kewensis the Schizonotus of Lindley and of Rafinesque are not discriminated, but wrongly united under Schizonotus Lindl.; while Schizonotus Gray is enumerated as valid. Also the Zhecanisia discolor Raf. is there erroneously made a synonym of Spiraea discolor, but it is an herbaceous plant allied to U/maria rubra, and probably based on escaped plants of the European UW. pentapetala Gilib. (Spiraea Ulmaria Linn.) It will be clear that on account of priority as well as of usage the name Sorbaria ought to be considered as the valid name, with Schzzo- notus Lindl. (1830) and Baszlima Raf. (1836) as synonyms; while Holodiscus (Koch) Maxim. should be substituted for Schizonotus Rat., at least by those who follow the Rochester rules in accordance with article IV, and by others on account of expediency, to avoid all pos- sible confusion with the other two genera, of which one is liable to be revived under certain rules of nomenclature. Sorbaria of course will not enter into the American flora if Chamae- batiaria is considered as constituting a distinct genus, which it may be, though structural differences in flower and fruit are hardly pres- ent. In foliage and habit, however, it is strikingly different. The observation of Rafinesque that Sordaria sorbifolia is a native of North- west America seems not to be based upon any reliable source, at least it has not been found again on American soil, neither has S. grandi- flora Maxim. (Basilima pygmaea Raf.), which was suspected by him to occur probably somewhere in the same region.—ALFRED REHDER, Arnold Arboretum. POTAMOGETON POLYGONIFOLIUS IN NEWFOUNDLAND. THE only note of the occurrence of this species in North America I know of is contained in Linnaea 2: 216. 1827; where Chamisso has the following: ‘Hujus loci forsitan est: Potamogeton de St. Pierre Miquelon prés Terreneuve in Herb. Brogniart, sed major.” I have not seen the specimens in Brogniart’s herbarium, which is at Paris. In the Journal of Botany for June of this year, I record it from Sable island, about 100 miles off the coast of Nova Scotia. I have now to record it from Newfoundland, from whence I find specimens the Kew herbarium from the herbarium of Harvard University. Igor] BRIEFER ARTICLES 59 As*this was gathered by American botanists I record it in an Ameri- can journal, the more so as I wish to point out that there is some hope it may occur in Massachusetts in the neighborhood of where Cad/una vulgaris L. occurs. The label of the Newfoundland specimens runs thus: No. 231. P. heterophyllus Schreb. Muddy banks of brooks. Whit- bourne. 17.8. 1890. Coll. B. L. Robinson (and) H. Schrenk. Distributed from the Herbarium of Harvard University. The two specimens are not P. heterophyllus, however, but P. polygonifolius in its usual “heath” form. As this is the first time it appears in American books it may be well to give the synonymy and distribution. . P. POLYGONIFOLIUS Pourr. in Acad. Toul. 3:325. 1788. P. oblongus Viv. Anal. Bot. 2: 102. 1802; alsoin Fragm. Fl. It. 1:1. f7. 2. 808 P. plantago Batard, Fl. dep. Maine et Loire 64. 1809. ; P. parnassifolius Schrad. (ined. 1818) ex Mert. et Koch, Deut. Fl. 1: 839. 1823. P. uliginosus et affine Boenning ex Cham. in Linnaea 2: 216. 1827. P. paludosus Bory in sched. ex Cham. Z. c. P. natans intermedius Mert. et Koch, Deut. Fl. 2 ¢. P. microcarpus Reut. et Boiss. Diag. Pl. Nov. Hisp. 2:24. 1842 is a variety, or perhaps a subspecies. Dist. Europe: Iceland to Piedmont, and Holland to Russia. ASIA: Siberia altai (Led.) to Mongolia, and China to India and Japan. AFRICA: Morocco to Transvaal? (Wilms), and } Madagascar. AUSTRALIA: New Zealand. The species varies greatly, the var. y erécetorum Syme, Eng. Bot. ed. 3,9:28. 1869, representing the small heath form, often growing in mud, the var. B pseudo-fluitans Syme, /. c., the other extreme, 7. ¢., a fluitant form often in deep or running water with thin translucent submerged leaves and coriaceous upper leaves. It may be distinguished from wafans by the small fruit, the blunt Stipules, the base of the leaves never having the flexible joint of zatans (there are no “strings” in the leaf of polygonifolius) which enables that species to turn its leaves in any direction; from heterophyllus by the style being central, by the nervation of the leaves, etc. ‘I hope to send specimens of this species to various American bot- anists next fall, so that they may know it and keep it in view.— ARTHUR BENNETT, Croydon, England. . 60 BOTANICAL GAZETTE [JULY A NEW SPHAERALCEA. Sphaeralcea martii, n. sp.— Perennial, low and rather spreading, 15 to 30™ high: leaves and stems whitish-green with an excessively dense white stellate pubescence: leaves subhastate, with broad short basal lobes, crinkled, edges coarsely irregularly serrate, about 27"" long and 15™" wide; petioles about ro™ long: flowers large, 23" in diameter, in clusters of six or seven at the ends of the branches; pedi- cels 5 to 7™" long: calyx 11™™ to tip of lobes, 4™™ to base of lobes: petals fully 15" long and broad, rather pale but vivid scarlet (dull pink in dry material): anthers orange; pollen bright orange: styles crimson : fruit maliform, densely white-hairy ; carpels with very small beaks ; ovules two in a cell. . Picacho mountain, Mesilla valley, New Mexico, in volcanic soil, March 25, 1900 (Cockere//); hills north of Picacho mountain, April 7, 1900, in fruit (£. O. Wooton). A specimen is in Herb. U.S. National Museum. Differs from S. sudhastata in its pointed leaves, clustered flowers, deeply cleft calyx with long acuminate lobes, and especially in its early flowering, wherein it resembles the annual species. The only- flowering date I find reported for S. suéhastata is in August.—T. D. A. COcKERELL, Last Las Vegas, N. M. CURRENT LITERATURE. BOOK REVIEWS. Handbook of systematic botany. THE first volume has appeared of a handbook on systematic botany by Dr. Richard R. v. Wettstein,' of the University of Vienna. We do not often have texts that confine themselves so closely and accurately to the subject as does this very excellent outline of classification. Its aim is to give a general view of the plant kingdom with especial reference to the evolution of phylo- genetic lines. Prefacing the special parts, which deal specifically with the groups of plants, there is a general treatment of a number of interesting topics. There is first an historical account of the development of systematic botany, which one wishes were longer. This is followed by discussions of the conditions which lead to the development of lines (phyla) of ascent and the sort of evidence of value in the determination of the same. There is an excellent account of the methods of systematic botany and the data of most value in studies on classification. Admirable illustrations are presented of analogous and homologous organs, rudimentary structures and juvenile forms. Finally the author considers the origin of new forms on the supposi- tion of phylogenetic evolution, or in other words the principles of Darwinism. This is in the maina simple account of evolutionary factors, and is particularly d tance of the so-called ecological factors in the definition and adjustment of species, Following the short general part, which is only forty-four pages long, comes the special part that will deal entirely with the various groups. The present volume ends with the thallophytes, but we are promised the completion of the subject next year. The author divides the plant kingdom into seven branches, as follows: 1) Myxophyta, 2) Schizophyta, 3) sushi phyta, 4) Euthallophyta, 5) Phaeophyta, 6) Rhodophyta, and 7) Cormo- phyta. The success of a division into great groups depends saree Se upon the balance maintained, and this one has some peculiar disadvantages in its lack of symmetry. It will hardly be questioned that branches I, 2, 3, 5, and 6 are well differentiated phyla or closely related groups of phyla. But this simplicity is all out of proportion to the complex conditions presented in the * WETTSTEIN, RICHARD V.: Handbuch der sg agi Botanik. Vol. 1. 8vo. pp, vi+201. figs. 762. Leipzig: Franz Deuticke. Igor} 61 62 BOTANICAL GAZETTE | JULY Euthallophyta and Cormophyta. Passing to the detailed subdivision of the main groups, the descriptions of orders and families and the illustrations are generally very good. We note that the Protococcaceae and Hydrodictyaceae are placed at the bottom of the large order Siphoneae, a position that is cer- tainly open to question. The Charales are reduced to a suborder of the same group, which seems curious for so highly specialized and well defined a line of development. Among the fungi the Brefeldian system is followed in part, with much of the arrangement in Engler and Prantl; and the classifi- cation of the Phaeophyta and Rhodophyta is a brief outline of the latter work. These two groups are not given the attention they deserve.— B. M DavIs. MINOR NOTICES. HERMANN VON SCHRENK has published’ his address on ‘“‘ Factors which cause the decay of wood,” delivered before the Western Society of Engineers on February 6 last. It deals with such topics as structure, chemical nature, and decay of wood, fungi and structural timbers, and preventive measures.— Ae a on THE THIRD FASCICLE? of the list of the genera of seed plants, according to the system of Engler, has just appeared. The general character of the work was stated in this journal in the notice of the first part. In the present signature 1352 genera are listed, bringing the number up to 3842, the list beginning ie Lychnis (Caryophyllaceae) and ending with Geoffraea (Legu- minosae).—J. M. C. F. LAMSON-SCRIBNER* has published a revised edition of the second part of his American grasses, the first edition having been exhausted. The work has been entirely rewritten, the synonymy has been revised or extended, and the descriptions are much fuller. The two parts now contain illustra- tions and descriptions of 627 species, and are invaluable to those who would name grasses.—J. M. C CHARLES V. Piper and R. KENT BEATTIES have published a manual of the flora within a radius of about twenty-one miles around Pullman, Wash. This includes some twenty-four townships in Washington and eleven in * Reprint from Jour. Western Soc. Engineers, May Igol. ?DaLLa Torre, C. G, DE, and Harms, A.: Genera Siphonogamarum ad sys- tema Englerianum conscripta. Fasciculus tertius (signatura 21-30). Small 4to. pp- 161-240. Leipzig: Wilhelm Engelmann. 1go1. 4/4. 3BoT. GAZ. 30:67. 1900. an grasses. II. Revised edition. 8vo. PP. 349. Bulletin 17, Division of eae: U.S. Department of Agriculture. 1901 e flora of the Palouse region. 8vo. pp. eS) Published by the Wash- ington Aevcnltuce College and School of Science, Pullman. May 14, Igol. 1901] CURRENT LITERATURE 63 Idaho, the region consisting mainly of rolling hills, destitute of trees and and western Idaho. These hills are generally called Palouse hills, and hence the title of the book. The manual certainly covers a region of very great interest in its unrivaled floristic riches. The keys and descriptions seem to be entirely adequate, and, checked as they have been by the large field experience of the authors, the manual must give as good a presentation of the flora as is possible at present. The nomenclature follows what are called the Kew and Berlin rules. Several new species are described, and the enumeration of species shows 14 pteridophytes, 9 gymnosperms, I14 monocotyledons, and 526 dicotyledons. The exceedingly varied conditions of the western mountain en will demand the publication of just such local manuals as this.—J. M. C. HE LIVERPOOL Marine Biological Committee is doing good work in publishing short popular papers on the more interesting animals and plants in the general region of its activities, the Irish sea. The first three papers described animals, but the fourth, just issued, is on Codium,® a very interest- ing genus of the Siphonales. Following a general introduction, we have presented an account of the structure, reproduction, habits, and distribution of this alga. The life history is still incomplete in certain phases of repro- duction. There are two forms of sporangia, one producing large green zoospores and the other small yellow elements, both however morphologically similar and biciliate. The larger green zoospores will germinate vegeta- tively, and the problems concern the fate and function of the small yellow bodies. They have been supposed to be sperms that should fuse with the green swarmers, but no unions have ever been observed. It is probable that the yellow zoospores are gametes, which under suitable condition will conju- gate with one another. The authors of the paper suggest that the plant is becoming apogamous, a view that has support, further than the mere nega- tive evidence, in the fact that the hypothetical gametangia at certain stages in their development may be reproductive. They are then adventitious buds, capable of growing out in a branching filament, which however appears to remain attached to the parent plant. The paper is illustrated with three very clear plates.— B. M. Davis NOTES POR STUDENTS. HE POLLEN tube in Cucurbita Pefo according to B. Longo’ traverses the tissues of the funiculus and outer integument before entering the °GIBSON and AULD: Codium. pp. viii +18. f/s. 3. L. M. B. C. Memoirs. IV. vi a mesogamia nella commune zucca (Cucurbita Pepo Linn.). Rendiconti della a1, R. Accademia dei Lincei ro: 168-172. IQOI. 64 BOTANICAL GAZETTE [| JULY micropyle. The writer proposes the r name mesogamy for this phenomenon.— CHARLES J. CHAMBERLAIN. Dr. N. WILLE® has begun a series of studies on the Chlorophyceae. The first paper contains numbers one to seven, and treats of the structure and phases in the life histories of Syézdion Droebakense, n. sp., Trochiscia, Prast- ola crispa, Ulothrix flacca and several new species of Ulothrix, Pseudendoclo- nium, nov. gen., and certain species of Rhizoclonium. Four excellent plates accompany the text.—B. M. Davis As A RESULT of experiments conducted in Maryland in the seasons of 1899 and Igoo, C. O. Townsend finds? that celery blight, caused by the fun- gus Cercospora apit, can be prevented by the use of fungicides. The best results, apparently, are given by ammoniacal solution of copper carbonate, rdeaux mixture being equally good as a fungicide but causing a slight stunting of the plants. To obtain the best results the first application should be made while the plants are still in the seed-bed, followed by other applica- tions every week until cool weather, which checks the progress of the disease. —ERnSsT A. BESSEY. _ THE SUBJECT of oat smut has received attention in two recent bulletins from the Illinois*® and Wisconsin™ experiment stations respectively. In both an estimate is given as to the annual loss in the state from this smut, about 14 per cent. for Illinois and about 6 per cent. for Wisconsin. The results of experiments with the hot water treatment at different temperatures are described in the former bulletin, the temperature found to be most efficient without injury to the germinative power of the grain being 135° F. for five min- utes. Both bulletins discuss the formalin treatment, which is shown to be even more effective than that with hot water and somewhat more easy to perform. — Ernst A. BESSEyY. A SHORT NOTE on the abundant and destructive occurrence of the dwarf mistletoe, Arceuthobium pusillum, in the upper peninsula of Michigan is given in a recent bulletin from the Michigan Experiment Station.? In connection with the recent rediscovery of this interesting plant in New England, it is of paca Dr. N.: Studien iiber Chlorophyceen. Med. f. d. biol. sta. v. Drébak. no. 2. 19 9 TOWNSEND, C. O.: Notes on aie’ blight. Bull. Maryland Agr. Exp. Sta. no. 74. pp. 167-182. igh 1-7. May Igor *SHAMEL, A. D.: Treatment of oats for smut. Bull. Ill. Agr. Exp. Sta. no, 64, pp: 57-72. 6 pls. Urbana. March 1901 ™Gorr, E. S.: Lvs Gina, of oat smut. Special Bulletin, Wisconsin Agr. Exp. Sta. pp. 1-4. fig. Madison. March 1go1. @ WHEELER; C. F.: The geology and botany of the Upper Peninsula Experiment ation. Report of the Upper Peninsula Experiment Station for the year 1900. Bull. ace Agr. Exp. Sta. no. 186. PP: 17-28. 4 pis. Dec. 1 SS 1901] CURRENT LITERATURE 65 interest to note that in Michigan it is in some places so abundant as to kill out nearly every tree. The parasite itself is attacked by a fungus, Wad/rothiella arceuthobtt Peck, which apparently serves to keep it in check somewhat. In the same bulletin are given notes on the trees and a list of the flowering plants growing on the station farm, as well as a list of the few of the com- moner diseases of cultivated plants observed there.— Ernst A. BESSEY. D. T. MAcDouGAL"*s has studied the bulbils which are formed in the axils of the aerial stems of Lysimachia terrestris, and regards them as repre- senting a new category of propagative bodies. They are branches of restricted development, and are formed under conditions unfavorable for seed formation, diffuse light and low temperature apparently being the prin- cipal inciting causes. noe a are a baie bean sues! organs of any kind, and resemble rhizomes ins which they are borne. The “germination”’ of the bulbil occurs without any 7 appreciable rest- ing period, and is followed by the final stages of the oe of the stele, which was checked during the formation of the bulbil. ulbil becomes the main axis of the new plant, and does not Pee except gradu- ally, after the manner of a rhizome, into which it be —J.M.C. MUSHROOMS ARE DISCUSSED in two recent Experiment Station bulletins, one from Idaho™ and the other from North Carolina. Both give rules for avoiding poisonous fungi and analyses showing the food value of the edible species. This has been greatly overrated, for, compared with other foods, these fungi have not only a small heating power but have also a low nitrogen content. In the former bulletin a few species are popularly described with the aid of good half-tone plates, while in the bulletin from North Carolina all or nearly all of the edible species reported in the state are described, rather too technically it would seem. is technicality, combined with the lack of illustrations, leads one to fear that it will not be serviceable as a popular guide. The glossary, five pages in length, is no doubt necessary but might have been made more accurate. The definition of basidia as “cellular processes of certain mushroom-bearing spores”’ is probably a typographical error, but is plainly very misleading as it stands.— ERNST A. BESSEY THE THIRD EDITION of Sturgis’s Literature of Plant Diseases** brings the *3 Propagation of ee terrestris. Bull. N. Y. Bot. Garden 2: 82-89. IgoI. ™ HENDERSON, L. ushrooms or toadstools: a poner food product. Bull. Idaho Agr. Exp. Sta. no. phy Pp. 27-54. figs. 1-72. Moscow. Marc *S Hyams, C. W.: Edible mushrooms of si Carolina. Bull. N. C. Agr. Exp. Sta. no. 177. pp. 25-58. West Raleigh. Dec. 1900. © Sturais. W.C., Literature of plant diseases. A provisional bibliography x the more important works ‘pabhabed by the U. S. Department of Agriculture and the cultural experiment stations of the United States from 1887 to 1900 inclusive, on ye gous and bacterial diseases of economic Saag hae Conn. Agr. Exp. Sta. for year ending Oct. 31, 1900. Part III. pp. 255-297. 66 BOTANICAL GAZETTE [JULY bibliography of these diseases nearly up to date. The first edition appeared in 1893 and the second in 1897. It does not assume to be a complete bibliog- raphy, but a reference list ‘‘to enable the practical observer of plant dis- eases to ascertain what are the principal sources of information regarding the specific cause of a certain disease and the method of prevention as recorded in the publications of our own Department of Agriculture and of the various state experiment stations.” As in the previous editions, the host plants are arranged alphabetically under their common names. A number of diseases formerly ascribed to parasitic organisms but since shown tobe due to other causes are omitted, while many new host plants whose diseases have been studied in this country only in recent years are added, making the list now somewhat longer than before. The work is exceedingly useful not only to the practical worker but also to the specialist.—ERNsT A. BESSEY. THE ZYGOSPORE OF SPORODINIA was studied six years ago by Léger, who found that both gametes contain hundreds of small nuclei which become scattered in the mingling cytoplasm when the membrane separating the gametes breaks down. The nuclei near the periphery are much smaller than those nearer the center. At a later stage all the nuclei disappear, and at each pole of the zygospore there is found an “embryonic sphere” contain- ing a large number of granulés. The spheres increase in size and fuse with each other, and soon afterward numerous nuclei again appear. Gruber” has examined Sporodinia, and he also finds a large number of nuclei in the zygo- spore. The nuclei are more numerous at the periphery, but those at the periphery and those at the center are approximately alike in size. This condition persists for a long time, and subsequent stages were hard to follow. No fusion, division, or disorganization of nuclei could be established with any certainty. The presence of “embryonic spheres” is regarded as doubtful. n germination the nuclei appear in greater numbers and pass into the germ tube. Although the writer was not able to observe any fusion of nuclei, he believes that a fusion of nuclei at the center of the zygospore is very probable. —CHARLES J. CHAMBERLAIN. THE EFFECT of fungicides upon the foliage of the peach is discussed by W. C. Sturgis in a recent report of the Connecticut Agricultural Experiment Station.” The experiments were made with various strengths of Bordeaux mixture, with a soda-Bordeaux, in which soda replaced the lime, with ammoniacal solution of copper carbonate, with copper acetate, and with potassium sulfid. The Bordeaux mixture was found to be injurious to the *7 GRUBER, EDUARD: Ueber das Verhalten der Zellkerne in den Zygosporen von Sporodinia grandis Link. Ber. d. deutsch. bot. Gesell. 19: 51-55. f/. 2. 1901 SturGis, W. C.: Peach-foliage and fungicides. Report of the Connecticut Agricultural Experiment Station for the year ending October 31, 1900. Part III, pF- 219-254. pls. 3-5. 1901 1901] : CURRENT LITERATURE - 67 foliage except when very weak solutions were used. The soda-Bordeaux was also injurious, as was the ammoniacal solution of copper carbonate. Potas- sium sulfid, however, proved to be harmless, and at the same time to bea fairly good fungicide. Normal copper acetate solution was harmless but the subacetate caused injury. Careful comparative examinations of leaves of plants susceptible to injury by copper-containing fungicides, viz., peach, Japanese plum, and apricot, with leaves of plants not so injured, viz., European plum, apple, cherry, quince, and pear, failed to reveal any constant difference in the thickness of the leaves as a whole, or of the epidermis and the different layers of tissue, or in the size or number of the stomata. The susceptible leaves, however, had a very dense spongy parenchyma, with small intercellular spaces, while the non-susceptible leaves had this tissue very loose in texture.— ERNST A. BESSEY. CLEISTOGAMOUS FLOWERS” are found in nearly all violets, but are espe- cially typical in Viola odorata, The normal flower which appears early in the spring has a handsome corolla, but it seldom produces good seed. The inconspicuous cleistogamous flowers which come later, usually after the nor- mal flowers have disappeared, produce an abundance of good seed. The stamens are larger in the normal flowers than in the cleistogamous, but the size of the ein ene is about the same in both. The structure of the anther wall is rent, the normal anther having the usual endothecium with lignified iiciencician whikess in the cleistogamous flower the endothecial layer retains its nucleus and cytoplasm. After the pollen is mature there is a resting period of various duration. Pollen tubes are then put out which penetrate the wall of the anther at its upper part where there is a region of small cells rich in protoplasm, a tissue comparable to the conductive tissue of the style. Oxalis acetosella, Linaria spurta, and Leersia oryzotdes were also studied. In typical cleistogamous flowers the pollen germinates within the pollen sac, and the structure of the anther wall is modified to meet the new mode o pollination. In Linaria and Leersia, where the pollen was not observed to germinate within the pollen sac, the anther wall has the same structure as in the normal flower.— CHARLEs J. CHAMBERLAIN. BULLETINS from the experiment stations of interest to botanists, and not heretofore mentioned in these pages, are as follows: A, S. HirTcHcock and G. L. CLoTHIER (Kans. no. 87, pp. 29) write upon the “ Native agricultural grasses of Kansas,” with many illustrations and charts of distribution. H. GARMAN writes on the agricultural grasses of Kentucky, with some fine reproductions from photographs, and A. M, PETER supplies some chemical analyses, the two articles forming one bulletin (Ky. no. 87, pp. 68, A/. 74). 7 Du SaBLon, LECLERC: Recherches sur les fleurs cléistogames. Revue Gén- €rale de Botanique 12: 305-318. s7 figs. 1900. 68 BOTANICAL GAZETTE [juLY F, A. WAUGH (Vt. no. 67, pp. 30) discusses hybridity among cultivated plums, and gives a systematic account of hybrid forms. Interesting trees of Vermont are described and figured by ANNA M. CLARK (Vt. no. 73, pp. 52), and those of Wyoming by AVEN NELSON (Wy. no. 40, pp. 52). W. W. ASHE (N.C. no. 175, pp. 8) gives technical diagnoses of 21 new species Crataegus and 8 new species of Panicum, E.E, BOGUE publishes “An annotated catalogue of the ferns and flowering plants of Oklahoma” (Okla. no. 45, pp. 48), D. A. SAUNDERS does the same for South Dakota (S. D. no. 64, pp. 127), and Henry L. BoLiey and L. R. WALDRON do the same for North Dakota (N. D. no. 46, pp. 91), all excellent beginnings toward complete floras of the respective states. C. F. WHEELER writes about the dwarf mistletoe in Michigan, with good reproductions from photographs, and on other topics (Mich. no. 186). L.H. PAMMEL describes the horse nettle (Solanum Caro- linense), bind weed (Convolvulus arvensis), and ground burnut (77ridulus henite as troublesome weeds in Iowa (Ia. no. 42.) D. A. BRODIE (Wash. 0. 45, pp. 12) gives facts establishing the poisonous nature of the Oregon seer nas (Cicuta vagans).—J. C. ARTHUR FERTILIZATION IN Ginkgo éiloba has recently been studied by Ikeno,” who gives a detailed account of phenomena from the cutting off of the ventral canal cell to the first division of the nucleus of the oospore. The nucleus of the ventral canal cell rapidly disorganizes, but in one instance it had enlarged part passu with the nucleus of the oosphere. In pr arations stained with methyl blue and acid fuchsin, the metaplasmic ground substance of the lar mass, also takes the red, while the nucleoli stain blue. e nucleus then undergoes a great change in structure, so that the metaplasm and chromatin can no longer be distinguished from each other. The further development of the nucleus of the oosphere agrees with the description of the correspond- ing phenomena in Pinus Laricio as described by the reviewer in 1899. e tube nucleus and the nucleus of the stalk cell disorganize within the pollen tube and do not enter the oosphere, and it is very probable that only one of the male cells is discharged, the other disorganizing without being able to enter. The nucleus of the male cell slips out from the cytoplasm mantle before fusing with the nucleus of the oosphere. The mode of fusion is like that already described for Cycas revoluta, that is, the male nucleus gradually penetrates the egg nucleus before losing its own membrane. At the time of fusion the sex nuclei are very unequal in size, the female being about ten times as large as the male. The behavior of the chromatin during fusion is not described. The spindle in the first division of the fusion nucleus is very broad and multipolar, and is never parallel with the longitudinal axis of the 7°Contribution 4 l'étude de la Seinen chez le Ginkgo biloba, Ann. Sci. Nat. Bot. VIII. 13: 305-318. pls. 2-3. 1gor } CURRENT LITERATURE 69 oospore. In the case figured the spindle is transverse. Fertilization occurs while the ovules are still on the tree—CHARLES J. CHAMBERLAIN DOUBLE FERTILIZATION in Zea Mays, which has been suspected for some time, and which is believed to be the cause of xenia, is described in a recent paper by Guignard.* The mature pollen grain contains, besides the vegetative nucleus, two very small elongated male cells, each in the form of a slender rod, curved or straight, and the ends often pointed. The cytoplasm of these cells is much reduced and difficult to distinguish, and their nuclei appear almost homogeneous. The synergids and oosphere are large, the former showing near the tip a conspicuous longitudinal striation, especially in material fixed in absolute alcohol. The nucleus of the oosphere is very large and contains much chromatin, and the cytoplasm is usually highly granular and much massed together at the time of fertilization. Near the oosphere, sometimes in the median plane, sometimes near the side of the embryo sac, are the two polar nuclei which do not fuse before fertilization, and have relatively large nuclei and a small amount of chromatin. As many as a dozen multinucleate cells may be found in the much narrowed antipodal end of the embryo sac. The pollen tube, after penetrating the embryo sac, usually seems to discharge its contents into one of the synergids. In one instance the two elongated male cells were observed resting against the base of a synergid; under high magnification their chromatin was distinct. One of the male cells unites with the oosphere, the other with the polar nuclei, which it binds together. Fertilization proceeds with such great rapidity that it could be observed in very few preparations. In general, the ovules at the base of the ear are first fertilized, and in hybrids many ovules are not fertil- ized. After fertilization one of the synergids usually persists for a time, ‘with the contents finely granular and refractive. Division of the definitive nucleus proceeds so rapidly that the author was not able to follow the course of cell division. The first two nuclei of the endosperm are large, each one having an enormous nucleolus and many smaller nucleoli. It is to be regretted that no figures are given.—W. J. G. LAND. THE EMBRYOLOGY of the Balanophoraceae presents many puzzling peculiarities. Accounts are somewhat divergent, but whether the divergence is due entirely to variation in the processes still remains to be seen. Writers agree that there is no ovule or placenta in Balanophora but that the mega- spore is situated in a tissue at the base of a prolongation incorrectly termed a “style.” Van Tieghem (1896) found that in B. zvdica the polar nuclei do not fuse and that fertilization occurs at the antipodal end of the sac as often as at the upper end. According to Treub (1898) in B. elongata the megaspore germinates in the usual manner. The polar nuclei, however, do not fuse but each divides 2t La double fécondation dans le mais. Jour. Bot. 15:1-14. 1901. 7° BOTANICAL GAZETTE {JULY independently. The egg apparatus breaks down and there is no fertilization, but an embryo develops.from one of the cells of the endosperm. Lotsy (1899) investigated B. globosa and esa Treub in every particular, including the peculiar origin of the em Chodat and Bernard have sean ‘iveieeuted flelosis guayanensts.” The archesporial cell becomes the megaspore directly without cutting off a tapetal cell or giving rise to a row of potential megaspores. The jacket or “tapetum” surrounding the embryo sac is sporogenous tissue. The two daughter nuclei resulting from the first division of the nucleus of the megaspore are quite different in appearance, the one at the upper end of the sac staining much more deeply. This nucleus gives rise to the egg, two synergids, and a polar nucleus in the usual manner. The other nucleus stains faintly and rarely divides at all, but soon degenerates so that no antipodals or polar nucleus are formed. According to Van Tieghem, the egg is fertilized in Helosis and Balanophora. The present writers find that in Helosis the egg becomes large, but also becomes very weak and feeble in appearance, so that while they were not able to prove or disprove the occurrence of fertilization, they believe that the feeble condition of the egg together with the position of the embryo in the endosperm favor Treub’s view that the embryo arises apogamously from the endosperm.— CHARLES J. CHAMBERLAIN. ITEMS OF TAXONOMIC interest are as follows: E,. P. BICKNELL (Torreya 1: 25-28. Igo01) has described a new Triosteum (7. aurantiacum) from the northeastern United States.—P/. Bakerianae 2: 1-42. 1901 contains Baker's collection of 1899, from fungi to grasses. Numerous new fungi are described y F. S. EARLE.—ALIcE EAstwoop (Bull. Torr. Bot. Club 28: 137-160. Js. 15-20. 1901) has published upon some small-flowered species of Nemophila from the Pacific coast, describing twenty-six new species M. A, HowE (tdem 161-165) has described a new Riccia from Georgia— E. P. BICKNELL (idem 166-172) has revised the eastern species of Teucrium, recognizing six species and describing four as new.—P. A. RYDBERG (édem 173-183), in further studies of the Potentilleae, describes new species of Potentilla (8), Horkelia, and Drymocallis.—W. A. SETCHELL (Zoe 5: 121-129. Igo!), in his ‘* Notes on Algae,” has described two new genera of Laminaceae (Hedophy!- lum and Pleurophycus) and a new genus of Dumontiaceae (Weeksia), besides several new species in other genera.—A. ENGLER (Bot. Jahrb. 36: 29-126. Ig01), in his 21st contribution to the African flora, presents the following papers: Fungi by P. HENNINGs, who describes numerous new species and two new genera, Fistu/inel/a (Polyporaceae) and Lactariopsis (Agaricaceae); Algae by W. SCHMIDLE; a revision of Schrebera (Oleaceae) by E. GILG; ODAT, R., and BERNARD, C.: Sur le sac embryonnaire de ]’Helosis guaya- nensis. Jour. de Botanique 14 : 72-79. p/s. 7-2. 1900. 1901 | CURRENT LITERATURE ay Leguminosae by H. HAkms, Scorodophiloeus, Rhynchotropis, and Scheffiero- dendron being new genera; Myrsinaceae by E. GILG; Amarantaceae by G. LoPRIORE, Argyrostachys being a new genus; Acanthaceae by G. LINDAU; Caricaceae by I. URBAN, Cylicomorpha being a new genus; and Gramineae by R. PILGER.— CARL MEz (zdem Beibl. 30: 1-20. 1901) has described numerous new species of Bromeliaceae and Lauraceae.—I. URBAN (édem 27-38) con- tinues his papers on new American plants.—A. ENGLER (édem 42) has’ described a new genus of Araceae (Protarum) from the Seychelles.— B. D GILBERT (Fern Bulletin 9:27. 1901) has described a new variety of Botry- chium ternatum (Oneidense) from central New York.—ACHILLE Fort! (Ber. deut. bot. Gesell. 19: 6-7. 1901) has published a new genus (//eferoceras) of marine Peridineae.—-W. SCHMIDLE (zdem 20-24) has described a new genus (Coccomyxa) of the Protococcoideae.—C. S. SARGENT (Rhodora 3:71-79. 1g01) has described six new species of Crataegus from the Province of Quebec near Montreal.-—G. P. CLINTON (dem 79-82) has described two new smuts on £riocaulon septangulare.— E, L. GREENE (dem 83-84) has segre- gated from Lupatorium ageratoides the bulk of the New England and north- ern forms that pass under that name, and called the species &. boreale.— C. DE CANDOLLE (Bull. Herb. Boiss. II. 1: 353-366. 1901) has published an account of the Brazilian Piperaceae and Meliaceae collected by W. Schwacke, including descriptions of numerous new species.—R. CHODAT (dem 395- 442), in continuing his account of the Hassler collection from Paraguay, has described numerous new species in various families—W. TRELEASE (Rep. Mo. Bot. Gard. 12:77. Af. 34. 1901) has described a new cristate variety of Pellaea atropurpurea from Missouri.—J. W. ToumMEy (zdem 75-76. pls. 32-33) has described a new Agave from Arizona. H. M. RicHarps (Bull. Torr. Bot. Club 28: 257-265. f/s. 27-22. 1901) has described a new genus Ceramothamnion) of red algae from Bermuda.—P. A. RYDBERG (idem 266- 284) has published a fifth fascicle of new species from the Rocky mountain region, among them being Piferda, a new genus of orchids.—J. K, SMALL (idem 290-294) has published a third paper on the shrubs and trees of the southern states, including a revision of the southeastern species of Pte/ea.— In Torreya (1: 54-55. 1901) J. K. SMALL has published a new Cornus from Kentucky, and N, L. BriTTON a new Crataegus from Washington.—F. Lam- SON-SCRIBNER and E. D. MERRILL (Rhodora 3: 93-128. 1901) have pub- lished a revision of the New England species of Panicum, recognizing thirty- five species.—J. K. SMALL (Bull. N. Y. Bot. Garden 2: 89~101. 1901) has published a synopsis of the Mimosaceae of the southeastern United States. He recognizes fourteen genera, and among them Siderocarpos and Havardia have been separated from Pithecolobium as new.—M. A. Howe (idem 101- 105. £7. 7z) has published an enumeration of the liverworts collected in the Yukon Territory by R. S. Williams in 1898-9, including a new species of 72 BOTANICAL GAZETTE (yuLy Scapania. Mr. WILLIAMS himself (zdem 105-148. ps. 75-24) has enumer- ated the mosses, including a new genus (Sryobrittonia) closely related to Tortula and Desmatodon, and a number of new species. L. M. UNDERWOOD (tdem 148-149). enumerates the pteridophytes ; while the seed plants are presented by N. L. BriTTON and P. A. RYDBERG (zdem 149-187), numerous new species being described.—P. A. RYDBERG (idem 187-233 Pls. 25- ~337) has given an account of the oaks of the continental divide north of Mexico, recog- nizing twenty-nine species, nine of which are new.—J. M. C PROFESSOR A. B. MACALLUM® has recently added an interesting contri- bution to the cytology of certain so-called non-nucleated organisms. His work is divided into three parts, each dealing with a separate group of low organisms—the Cyanophyceae, Beggiatoa, and the yeast cell—and was undertaken with the hope of throwing some light on the origin of the cell nucleus, and to obtain data to determine the morphological character of the primal life organism. In his investigations Macallum not only used the ordi- nary cytological methods, but microchemical tests were also employed to advantage. Picric acid and corrosive sublimate afforded the best results as carmin, and methylen-blue. The microchemical methods employed for obtaining the reactions for ‘masked iron” were practically the same as those used in his earlier work published in 1896. The iron liberated by sulfuric acid alcohol was converted into Prussian blue, the trichomes were then stained with a picro-carmin solution for twenty-four hours, when the cyano- phycin granules acquired a deep red color which contrasts markedly with the Prussian blue tint of the iron-holding granules. The results obtained on the _ Cyanophyceae are briefly as follows. The cell consists of two portions, the central body and the peripheral zone holding the pigment. There is no evi- dence of the presence of a special chromatophore. There are two types of granules present in the cell. The one stains with haematoxylin, contains “masked iron” and organic phosphorus, and therefore resembles chromatin. The other type is found in the peripheral layer, and chiefly adjacent to the cell membrane. It stains with picro-carmin, and is free from organic phosphorus and “masked iron.” It is probably a proteid. There is no nucleus or any structure which resembles a nucleus in the Cyanophyceae. In Beggiatoa there is no differentiation of the cytoplasm into a central body and a periph- eral layer, such as Biitschli describes. The compounds of ‘masked iron” and organic phosphorus are uniformly diffused throughout the cytoplasm in the threads. In the “spirilla,” “comma,” and “coccus"’ forms the cytoplasm shows characters like those of the threads, but there are also granules present 23On the nin ie of non-nucleated organisms. University of Toronto studies. Physiological series 2. 1900. Igol] CURRENT LITERATURE 73 which give a slight reaction for “masked iron” and organic phosphorus, and therefore are considered analogous tochromatin. No specialized chromatin- holding structure in the shape of a nucleus was found in any of the forms of Beggiatoa studied. In his studies on the yeast cell, Macallum finds that the cytoplasm takes a stain with haematoxylin and gives a diffuse reaction for “‘masked iron’’ and organic phosphorus. In addition to the chromatin-like substance diffused throughout the cell, there is usually present a homogeneous corpuscle. This is not considered to be a nucleus, although held to be such by other investigators. The chromatin-like substance in Saccharomyces is soluble in artificial gastric juices, thus differing from the chromatin of the higher plant and animal cells. The paper is illustrated by a colored litho- graphic plate. It is a valuable and highly interesting addition to the litera- ture of this important problem.—A, A. Lawson. NEWS. Dr. G. T. Moore, of Dartmouth College, has been appointed phycologist in the Department of Agriculture. Dr. J. C. ARTHUR sailed July 6 and will spend July and August in Europe with his bride. He will attend the botanical conference at Geneva in August. ; Dr. J. B. OVERTON, who received his doctorate in June from the Uni- versity of Chicago, has been appointed professor cf botany in IIlinois College, at Jacksonville. Mr. A. A. Lawson, fellow in botany at the University of Chicago during the past year, has been appointed assistant in botany in Leland Stanford Junior University. HENRI PHILIBERT, honorary professor in the Faculty of Letters, dis- tinguished for his bryological studies, died at Aix on the fourteenth of May, in his seventy-ninth year. PROFESSOR G. J. PEIRCE, of Leland Stanford Junior University, takes charge of the botanical work during the summer at the Hopkins Seaside Laboratory near Pacific Grove, California. WE LEARN from Science that a memorial tablet of the late Thomas Con- rad Porter, long professor of botany in Lafayette College, was unveiled in connection with the recent commencement exercises ProFressor C, E. Bessey, after attending the meeting of the American Association for the Advancement of Science in Denver, will go to the Pike’s peak region to join the University of Nebraska camp for a month, to study the mountain flora. THE REGENTS of West Virginia University have abolished the professor- ship of botany in that institution without notice to the present incumbent, Dr. E. B, Copeland. Dr. Copeland will spend the summer at the Cold Spring Biological Laboratory. Mr. H. N. WHITFORD, assistant in botany in the University of Chicago, has been appointed a collaborator of the Bureau of Forestry for the study of the ecology of the forest. He is to carry on work this summer in the neigh- borhood of Flat Head lake, Montana. . C. Cowes, of the University of Chicago, will conduct a field party selsinns August in northwestern Montana. The party will number about 74 [JULY gor] NEWS 75 twenty, and the work will be along ecological lines. Most of the work will be in the neighborhood of Flathead lake. where the state biological station is located. THE AMERICAN ASSOCIATION for the Advancement of Science will meet at Denver, Colorado, August 23-28, rgot. A preliminary program of the meetings of section G (Botany) will be issued about July 15. One day will be given up to a joint session with the Botanical Society of America. It is also planned to devote one day’s program to the subject “Adaptations of desert plants.”’ Dr. F. E. CLEMENTS is to spend the summer in the Pike’s peak region in Colorado, engaged in ecological studies of the flora. He will be accom- panied by a party of botanists from the University of Nebraska, numbering a dozen or more. Instruments for accurate observation of ecological factors have been provided for the party. THE LATEST Bulletin of the New York Botanical Garden, issued May 27 last, contains much information of interest to botanists. The planting of the grounds is proceeding rapidly, and the present showing of species grown during the year is as follows: herbaceous grounds 2300, fruticetum 450, salicetum 40, arboretum 220, and viticetum 60. During the year 48,895 specimens have been added to the herbaria. JOHN J. THORNBER has been appointed special botanical collector for the University of Nebraska for the summer of 1901. He is to accompany the field party of the United States Forestry Division now at work in Nebraska, and is to act as its botanist, at the same time being the botanical representative of the university. In addition to the collection of specimens, he is to make careful ecological studies throughout the territory traversed. UNDER A commission from the United States government, Dr. H. von Schrenk, of the Shaw School of Botany, is to spend the summer in Europe, in an investigation of the problems connected with the decay of railroad ties on the principal roads, this work being done in connection with an extensive series of investigations into the same subject which he is undertaking for the Department of Agriculture and in which the principal American railroads are cooperating. On MAY THIRTIETH there occurred the unveiling of a memorial tablet to Asa Gray in the Hall of Fame of the University of New York. The ceremony was committed to the Botanical Society of America. Most appropriately Dr. B. L. Robinson, professor of systematic botany in Harvard University and curator of the Gray herbarium, a ae N. L. Britton, nese of the New York Botanical Garden, Pp I f the society, Dr. Robinson unveiling the tablet. SUMMER FIELD WORK will be undertaken by the staff of the School of Botany of the University of Texas as follows: One party will make an exploration 76 BOTANICAL GAZETTE [yuLY of Padre island between Corpus Christi and Brownsville. A member of the staff will accompany a party conducting the mineralogical survey, which will make an expedition into the trans-Pecos region of the state. A party of students will collect plants and study ecological questions from Cuero west- ward toward the Pecos river. These various expeditions should do much in making known the state botanically. THE BOTANICAL staff in the State University of Iowa has organized a summer school of botany to be conducted along the shores of Lake Okoboji in northern Iowa. The region is not only one of the most delightful of western summer resorts, but has unusual attractiveness for the naturalist. The uni- versity offers this year to students of botany a laboratory with all essential apparatus and libraries. Abundant boats put all the lakes into easy com- munication with one aaother, and in addition the laboratory will have its own equipment for the use of students. Professor Shimek will act as director and will be aided by special assistants. It is expected that hereafter the work in botany for the summer session of the State University of Iowa will all be con- ducted at Okoboji. Dr. J. N. ROSE left about the 2oth of June for his third botanical trip to Mexico. He expects to go first to the City of Mexico, working out from this point as a base southward towards Acapulco and eastward toward Vera Cruz, probably ascending Mount Orizaba and Popocatepetl. The objects of his trips are to make a general botanical collection; to collect at type localities certain species of Humboldt, Galeotti, Schiede, and other early collectors ; and to acquire information regarding the economic uses of Mexican plants, especially such as will supplement a second paper on the useful plants of Mexico, which is now nearly completed. ERRATA in the last volume reported too late for inclusion in the usual list are as follows: P. 109, line 4, after pellucido-punctata insert comma. P. 121, line 8 from below, after corticata insert comma. P. 392, line 14, for ERIGERON read ERIOGONUM. P. 393, line 6, for TROGANUs read T1oGanus. 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DBAS WE WE WE DE WS SIE WEBER Giardina | Waukesha Hygeia Mineral Springs Water OM IT IS MADE THE FAMOUS BORO. LITHIA WATER GINGER ALE AND WILD CHERRY PHOSPHATE y THE WAUKESHA WATER co. 132 N. Jefferson Street, Chicago Telephones - - Monroe 1166 and 1168 “debility pra hired a tardy valesc ~ Sheet and Beau dy conv. Ses ai Sy miner Bete. | mineral vi eps have ‘ seen the ~ IP Hg aluable Adjunct to the BUFFALO LiTHA WATER *2 Dr. — Ben’ Johnston, Prien of Surgery in the Virginia Medical College, mond, Va., 4s L THK in my opinion has a larger range of usefulness any other mineral water. Among the nu- onditions to which | have Ko it with good results may be ia correspondence regardi: ng su bscript 107 The Usher of Chicago Press, Chica cage, il. As dt Til d-class mail matter. ] 4 ‘oY > Every Botanist Should be familiar with the prominent works of GEBRUDER BORNTRAEGER Publishers. Die Glykoside. Chemische Monographie der Pflanzengly- koside nebst systematischer Darstellung der kiinstlichen Glykoside von Dr. I. I. L. von Ryn, Director der Reichsver- suchsstation in Mastricht. 8vo. Cloth, $2.50. Das Werk giebt—wie es bisher noch nirgends geschehen — eine eingehende chemische Behandlung der Glykoside—nicht nur eine kurzgefasste Zusammen- stellung der chemischen Eigenschaften dieser Korperklasse, sondern die Darstel- lungsmethode, die Griinde, welche zur Aufstellung der Constitutionsformeln gefithrt haben etc., so dass das Buch in chemisch-pharmaceutischen wie pharma- hkologischen Kreisen sowte unter den Studirenden und sonstigen Freunden der phytochemischen Forschung sicher mit grosser Freude begriisst werden wird. Die Harze und die Harzbehalter. Historisch kritische und experimentelle, in Gemeinschaft mit zahlreichen Mitarbeitern ausgefiihrte Untersuchungen yon Prorrssor DR. A. Tscuircu, Director des pharmaceutischen Institutes der Unt- versitat Bern. Mit 6 Tafeln. 8vo. 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PRI ICE $1.50, NET This book contains directions for omni igs 5 cage te sei plant material eis microscopic sponge: sage It is based rse botanical mic technique and is the first complete soles to i iabllahed on this par in rsi y: It aims, therefore, to meet the requirements only of the student who has the assistance of an a in a fully peste laboratory, but, also, the student who must work by himself and with limited apparatus. hand ne the pa araffin acsed, 5 prea method, and the glyce method are treated in considerable det ater chapters specific Ti sectican are given for star such prepar nae as are needed by those who wish to ae wh structures. Formulas are given for the reagents commonly used in the Histo- logical] Laboratory. For Sale by Dealers, or by the Publishers THE UNIVERSITY OF CHICAGO PRESS, CHICAGO, ILL. [JUST PUBLISHED] Send 12 cents, U. S. stamps, for ; Wm. Wesley 8& Son’s Botanical Catalogue, 1901 J i Being CONTENTS: No. 137-138 of Transactions of Scientific Societies Periodicals Bibliography § History Biographies and Portraits Herbals Early Botanical Science Linnaeus NATURAL HISTORY and —— . SCIEN Microscopy Morphology aad Physiology << Encyclopaedic works Classification Nomenclature Cryptogams Phanerogams Fossil Plants Natural distribution of Plants (Floras) Agriculture and Horticulture to the end of the J8th century Gardening Landscape Gardening The Flower and Ornamental Garden Husbandry Tropical Agriculture Commercial Plants _ Medical Botany Forestry Diseases of Plants William Wesley & Son, Booksellers, 28 Essex Street, Strand, ee London, England. SISSIES BOOK CIRCULAR More than 3300 works, classified under 42 headings BRS S RES RRR, — The Journals of the University of Chicago BEING THE DESCRIPTIVE LIST OF ONE WEEKLY, FIVE MONTHLY, ONE BI-MONTHLY, THREE QUARTERLY, AND ONE SEMI-QUARTERLY PUBLICATIONS & #% & THE BIBLICAL WORLD Edited by President W, R. Harper, A mon sis aaaaenk Subscription price, i in the Uni $2.00 a year; foreign, $2.50; single copies, 20 ce The Biblical World i is devoted ete wa to ‘bibli- opular ae oe tes SY — Sunday-school teacher, and the thi siti laym THE SCHOOL REVIEW tape monthl yin the P ies, 20 , except in July and August. Subscription nited Priguagen $1.50 a year; foreign, $2.00; So adequately has the Sc, apt re eatidys served th interests of high-sch work that it has come to be rec secondary education in the U devoted exclusively to thi Pg and helpful, and is indispensable to every THE BOTANICAL GAZETTE Edited oy ge M. Coutter. Published monthly, with illus- ni sg 9p on dp rico in the United States, $4.00 pees ‘forei n, $4.50; single copies, 50 cents. e Bota me Goxette is ee illus he bea 9s journal veiges ahei botany in — more than twenty years it has ee the nepeon tive huceriean journal of botany, pete ning contri- butions from the leading botanists of America and Europe. THE AMERICAN JOURNAL OF SOCIOLOGY Edited by AtBion W, Smatt, Published bi- pred with phon Vege “Subscription price, in the United = es, $2.00 oie 3 foreign, $2.50 ies, 35 ¢ eee aim of the American Seas of Soct- problem” is both me out the general tenden- cies in the rapidly changing field of sociology. - THE AMERICAN JOURNAL OF SEMITIC LANGUAGES AND LITERATURES Edited by Presid ILLIAM R, HARPER. sy apne + Subscription p 8 Aa in the = aa Saas $3.00 a year; for eign, $3.25; single cop The object of this journal to encourage the study of the roc engi a s and ggg to ish information c ing the of Semitic Pp at home and abroad, ,and to Minis asa medium rtments. Articles are published in German, ene and Latin, as well as in English. THE JOURNAL OF GEOLOGY Edited by T. C. CHamBERuIN. Published semi-quarterly, with illustrations. Subser iption price, in the United States, $3.00 a year; foreign, $3.50; single copies, 50 cents. Devoted to see ages of geology and the — sciences, and con rticles covering a wide range of subjects. Adapted ‘ss young geologists, nivaaell students, and teacher THE ASTROPHYSICAL JOURNAL Edited by Gzorce E, Hare. Published a cept February and August, with illustrations. baci price, in the United States, $4.00 ar; foreign, $4.50 =e ae 50 cents, An ernational anid of cag 8 and a iin erae physics uable are in- terested in astronomy ei poreaal ens THE JOURNAL OF POLITICAL ECONOMY Edited by J, Laurence LAUGHLIN, Published bed | ae ription price, in diet nited States 3 for- n, $3.40; single copies, re cents, This publication promotes the a treatment of problems in — cal economics, and also con- tains contributions on topics of ‘theneetiell and speculative arses t. THE AMERICAN ‘scan OF THEOLOGY of Chicago. nitedStates 30 a pears got ate world so catholic in its nt me of modern Pasi” rch in all the different of Scclsnical though rahilen by _— felds and particular schools The pe ae as sto THE ELEMENTARY oe TEACHER AND THE COURSE OF drone Edited by Francis W. P. . Published monthly, except i August sad eaebe "Subecriptiois price, in the Uae Ngan $1.50 a year; foreign, $2.00; single copies, eth onthly periodical for teachers and pare ents. . et ‘husbee seca practical plans i teaching n every grade the kindergarten through the high and ae schools. THE UNIVERSITY RECORD lished weekly, oY hectigaieas $1,005 he University Baa’ is the — — Lies? Hessen of the University of Chi ea sda 3 number, enlarged in size, is issued pace SAMPLE COPIES FREE ON REQUES ail THE UNIVERSITY OF CHICAGO PRESS #4 CHICAGO, ILLINO! Sa kc -ehithateididiieeindliebics= 5, SECOND-HAND BOOKS ON BOTANY AND ! will also furnish — that can be aes MATHEMATICS either in or out of print. Send for circ isis L. SMITH, eae Ksell FoR FALLS owa City, Io aS ae Che Elementary School Record A SERIES OF MONOGRAPHS ON THE EXPERIMENTAL SCHOOL OF THE PEDAGOGICAL DEPARTMENT OF THE UNIVERSITY OF CHICAGO Edited by JOHN DEWEY and LAURA L. RUNYON HE object of the Zlementary School Record is to make possible for use in other schools the details of subject-matter and method in the application of modern psychology in education, as demonstrated by the University Elementary School. The plan includes a series of nine monographs, each number contain- ing a record of work done by a group in the school, and also an article concern- ing the work of one department in all grades. The complete series is now ready. THE SERIES, ROYAL 8vo, PAPER $1.25, CLOTH $1.75, NET. | | THE UNIVERSITY OF CHICAGO PRESS | CHICAGO, ILLINOIS / U7, ‘I 6 FOR $38 \" é. fA 2==>74 In 12 Little Monthly Payments Three thousand dailies, weeklies, and a are Je igh tg produce one copy of PUBLIC OPINION. 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No better company than The Prudential Insurance ae of America JOHN F apne ices HO stakes Newark, N. J. TE FOR INFORMATION, DEPT. VOLUME XXXII NUMBER 2 BOTANICAL (,AZEFTE AUGUST, roor GAMETOGENESIS AND FERTILIZATION IN ALBUGO. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. X XIX. FRANK LINCOLN STEVENS. (WITH PLATES I-IV) BEForE the publication of my paper (1899) describing the fertilization of Albugo Buti, the mature oosphere in the Phycomy- cetes was generally conceded to be a uninucleate structure. The question of homologies in the group did not then seem difficult. A. Bliti, however, presented a unique condition, in that about one hundred antheridial nuclei fuse in pairs with an equal number of egg nuclei. This condition seemed remarkable and inexpli- cable, inasmuch as Wager (1896) had studied the fertilization in A. candida, and had described a fusion of single male and female nuclei in a uninucleate oosphere. Such divergence in the same genus led to some speculation. Davis (1900) reexamined 4. candida and fully confirmed Wager’s account, as indeed Berlese (1898) had already done. It is well established, therefore, that there is a simple fusion in A. candida, and my own account (1899) stands for a multiple fusion in A. Buti, Interest also attaches to this genus on account of the presence of a coeno- centrum, a structure of prominence, but of hitherto unknown function, The study of A. Portulacae and A. Tragopogonis was under- taken in the hope that these species might help to explain the 77 78 BOTANICAL GAZETTE [ AUGUST apparent discrepancy between the two species previously studied. The selection of A. Tragopogonis has proved particu- larly fortunate, for it has led directly to a solution of the prob- lem. The conditions in this species, while to a certain extent similar to those described for A. candida, made evident the necessity of a critical reexamination of the latter. I have included A. candida, therefore, in this investigation. The material of A. Portulacae on Portulaca oleracea, and A. Tragopogonis on cultivated Zragopogon and Ambrosia artemisae- folia, was collected at Syracuse, New York. The material of A. candida on Bursa was collected at Columbus, Ohio. All the material'was fixed in chrom-acetic killing fluid and imbedded in paraffin in the usual way. The methods in general were the same as those employed in the study of A. Blitz, and are described in detail in Bor. Gaz. 28: 233. 1899. The Flemming triple stain was used throughout, and I am convinced that it is best for the study of Albugo, although it is exceedingly sensi- tive to manipulation, and the different species of Albugo require quite different treatment. The results of this investigation are presented in three sec- tions, entitled “descriptive,” ‘ phylogeny,” and ‘general consid- erations.’ The first section describes the phenomena observed, avoiding theoretical considerations and presenting only what may be regarded as established facts. The second section considers the phylogenetic bearing of this research. The third section considers the genus from the comparative standpoint, an attempt being made to interpret the significance of the phenomena, and to call attention to anaiogous phenomena in other organisms. The species discussed were kindly determined by Dr. Paul Magnus, who assures me that the names and synonymy are as follows: Atsuco PortuLacar (DC.) O. Ktz. (Uredo Portulacae DC., Cystopus Portulacae Lév.); A. Tracopoconis ( Pers.) S. F. Gray (Uredo candidus Tragopogonis Pers., Cystopus Tragopogonts Schrot., Uredo Tragopogonis DC., Cystopus cubicus Lév., C. spinu- losus DBy., Uredo cubica Strauss); A. CANDIDA ( Pers.) O. Ktz. ( Uredo candidus Pers.). SS eee a ail 1901} GAMETOGENESIS AND FERTILIZATION IN ALBUGO 79 I wish to express my chief thanks to Professor Strasburger for daily counsel during the progress of this research; to Pro- fessor Magnus for names and synonymy; to my wife for the preparation of the slides, and for Plate IV; and to the Univer- sity of Chicago for the advantages derived from a traveling fel- lowship. lx DESCRIPTIVE. ALBUGO PORTULACAE. The early stages of the sex organs in A. Portulacae differ in no essential detail from those described for A. Buti (Stevens 1899, figs. 42, 43, 45, 59-65). The nuclei are distorted as the protoplasm flows into the developing oogonium, but’ as this structure attains its full growth and recovers its turgor they regain their spherical form and enlarge, rapidly assuming the spirem condition. They are more numerous than in any of the other species examined, ranging from 300 to 400 in each oogo- nium, and are smaller than in A. Bit, rendering this an unfavor- able type for cytological study. The aggregation of the cytoplasm into several regions of greater density is the first indication of the development of the oosphere (jfig.z). The aggregations soon coalesce, forming one large mass of fine uniform cytoplasm. Apart from the struc- tural differences between the dense alveolar center and the vacuolate filar periphery, there is a very distinct difference in stain reaction. The dense fine-grained cytoplasm refuses the gentian, but takes the orange G lightly, while the vacuolate peripheral cytoplasm takes the gentian strongly. The uniform dense alveolar region is the rudimentary oosphere, and is cen- trally placed in the oogonium (jig. 3). Throughout this differentiating process the nuclei, which are now in mitosis, are crowded out of the denser masses and come to lie near the larger vacuoles (fig. 2). Therefore, after the denser masses have coalesced and the larger vacuoles are forced into the periplasm, the nuclei are to be found near the rather indefinite boundary between periplasm and ooplasm. This boundary, however, becomes distinct and sharp immediately . 80 BOTANICAL GAZETTE [AUGUST after the nuclei have passed to the periphery. Meanwhile the mitosis advances from prophase to metaphase. The oogonium in this condition presents typically a region of uniform finely vacuolate cytoplasm devoid of nuclei, sur- rounded by a zone of cytoplasm bearing large vacuoles and con- taining the nuclei in metaphase. This condition I have termed the stage of zonation (fig.3). A full understanding of this stage is necessary to interpret it in other species. Zonation presents the first clear differentiation of ooplasm from periplasm at a time previous to the existence of any wall between these parts, and finds the ooplasm nearly or quite devoid of nuclei. Zonation is very definitely and clearly marked in A. Bhi and A. Portulacae, the periplasm and ooplasm being as sharply separated as though an actual wall existed between them, but is much less conspicu- ous in A. Tragopogonis and A. candida, thus rendering these spe- cies more difficult of interpretation. ; Immediately following zonation the nuclei divide, many of them lying in such a position that one of their daughter nuclei reenters the ooplasm. In A. But nuclei are frequently found in late anaphase with one daughter nucleus lying inside of the oosphere and the other in the periplasm, thus affording direct evidence of the derivation of the primary oospheric nuclei. This rarely occurs in A. Portulacae, as the dividing nuclei usually lie quite outside of the line separating ooplasm and periplasm. The results, however, are precisely those presented in A. Bun, namely, one of the daughter nuclei of each mitotic figure, lying with its long axis approximately perpendicular to the boundary of the oosphere, reenters the ooplasm (figs. 4, Ss). 2 Re result- ing oosphere contains many nuclei, the number usually varying from 50 to 100 in this species. These primary oospheric nuclei now divide mitotically, and their products function as the female sexual elements. It should be carefully noted that there are here, as in A. Bliti, two mitoses in the oogonium. The first occurs during the differentiation of the oosphere and provides the primary oospheric nuclei. The second occurs in the oosphere (figs. 6, 7) and results in the ont « a Tenn —s —— 1901 | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 81 female nuclei. These two divisions may be recognized at a glance, inasmuch as. the mitoses are simultaneous for all the nuclei concerned. The nuclear figure of the second mitosis dif- fers from that of the first in that the nuclear membrane disap- pears and there are fewer spindle fibers. The second mitosis usually affects only those nuclei that lie in the clearly differen- tiated oosphere. Peculiar activities are present in the center of the oogonium shortly before zonation, which lead to the formation of the coe- nocentrum. This structure, much less conspicuous and much more ephemeral in A. Portulacae than in A. Bit, is of the same general nature, however, and needs no special description. The central globule is found only rarely and is very small. The development of the antheridium of A. Portulacae is sim- ilar to that of A: Bliti in its general features. Simultaneously with the mitosis of the oogonium two mitotic divisions occur in the antheridium, but the difference which is so clearly apparent between the nuclear figures of the first and second mitoses in the oogonium is not found in the antheridium. The antheridial cytoplasm stains darkly with the gentian violet, and resembles the periplasm rather than the ooplasm of the oogonium. The antheridia lie closely appressed to the oogonia, and it is not unusual to find more than one adhering to the same oogonium, a condition much more rarely seen in A. Bhi. A slight papilla is developed from the oogonium into the antheridium at their point of contact. This is the ‘‘receptive papilla,” first described by Wager (1896) tor A. candida, and it reaches most remarkable proportions in A. Portulacae (figs. 1, 5; 6). In its early stages it resembles the receptive papilla of A. candida and A. Bliti, being merely a slight protuberance extend- ing from the oogonium to the antheridium.t. In A. Portulacae, however, it reaches much greater proportions ( jig. 5 ). Its development occurs at a later period than in the other species, usually after the differentiation of the oosphere or even during * The account of the structure in 4. Bliti (Stevens 1899) traces the growth of the papilla in detail. $2 BOTANICAL GAZETTE [ AUGUST the second mitosis, while in A. But it precedes the preliminary massing of the cytoplasm. In A. Portulacae the oosphere some- times protrudes into this papilla, a phenomenon not occurring in A. Bliti, since its oosphere has not been differentiated when the papilla is formed. The condition presented in fig. 6 shows that sometimes the papilla becomes ruptured. This I cannot regard as normal, however, since only one of the hundreds of prepara- tions examined shows such a condition. The mode of formation of the antheridial tube is uncertain. In somewhat advanced stages it is found extending nearly to the center of the oosphere. It has a well-defined wall, and is firm and straight (fig. 8), differing from the swollen gelatinous tube of A. Tragopogonis and A. candida, but agreeing closely with that of A. Blin. In no case was the antheridial tube in a mature oogonium seen to contain less than fifteen nuclei, and in most cases there are probably 150 or more. I have seen the tubes in transverse, longitudinal, and oblique sections, and they were always multi- nucleate and extended into multinucleate oospheres (figs. 7-9). The male nuclei are oval as they lie in the end of the antheridial tube, and stain more darkly at the anterior end. Whether there is any constant numerical relation between the sperm nuclei and oospheric nuclei could not be determined with any accuracy, owing to the great number of sperm nuclei crowded into the antheridial tube. When fully developed, the antheridial tube opens as in A. Butt. There is first a softening of the wall, which is probably due to the presence of an enzyme, and it is finally. dissolved, allowing the nuclei to escape and pair with those of the oosphere (figs. ro, zz). Before fusion both male and female nuclei enlarge somewhat, although they maintain a typical rest- ing condition during fusion and regain their original size soon after. The result is an oospore containing between 100 and 200 nuclei, which passes the winter without further change. A very few winter spores were seen with a relatively small number of nuclei, in one case as few as six, but this condition must be regarded as very exceptional. Such cases probably i ; ; ; Igor | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 83 result from a fertilization of oospheres containing few female nuclei, and such are rare. It is unnecessary to describe further the development of the spore, since its course is apparently parallel with that of 4. Biz. It consists simply of an accumulation of food stuffs and a growth of protective structures. A portion of a mature wall is shown in fig. 72. The endospore consists of one layer only, not two, as I have described in A. Blitz. Berlese, in a recent paper, entitled ‘‘Ueber die Befruchtung und Entwickelung der Oosphare bei den Peronosporeen,”’ dis- cusses some of the cytological phenomena of this genus. In | this study he has included observations on A. Portulacae, giving six figures to represent the species. His results differ essentially from mine, a difference which is the more interesting since simi- lar technique was employed, and in both cases the host plant was Portulaca oleracea. Yor the sake of clearness I will indicate some of the discrepancies between his work and my own. _ Ber- lese finds thirty to forty nuclei in an oogonium before mitosis, and says that these nuclei divide several times, the nucleoli van- ishing during prophase. My preparations show 300 to 400 nuclei which divide twice, their nucleoli persisting till late ana- phase. He finds ten to twelve nuclei in an antheridium, while my material shows a number several times greater. Notwith- standing his statement (p. 176), ‘‘das Cytoplasma, welches den in der Gonosphare zuriickgebliebenen Kern umgiebt, ist dicht und sehr fein gekérnt,” the ooplasm which he represents is much more coarsely vacuolate (Berlese, 1898, figs. 2-4) than in any Albugo I have examined. Indeed, it does not resemble the ooplasm of a typical Albugo, which varies but little, so far as my observation goes, but rather presents the features of the ooplasm of Peronospora. He notes no “ receptive papilla,” and the antheridial tube in both structure and content is very unlike that described in this paper. He describes the chromosomes as visible in the male nucleus during fusion (p. 179), and counts them. None of the species I have examined presents a possi- bility of counting chromosomes at this stage, inasmuch as the 84 BOTANICAL GAZETTE | AUGUST nuclei fuse in typical resting condition, and the chromosomes are then absolutely indistinguishable. Many other discrepancies might be enumerated. Such marked divergences, aside from the fact that Berlese has described a uninucleate oosphere and antheridial tube, while I find them both clearly multinucleate, are sufficient to warrant the suspicion that we are working with quite different species. Still, only one species is as yet described on Portulaca oleracea, and the configuration of the spore wall represented and described by Berlese agrees fully with that of the form dis- cussed in this paper. I have taken particular care to convince myself of the iden- tity of the species used, and have the assurance of Professor Magnus that my species is truly A/bugo Portulacae. Furthermore, I have been able to compare this material with authentic Euro- pean herbarium specimens, and they agree precisely. Not only do my preparations agree with European material in gross char- acters, but the cytoplasmic structures agree with those carefully figured by Istvanffi (1895, p/. 36, figs. 24, 25). This agreement emphasizes the identity of the American with the European spe- cies, and further shows that the species from widely separated regions? do not vary appreciably even in the cytoplasmic phe- nomena regarding which Berlese and I report such diametrically opposed observations. It is difficult to explain these numerous and serious discrep- ancies. Great variation in cytological detail is to be expected in this genus, and possibly the Italian form differs from the American as well as from that which Istvanffi studied. This is improbable, however, in view of the close agreement between these two latter forms. The problem can hardly find definite solution until the Italian form has been studied more closely. A mere difference in the number of functional nuclei, even a difference so great as that between the multinucleate and uninu- cleate oosphere, is readily conceivable in the same species when ? I have been unable to learn where Istvanffi collected his material, but assume that it is of European origin. } ; SANA EOE Ltt fees teen eBee om 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 85 the condition presented by A. 7ragopogonis is borne in mind. Berlese, however, describes no such phenomena as are seen in A. Tragopogonis, nor does he see the coenocentrum. Thus it appears that in the Italian form the reduction to a uninucleate condition, if such a condition really exists, is of an entirely dif- ferent nature from that described in this paper. ALBUGO TRAGOPOGONIS. In A. Tragopogonis there is a simple fertilization. One male nucleus is conveyed into the oosphere, where it fuses with one oospheric nucleus. To understand the variation between this form of fertilization, also characteristic of A. candida, and that shown by A. Bliti and A. Portulacae, it is necessary to follow care- fully the stages in the development of the oosphere in the two former species, and to homologize them with those presented by the two latter. In A. Tragopogonis the oogonium and oospore are smaller than in A. Bhtt or in A. Portulacae. The number of nuclei is less in proportion to the size of the oogoniun, however, so that for each nucleus there is more available space. This obviates much of the dense crowding of mitotic figures seen in A. BA and in A. Portulacae at the time of zonation, rendering this a favorable type for study. The small size of the oosphere, however, makes it much more difficult to recognize the stages of development, so that even the preliminary examination must be made under an immersion lens. The early stages are similar to those described for A. Portu- lacae. The protoplasm flows into the oogonium, the nuclei enlarge, the cytoplasm masses in the center, and finally the fully developed stage of zonation is reached. As in A. Bliti and A. Portulacae, this is a well-marked developmental stage (compare figs. 28 and 3), although it does not stand out with quite the clearness that obtains in those species. There is one central area of dense fine-meshed alveolar ooplasm, which is very clearly differentiated from the filar deeply staining periplasm. The central region, which at this time contains a very prominent 86 BOTANICAL GAZETTE [AUGUST coenocentrum, is entirely devoid of nuclei. They surround the central region, lying wholly within the periplasm, and are usually in metaphase. In this species, as in A. Portulacae, the mitosis is nearly completed in the periplasm, and the daughter nuclei do not begin to push back into the ooplasm until the chromosomes have retreated from the equatorial region of the spindle. It is also apparent that only those nuclear figures which are appropriately oriented contribute daughter nuclei to the ooplasm, and that only one of the pair gains entrance. From their mode of entrance it follows, as in A. Portulacae, that the chromatic content of the nuclei is situated at the end most distant from the sister nucleus. The nuclei number about fifty, and the tendency toward lowering the number that is occasion- ally seen in A. Portulacae is not apparent here. The primary oospheric nuclei now undergo a second mitosis, which is clearly distinguishable from the first by the character of the chromatic figure, which is similar to that described for A. Portulacae. The fact that the second mitosis involves only the oospheric, not the periplasmic nuclei also distinguishes it from the first (jigs. 32, 34). The definiteness with which ooplasm and periplasm are delimited at and after zonation (figs. 28-31) precludes any pos- sibility of confounding pre- and post-zonation stages. The clear- ness of the nuclei, which are seen as they enter the oosphere (fig. 29), and which can be followed through all the stages of the second mitosis (figs. 32, 34), renders it equally certain that in A. Tragopogonis the oosphere is multinucleate. So far there has been no deviation from the course followed by A. Bliti and A. Portulacae. These potential female nuclei appear to differ from each other in no important respect, unless it be in their distance from the coenocentrum, yet under the usual conditions only one of them is destined to function as a sexual nucleus. A study of later stages shows that one (or very rarely two or three) of these nuclei comes to lie in close contact with the coenocentrum, and there grows (figs. 35-37) until it becomes many times its former i : } A t { i ———# ee ee = er 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 87 size and much larger than the nuclei found at earlier stages in the developing oosphere. A study of oospheres after the second division shows the potential female nuclei in various stages of degeneration. They appear first to lose their chromatin network, the nucleoli and membrane persisting longest. Such degenerate nuclei can be found in abundance in the stages indicated. Frequently one or two rather large and apparently normal nuclei are found in an oosphere, which also has its one large nucleus in contact with the coenocentrum. In other cases, many very small nuclei are to be found in the ooplasm, and one large nucleus near the coe- nocentrum. These small nuclei range from the normal size to such small dimensions that only the nucleolus can be perceived with certainty. It would be impossible to recognize the small- est as nuclei, were it not that they are connected with undoubted nuclei by a series of complete gradations, and from the fact also that they lie in a cytoplasm remarkably clear and free from granules. Occasionally these extremely small nuclei are found in mitosis. Sometimes I have found groups of supernumerary nuclei, and often small ones are found in pairs, as though trying to fuse (fig. 38). Since this condition has never been found before the opening of the antheridial tube, it may represent the pairing of a supernumerary oospheric with a degenerate anther- idial nucleus..- That these inclusions described as degenerate nuclei are really nuclei is established by their structure and behavior. That they do not result from the division of a fusion nucleus is shown by their presence when the male and female nuclei are found lying together, but still unfused in the same oosphere (fig. 38). The degenerate nuclei, as would be expected, are most numerous immediately after the second mitosis, while the one functional nucleus which lies near the coenocentrum is yet small. Before the opening of the antheridial tube nearly all have disappeared. There is convincing evidence that the oosphere is at first multinucleate (figs. 30-32, 34) and eventu- ally uninucleate (jig. 36). There is no evidence to indicate 88 BOTANICAL GAZETTE [AUGUST that any nuclei escape through the boundary into the periplasm, but they are seen in the ooplasm in all stages of degeneration. The coenocentrum in A. Tragopogonis, as in A. Buti and A. Portulacae, makes its appearance when the central mass of ooplasm is formed just before zonation (figs. 27, 28). It first consists of an area of cytoplasm which takes the orange G with great avidity, while the neighboring cytoplasm stains deeply with the gentian violet. A trifle later than zonation the coeno- centrum is very highly developed, and appears in section as several zones of cytoplasm differing in density and stain reac- tion. The innermost area is coarsely vacuolate, and stains lightly with orange G. This region is surrounded by a narrower zone of dense granular cytoplasm, which is in turn encompassed by a less dense zone, and this finally by a broad zone of cyto- plasm which stains more deeply with gentian violet. This con- dition is not greatly changed at the time of entrance of the primary oospheric nuclei represented in fig. 30. In later stages the outer zones are lost, and the innermost region assumes a characteristic homogeneous oily appearance and is quite spherical (fig. 31). About the time of the second mitosis the innermost region, all that remains of the coenocentrum, loses its clear appearance and becomes coarsely granular (fig. 32), and in its stain reaction shows the probability of nuclear contents. The vacuolate area seen in the center of very young coenocentra (fig. 30) is probably a reservoir for the reception of foods that are elaborated by the- surrounding (presumably zymogenic) zone, which is in turn encompassed by typical trophoplasm. As the coenocentrum becomes older, these vacuoles, or rather globules of food stuffs in the protoplasm coalesce, and form the one central globule, which at first has a clear oily appear- ance (fig. 37), although it is not a true oil. The coenocentrum possesses an attraction for the nuclei similar to that noted by Wager (1900) in Peronospora parasitica. Nuclei in greatly elongated condition, apparently moving toward the coenocentrum, give sufficient evidence of this (jigs. 30,31). As a result, several nuclei come into actual contact = 1901 | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 89 with and even penetrate the coenocentrum (jigs. 32, 34), and are thus found during all stages of the second mitosis. In some cases only one nucleus in mitosis is found thus attached; in others as many as three seemed to be so anchored, and prob- ably, if the sphere could be viewed from all sides, as many as six or seven nuclei could be seen attached to the coenocentrum. The other primary oospheric nuclei are found lying free in the neighboring ooplasm in a similar stage of mitosis. It is not probable that any nuclei actually enter the coenocentrum, inas- much as it maintains its homogeneous appearance until consid- erably later. After the completion of the second mitosis, one small nucleus is found lying very close to the coenocentrum, possibly attached to it, although the evidence in A. Tragopogonis is not clear. Older stages show a larger nucleus. As the size of the nucleus increases, the coenocentrum becomes more granular ( figs. 35-37), and loses its definite form, eventually appearing simply as a granular mass partially enveloping the female nucleus. As has been said, there is usually only one nucleus lying beside the coenocentrum after the completion of the second division, although several are in contact with it during this process. The oosphere when ready for fertilization contains one large nucleus which lies beside the remains of the coeno- centrum. are employed in this paper, except that the Great plains region is made to include all east of the front ranges of the trans-Pecos Texas and north of the Rio Grande plain, with meridian 97.5° as an arbitrary boundary on the east. This is done because the physiographic ecology of the vegetation is most significant when it is borne in mind that the present condi- tions are but a stage in the leveling down of a former higher plain which covered this entire area, and whose remnants we have in the Great plains proper, the Edwards plateau, and buttes 5 Topographic atlas of the U.S. Texas folio. 116 BOTANICAL GAZETTE [AUGUST and mesas scattered all over the central prairie provinces. The process of leveling is in various stages in different places, such as deep dissections in the southern half of the Edwards plateau, undulating prairie in the Grand prairie province, a flat plain in the granite area with rugose surface caused by projecting granite ve oe aS RiTOF eel ae ag Fic. 6.— Vegetation provinces of the west Texas region, chiefly on the basis of pploniasts and geology: 7 to 74, Rio Grande plain, ae region (7, cee mber province; 2@ Edwards plateau plains, grass formations; 26 Grand prairie transition, between Lower Sonoran and Austro- -riparian, grass Ens lea. adobe vegetation, butte and escarpment timber; 2c, post oak formation on granite, Carbon- iferous, and upper cross timber sands and gravel; 2d, grass prairie formations of Red beds province, Lower Sonoran with elements of Upper Sonoran; 2¢, erosion remnants (buttes) of Staked plains and Cretaceous area, xerophytic timber; ey, Staked plains Lower Sonoran; 2g, Staked plains Upper Sonoran ; 24, Toya ah basin, f belts); 3 to 34, provinces of Rocky mountains and south plateau slope (7, extreme Lower Sonoran, bolsoa flora, chaparral, Yucca- -Agave-Cactus formations ; 74, UppeT Sonoran ; 34, Rocky mountain transition). 1901] VEGETATION OF WESTERN TEXAS i rg masses. These various phases in the process of wearing away the former plain and leveling up the denuded areas constitute the edaphic conditions which determine the general types of vegetation formation prevailing in the several provinces. These might serve almost equally well as designations of vegetation provinces. The provinces here referred to are the following: the Rio Grande plain; the Great plains region, embracing the Edwards plateau, the Grand prairie, the granite area, the Car- boniferous area, the Red beds prairies, the Staked plains, Toyah basin, and the Stockton plateau; the south plateau of the Rocky mountains, embracing isolated mountain masses and cafions, grass plains, and bolson deserts. PLANT FORMATIONS. The classification of plant formations emploved in the fol- lowing pages is based chiefly upon local conditions of soil, geo- logic structure, and physiographic features, that is the forma- tions are edaphic. For example, rock formations, forest forma- tions, and salt basin formations exist because of local soil structure or content. In the case of grass formations, climatic factors, especially moisture, play an important part, not only in determining the existence of a grass formation as opposed to a forest formation, but also in determining the special association of species in the different formations. For example, although the physical structure of the Staked plains is most favorable to forests, such formations are naturally excluded by scantiness of rainfall; and in the Rio Grande plain the pigmy forest of chap- arral succeeds the dense mesophytic forests of the Atlantic coast plain because the factor of moisture has suffered so great reduction. In every case the particular type of formation exist- ing upon a given local area depends upon the local condi- tions of physiography and geology. For example, of the forest formations the post oak type is always present upon sand and gravel beds. The streamway cafion, hill bluff, and escarpment forests are all products of the soil conditions prevailing where they occur; for, although their differences are due to differences 118 BOTANICAL GAZETTE [AUGUST of moisture, these are a result of the physiographic and geologic peculiarities, and may occur independently of rainfall and humidity. The plant life of the region in general may be included in the following formations: (1) grass formations, (2) woody for- mations, (3) succulent formations, (4) rock formations, (5) halophytic formations. I. GRASS FORMATIONS. The consideration of p'ant formations in western Texas may begin appropriately with the grass formations, for, excepting only the highest mountain summits of trans-Pecos Texas, the climate is a “‘ grass plains climate,” and the grasses may be said to form the matrix of the vegetation of the region. Texas is thought of commonly as a land of grasses, and properly so as regards the portion considered in this paper. Under what may be called natural conditions, to distinguish them from conditions which prevail under the present era of exploitation, the grass formations held their own in the perpetual struggle against woody vegetation. With the advent of the cattle business, however, this advantage was lost, and the present is an era of the rapid encroachment of timber formations. These phenom- ena and their causes will be-specially considered in a subsequent paragraph. Mention is made of the matter here to explain that in discussing the grass formations as they now exist we are dealing with a vegetation which, though still the dominant type, has not only a more restricted distribution than formerly, but is undergoing perceptible changes, not only in restriction of its area as the dominant formation, but in the association of species within the formation. Along with the grass plains vegetation will be discussed those types of formation which, though distinct enough as for- mations periodically, never for more than a brief period form the dominant vegetation, and are in every case distinctly a prairie feature. Such are the prairie annuals (generally meso- phytes) and the lignescent perennials (tropophytes). OO ae ET TT SS eet ETNIES ging wc 4 INE ahaa Igor] VEGETATION QF WESTERN TEXAS 11g In the present discussion the formations are taken up by geologic and physiographic provinces, because they seem not to be distinguished so much upon floristic as upon ecologic grounds. Temperature conditions will be seen to paly a réle as between a province in the extreme south and one in the extreme north, but even then chiefly in the floristic content of the annuals and lignescent perennials of the prairie formations. Again, the breadth of area from east to west, following the lines of decrease in rainfall, gives the grass formations of the eastern provinces a physiognomy differing from that of the extreme western. Finally, as previously mentioned, since edaphic fac- tors exert such a marked control upon formations, the consider- ation of the grass formations by provinces will give the full force of these factors. A preliminary word may be said in reference to the grass formation of the region asa whole. It was stated above that the climate of the region is a grass plains climate, and that the grasses form the matrix of the vegetation, this being true even where they are not the dominant element. Taking the region as a whole, there is a wide range of climatic (hydrometeoric) conditions between the east and west boundaries. But even in the province of greatest rainfall, climatic conditions, together with geologic and physiographic conditions, result in a decidedly xerophytic vegetation, The grass vegetation is the chief expo- nent of the xerophytic conditions, and certain ecologic types of grasses are found through the entire region, as Bulbilis dactyloides, the specifically designated ‘buffalo grass.” Throughout the entire region, also, the dryness of air and brilliancy of sunshine cause adaptations to rapid transitions from active growth to dormant conditions, great quantities of nutritive materials being stored in the dormant parts, THE RIO GRANDE PLAIN. PHYSIOGRAPHY AND GEOLOGy.—The area here included in the Rio Grande plain is really the continuation of the Atlantic coast plain west of the 98th meridian. It is in general a triangle, 120 BOTANICAL GAZETTE [AUGUST whose vertices lie at San Antonio, Del Rio, and Brownsville.. At the north, along the base of the triangle from San Antonio to Del Rio, the plain ends abruptly at the southern margin of the Great plains region, which is here marked by a sudden down- fall, the Balcones fault escarpment, which here has an altitude of about 1000 feet. From the foot of this escarpment the plain slopes gradually to Gulf level. The Rio Grande plain of geolo- gists, called also the Rio Grande embayment, is described as a constructional plain lying between this escarpment of the plains and the east front of the Mexican cordilleras. Its surface con- sists of the sheet flood débris of these two border regions, and its individuality, as distinct from the Atlantic coast plain east- ward, lies in its construction and its surface weathering under far drier conditions than those which prevail eastward. The sheet flood débris from the margin of the plains does not cover all of the Rio Grande plain as here defined. The flat coast prairie with its compact clay structure still extends along a narrow belt toward the lower Rio Grande. From Brownsville northward to the middle of the region extend vast sand plains,. tongues of which reach well up toward the escarpment border. The flood débris lies in coarser or finer beds over the northern. half of the plain, with exposures of arid clays, flat silt plains, or ridges of coarse gravel. Add to these features the basalt extru- sions both along the northern margin and in the southern sand plains, and the streamway erosions through the various deposits,. and we have the factors which not only combine to determine different types of grass formation, but have also figured promi- nently in favoring the encroachment of the woody vegeation,. chiefly the chaparral formations. CLIMATIC CONDITIONS.—The temperature conditions are of great significance to vegetation in this province, but only indi-- rectly do they react upon the character of the grass formations. This indirect control consists chiefly in permitting the occur- rence of woody species that require high annual temperature (Mimoseae for example), which, with certain artificial barriers- removed, the burning of the grass notably, are capable of. Igor] VEGETATION OF WESTERN TEXAS 121 waging a successful struggle against grass vegetation. A further result of temperature conditions upon the grass formation is to determine, in conjuncticn with moisture, the floristic content of the subordinate elements in these formations — the mixture of annuals and lignescent perennials with the grasses. With regard to moisture conditions in general, scant rainfall, low humidity, and brilliant sunlight, are such as to give the grass vegetation a pronounced xerophytic structure. In this respect the grass formations agree with those of the dry plains westward, in which, with the approach of the resting period, coincident commonly with rainless periods, the aerial parts become ‘“‘cured;’’ that is, they die, retain without loss their nutritive qualities, and remain in a good state of preservation. Since the’ 98th meridian is a mere arbitrary boundary, selected with reference to separating approximately the xero- phytic vegetation regions from the mesophytic, it is evident that at the eastern border of the Rio Grande plain, especially in the coast region, the grass formations approach more nearly the mesophytic structure; while at the west, along the Rio Grande, they approach the pronounced xerophytic aspect of distinctly arid regions. With respect to the relation of grass formations to woody formations in the Rio Grande plain, the encroachment of the lat- ter has been so vigorous as practically to destroy continuous areas of open grass formation. Much of the province is cov- ered by impenetrable thickets of chaparral. There are broad stretches of savanna where the grass formation is more open, but the areas are studded with isolated individuals or clumps of live oak, or by open post oak formation. On the basis of geologic structure and soils, three types of grass formation may be distinguished: (1) those of the flood débris plains; (2) those of the sand plains; and (3) those of the coast prairie. The first two are by far the most extensive and important. The third is but a slight extension of the coast prairie formation which is so characteristic on the Gulf coast from central Louisiana westward to our region. 122. BOTANICAL GAZETTE [ AUGUST THE FLOOD DEBRIS PLAIN.—The flood débris from the Cre- taceous formations of the Edwards plateau covers approximately the upper half of the Rio Grande plain, the altitude being from 500 to goo feet. The level stretches of this part of the plain are covered with finer silt débris, the mesquite-chaparral plains. There are arid clay hills in the Eagle pass region, and coarse gravel and stony slopes nearer the escarpment, besides basalt cones or ridges and the outlying block of rough hills (Anacacho mountains ). Floristically the grasses are chiefly of genera making up the buffalo grass range of the plains northward. On the rougher areas, especially westward, the extreme xerophytic conditions give the aspect of the arid plains. The associated species are chiefly lignes- cent perennials, or perennials with thick fleshy or tuberous roots, such as Jatropha spathulata sessiliflora on stony or gravelly soil, /. macrorliza on loose silty soil, and numerous other Euphorbiaceae and Nyctaginaceae peculiar to warmer lower Sonoran areas. The grass formation on these areas has been very much reduced by over-pasturage, so that during drouth periods vast tracts lie quite bare of grass vegetation. In this condition pas- tures not wholly beset with chaparral have the appearance of fields lying beaten and fallow. So great has been the depletion of grasses that during certain dry years it was stated that from fifteen to twenty-five acres of land were required to pasture a single cow. What permanent effects on the grass formation will. result upon areas so. denuded it is not yet possible to say, except that the chaparral will cover the entire plain. The grasses have great recuperative power, and it is said that after periods of abundant rainfall the earth is covered again with a close grass formation. No doubt a period of rest from excessive pasturage would enable them to recapture fully much lost ground. Of course, with the presence of the chaparral and the new relations it involves, the original grass formation of open sunny plains will suffer some material changes. This subject offers a field for ure investigations. ITH, JARED G.; Grazing problems of the southwest. Bulletin 16, Division of 5 eee U. S. Department of Agriculture. 1899 1901 | VEGETATION OF WESTERN TEXAS 123 THE SAND PLAINS.—These sandy grass plains constitute the most considerable feature of the southern half of the Rio Grande plain, and arms from them reach well toward the northern boundary. In some areas the sands are so deep and shifting as to render any stable vegetation impossible. Such areas are not yet well enough known to discuss here. The conditions as regards earth moisture in the sand plains are such as to leave them covered with grass vegetation and accompanying herba- ceous plants when the flood débris plains are bare of the corres- ponding formation. This is because they are not only better receiving areas, but the underground water is more available, for as the altitudes are low the distance to water is not great, and the open texture of the ground aids in that short distance in bringing water to the surface vegetation. The grass formations of the sand plains are different from those of the flood débris plains in two important respects ; first in the more open character of the formation, and second in their floristic content, in which the secondary elements —the prairie annuals and lignescent or succulent perennials—are especially involved. The open character of the grass formation permits an uncommonly varied growth of sand plains species, Many of the more important of these are of semitropical affinities, and hence not found in the more northerly or more elevated provinces. THE COAST PRAIRIE.—This is typically a sod prairie with grasses and sedges of mesophytic requirements, and annuals similarly adapted to wet, Jow, coast lands. Such prairie is con- tinuous between Houston and Corpus Christi, except for stream- way interruptions; though in passing westward along the line of decreasing rainfall the formation gradually becomes of xero- phytic stamp, and the semi-marsh land species are succeeded by grasses of the plains. Within the province of the Rio Grandc this formation passes into that of the sand plains. UNIVERSITY OF TEXAS. [ Zo be continued. | SKtaeG A STUDY OF THE SPORANGIA AND GAMETOPHYTES OF SELAGINELLA APUS AND SELAGINELLA RUPESTRIS. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY XXXI FLORENCE MAy LYON. (WITH PLATES v-IX) SELAGINELLA APUS. Sporancium.— It is by no means an easy matter to determine the origin of the sporangium in S. apus. It is quite possible, from a number of slides, to select a series which shal] seem to prove that the archesporium consists of two superficial cells originating just above the axil of a sporophyll (figs. 3, 4). It is equally easy to procure evidence that a single epidermal cell initiates the sporangium (figs. 7, 2, 5). This single cell may be either upon the sporophyll or removed by several intervening cells from its base. The exact line of demarcation between sporophyll and axis is indeterminate. The difficulty lies in the fact that until the sporangium is well established, consisting of some half dozen or more cells, there are almost no indications in the structure, or size, or staining qualities of these cells to distinguish them from the vegetative tissue. Moreover, a sporo- phyll originates in close proximity to a sporangium (fg. 5), and at about the same time and until it has established an apical cell there is no way of distinguishing it from a young sporangium. These facts render cross and tangential sections well nigh useless for an interpretation of the earliest stages. From serial radial, 7. ¢., vertical, sections of the strobilus, if cut with due reference to the phyllotaxy, it is possible to form con- clusions by comparison of the series of sporangia and sporo- phylls in the same rank and of different ages that appear in an exact median section. It is evident that the number of such 124 [AUGUST Pare RTT A a IRIE NT EEE IE 7 * 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 125 sections which may be obtained by ordinary means is very limited. I found it necessary to imbed the strobili singly, examine each section as its plane approached the median region, and constantly alter the angle of the paraffin block in the micro- tome, until by the vascular strand of the axis and the leaf traces of the older sporophylls I could estimate approximately the section sought. Even with these precautions, I was not infre- quently doubtful as to whether a cell that appeared in the right place theoretically for an archesporium might not be the initial cell of a sporophyll from the next rank that a slightly oblique section would display. These explanations are necessary, for after much painstaking study of many sections of tips of very young strobili, I find myself becoming less certain of there being a definite rule governing the initial phases of the sporangial growth. In many median longitudinal sections cut in the man- ner described above, a single epidermal cell projects from the surface in a vertical line between the youngest sporophyll and the apical cell of the strobilus. Occasionally it takes a deeper stain than the neighboring cells. There are usually from ten to twelve cells between it and the apical cell, and two or three between it and the subtending sporophyll (fg. 5). Quite as often this projecting papilla is composed of fwo cells of equal size lying in the vertical line (figs. 3, 4). In either case peri- clinal walls are formed, cutting off one or two cap cells (figs. 2, 5,6). From the hypodermal cell or cells, thus formed, origi- nates the sporogenous tissue. From the cap cell, which divides much more rapidly, the sporangium wall is formed. It is possible, I think, to determine in sporangia of various stages, nearly up to maturity, whether in a given case it originated from a single epidermal cell or from the two superimposed cells. In the second case, the complex of cells resulting from each of the two primary cells consists of more regular radial rows, and there is a quite definite plane of cleavage between the two groups. The only other interpretation of appearances like figs. 6 and 8 is that the primary single archesporial cell divided first anticlinally, thus producing the superimposed cells, each of which then cuts 126 BOTANICAL GAZETTE [AUGUST off a cap cell. I have sought in vain, however, to establish that the first two are sister cells. When the sporangium consists of from four to six cells (figs. 6, 7, 8), the hypodermal cells assume a different appearance when stained, which distinction is main- tained thenceforth in the development, and establish their identity as primary sporogenous cells. There is no apparent regularity in the order or plane of their division, with the excep- tion above stated. The tapetal cells are differentiated very early in the history, and are the peripheral sporogenous cells which assume a more symmetrical shape and regular arrange- ment (fig. zo). A few of the cells in this layer, lying directly above the pedicel of the sporangium, appear to be derived from the vegetative cells in that region, and are not the lineal descend- ants of the archesporial cell. Simultaneously with the differentiation of the tapetum the sporangium wall divides into two layers by anticlinal partitions (fig. 10,. The outer layer soon surpasses the inner in the size of its cells (fig. rz). 128 BOTANICAL GAZETTE [AUGUST formed upon its inner surface. This is the exospore ( figs. 20-22). The spores increase in volume, but the exospore expands so much more rapidly than does the protoplasmic content, that the latter is left in the apical portion of the spore cavity asa | tenuous spherical vesicle of protoplasm filled with a limpid fluid (fig. 377). A single very small nucleus, in which with difficulty may be seen one or two nucleoli, lies in the region nearest the spore apex, _The rest of the spore cavity between _ the vesicle and the exospore is filled with a limpid fluid of the same nature as that which occupies the space between the tetrad and the mother cell membrane. Soon hair-like radiations appear traversing these fluid regions (figs. 42, 43), whose general direction is from the spore mother cell membrane toward the * “* center of the tetrad. A thick layer which stains very deeply | appears upon the inner face of the exospore (fig. 37). The i latter maintains its more rapid rate of growth and soon after- ward is widely separated from this layer, by a space filled plage with the liquid and the radial fibrillae (figs. 38,42). The sculpturing of the exospore begins almost immediately upon its ¢ inception. The spines are laid down by depositions of matter derived from the liquid between the teres ane and the — Pe cell membrane ( figs. 42-46). ~-#e ern As the megaspores increase rae size, “nef are fotsue i speek iy the liquid in which they float seeping in between them (figs. 35,» 4-<< 42, 43) and the exospore is ruptured between the spores in such »¢« ee manner that each presents the appearance of a tetrahedron with a 4 2 ~ . Ss % -h hemispherical base and three plane triangular faces. The tearing apart of the spores leaves a trefoil-shaped cleft extending from the apex along the three ridges between the triangular faces, and bounded by the flaring flaps of the torn exospore ( fig. 58). The sterile mother cells in part disappear by dissolving in) the slimy fluid in the sporangium cavity, but not until the exo- ~~~ vie spore is well developed. Some persist however throughout the 7 development of the gametophyte and perhaps may grow slightly; ie they never divide. ne FEMALE GAMETOPHYTE.—The spore is but a small fraction of 1901] SPORANGIA AND GAMETOPAYTES OF SELAGINELLA 129 its final volume when the sexual generation begins, thus over- lapping the asexual (fig. 44). The initial steps of the female ° ' gametophyte development are the rapid expansion, without cor- responding increase in thickness, of the protoplasmic vesicle, and the division of its nucleus. The nuclei divide by karyoki- nesis, and with each successive division become larger (/fgs. 45,46). The thick envelope surrounding the vesicle stretches, becoming proportionately thin as its surface increases, until it comes to lie against the inner surface of the exospore. At this stage it consists of two distinct layers, the endospore_and The gametophyte at this stage thus consists of (1) exospore, still growing, (2) mesospore; (3) endospore; (4) the proto- plasmic vesicle, consisting of a very thin and homogeneous layer of protoplasm applied to the inner surface of the endo- spore, in whose apical region are imbedded numerous large, ovate, flattened nuclei; and (5) a large central vacuole filled with a watery fluid in which are suspended many oil drops. Beyond the increase in the number of nuclei, which preserve about the same relative distance from one another, there is no further change until the spore membranes complete their growth. When the maximum size is reached, fibrillae arise in the protoplasmic vesicle at the apex, and radiate downward over its surface, blocking it off into irregular areas, each of which contains one or more nuclei (figs. 49, 52, 52). Simultaneously the protoplasm becomes invaded with masses of granular matter, and encroaches on the space occupied by the vacuole. Nuclear divisions take place radially, and so rapid is this process that frequently the spindle fibers of three and four generations of nuclei may be seen in a single section. The fibrillar radiations permeate the protoplasm, keeping pace with its increase in thickness. As the central vacuole diminishes in size, its contents change in appearance from an emulsion to a turbid fluid thickened with granules (fig. 53). The proto- plasmic fibrillae apparently are concerned with the distribution nt ~» 130 : BOTANICAL GAZETTE [ AUGUSI1 of this nutriment before cell walls are formed. They have the appearance of streams bearing granules. The final and penul- timate divisions of the nuclei are distinguished by the appearance of nuclear plates and spindles of quite different appearance, which result in the formation of definite cells with cell walls, each containing a single nucleus ( figs. 26-28). At this stage, each nucleus is so surrounded by a mass of proteid as to be completely obscured. The only cells not thus gorged are a limited number which lie in the upper layer nearest the apex, beneath the trefoil-shaped cleft in the exospore (fig. 54). By continual multiplication they form a prothallial cushion, which widens the breach and bulges through. These cells are much smaller than those filled with the food supply. There may be from three to five layers inthis cushion. There is no diaphragm ) between the region designated the prothallial cushion and the | mass of storage cells underlying it. ARCHEGONIA.—A few cells in the prothallial cushion soon become conspicuous by reason of their large nuclei. Each divides by two periclinal walls, forming a tier of three cells. The uppermost of these divides by two anticlinal walls at right angles to each other into four cells. These again, by periclinals, form.the four cover and the four neck cells. The middle one of the original tier does not divide and becomes directly the single neck canal cell. The lowermost divides into the egg and the ventral canal cell ( figs. 29-32). A suggestive irregularity sometimes occurs in the last mentioned division. The central cell, that is, the lowermost cell of the original tier, may divide in such fashion that the egg and its sister, the ventral canal cell, may lie side by side in the venter of the archegonium, instead of in the normal fashion of ventral canal cell above the egg (fig. 32). The cover cells project very little from the surface x of the prothallium. Thus the archegonia are imbedded in the a surrounding tissue, whose cells in immediate contact with the egg and the ventral canal cell become more or less modified in form. The neck canal cell pushes up like a wedge, spreads apart the four neck cells, and dissolves. The ventral canal cell 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 131 also disappears (with a possible exception noted hereafter), and the egg lies free in the venter. There is a large receptive spot on the oosphere, and its nucleus is not centrally placed ( fig. 33). I have never seen more than five archegonia in a single gametophyte. FERTILIZATION.—Not only are the archegonia formed in the unshed spores, but frequently, at least, fertilization and the early phases of the sporophyte development take place while the sporangium with its prothallus are still in the strobilus. The strobili cease to grow, fade, and may separate from the plant before fertilization, but the spores do not fall from the spo- rangium. It was not until I had collected from the soil several hundred spores which had been shed, with the expectation of finding fertilization stages, that I thought to examine the with- ered strobili. Almost without exception in these I found embryos, whereas im no case have I found any evidence of fer- tilization in the spores that are shed. FURTHER DEVELOPMENT OF THE MEGASPORANGIUM.— The glandular tapetum is very active until the megaspores have stored their maximum amount of nutriment for the growth of the embryo. At this stage they quite fill the sporangium cavity, which in consequence has assumed a four-lobed appearance. The tapetum then declines in importance. Its cells collapse and form a pavement-like layer. The outer layer of wall cells becomes greatly modified. In section it appears precisely like a similar section of the annulus of a leptosporangiate fern (jig. 52). Four areas of the larger thick-walled cells, corresponding to the protuberances caused by the spores lying within, are separated from one another by narrow strips of small cells with thin walls. The latter are the lines of dehiscence. The spo- rangium splits into two valves along these lines, but the halves do not separate so widely as to allow fertile spores to escape. Apparently they may open and close more than once. The Sporangium appears fresh and active, and its wall contains chlorophyll until after fertilization has occurred. With the decline of the tapetum the lower stratum of the wall becomes 132 BOTANICAL GAZETTE [AUGUST more vigorous, as does a group of cells that lies just above the pedicel, and which projects into the sporangial cavity (fig. 58). This cushion is in close relation to the vascular strand and prob- ably facilitates the supply of nutriment to the sporangium wall, until the embryos begin to form. Frequently I have found microspores (in which the sper- matozoids had formed) within the megasporangium at the period of fertilization, and it is possible that the microspores are hurled into a gaping megasporangium when ejected from the micro- sporangium.? This is rendered more probable by the frequent occurrence of microspores caught in the angles between spor- ophylls and stem. Moreover, if plants that have become some- what dry be profusely watered, the mature microsporangia open explosively and discharge spores. Mrcrospores.— The microspores of S. aus are much smaller than those of S. Kraussiana and S. Martensii. They early develop a pebbled, thick exospore, which causes much trouble in imbed- ding and sectioning (fig. 69). Moreover, a comparatively small number, in proportion to the immense output, mature. Curious aberrations in growth are constantly found. It was necessary to study mature gametophytes discharging spermatozoids and trace back the different stages to the mother cells. Frequent com- parative measurements finally afforded a clew to detecting abnor- malities in the early stages. Not more than five sixths of the potential mother cells divide into spores; the others rapidly disappear. The division is accomplished in a manner analogous to the division of the megaspore mother cell, which is very little larger. Two spore coats develop, a thick spiny exospore, and the delicate mem- branaceous endospore. The microspores are shaped like the tI have examined S. Martensii, S. Kraussiana, and S. denticulata, growing in the greenhouse, with reference to this point. The two former do not shed their strobili, and I have found loose megaspores containing sporophytes in the soil on the enches where the plants are growing. The last mentioned species, on the other hand, sheds its strobili in profusion. I find, however, that their spores are invariably sterile and aborted, and therefore am unable to form an opinion as to whether this shedding is the normal habit, or due to cultivation in an unnatural environment. eae 190t] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 133 megaspores, but, unlike the latter, separate from one another as soon as the exospore develops. As was the case with the female gametophyte, the sexual generation begins before the microspore has ceased growing. At the moment that the microspores separate from one another, each possesses a thick and a thin coat, a layer of protoplasm parietally placed, with one nucleus, and a central cavity filled with fluid (jig. 68). The nucleus increases in size and divides. This process often takes place in that part of the spore where one of the lateral ridges meets the hemispherical base (fig. 78). The proto- plasm increases in quantity and encroaches on the central vacu- ole. Granules of various sizes make their appearance in the cavity and in the surrounding protoplasm. These bodies stain pre- cisely like the nuclei and the more regular ones may be mistaken for them, as I frequently discovered in the early period of my own work (fig. 77). By precautions in decolorizing, the presence of nucleoli always distinguish the nuclei from the granular masses. These bodies are formed by the agglomera- tion of many smaller granules. One of the nuclei formed by the first division remains against the wall. It may grow larger and the protoplasm immediately surrounding it is some- what denser, but no wall separates it from the rest of the spore. This may be the vegetative prothallial cell (figs. 70, 75). The other nucleus passes into the center of the spore. The protoplasm that envelops it sends out radiating processes that incompletely divide the spore cavity into irregular cham- bers, each of which is filled with granular masses of various sizes and shapes (figs. 76, 80, 81, 83). These strands of proto- plasm are continuous with a thin layer in contact with the endospore. The central nucleus with its envelope of proto- plasm divides into two cells which usually separate from each other (jigs. 79-83). By repeated division of each of these cells a complex results which consists of two uniform masses of sperm mother cells. There is no law of sequence which the cells follow in dividing, although the final product consists of cells very regularly arranged (jigs. 95-97). The male 134 BOTANICAL GAZETTE [AUGUST gametophyte then is made up of one prothallial cell and a naked mass of sperm cells, which later come to float in the slime produced by the disorganization of the food granules. Some- times at this stage the large deeply stained prothallial cell may be seen flattened against the endospore (jig. 97). There are about 128 sperm cells. The exospore splits along the three ridges from the apex downward, and the endospore, dilated with fluid, protrudes through the gap (figs. 98, 99). The sperm cells separate from one another, and a single spermatozoid is organ- ized in each. These are spirally coiled like those of Osmunda— two complete turns and a part of a third—but I can demon- strate no cilia (figs. 33, r00). Neither in appearance nor in movement do they resemble any bryophyte spermatozoid with which I am familiar, or have seen figured. The movement of biciliate gametes is characteristically different from that of an ordinary fern. These spermatozoids progress with a screw-like motion. The latest stages of development occur in the strobili after they are shed. MicrosporanGium.—At maturity the microsporangium wall consists of two layers of cells, of which the outer is divided into regions of thick and thin-walled cells, which enable the spor- angium to open lengthwise into two symmetrical valves (jig. 62). The microspores are discharged with much force. If plants that have become somewhat dry are watered copiously and covered with a bell jar, the ripe microsporangia burst open and jerk back their valves, which instantly recoil and hurl the microspores. These may be seen, looking like red powder, lodged in the axils of leaves or upon any other part of the plant where they have chanced to fall. Of necessity the waste must be great. _If this is the method adopted by the plant to secure fertilization it may account in a measure for the extremely small number of sporophytes that are developed. S. apus grows most profusely in a neighboring locality, where almost daily observations have been made during the past year. Early in May 1900, when this work was undertaken, the strobili were well advanced. It was observed that if several eggs in one Ig01] SPORANG/A AND GAMETOPHYTES OF SELAGINELLA 135 strobilus are fertilized, the formation of new sporangia is checked. The sporophylls begin to decay, as finally does the axis imme- diately below the fruiting head. The production of new strobili went on until late in August, at which time all the strobili were shed, whether fertilized or not. In September vigorous vegeta- tive growth took place during the fall rains, and continued until checked by several days of very cold weather in December. In the latter part of the same month, upon removing the snow that covered the plants to a depth of six inches, they were found to be green and ready to grow at any favorable moment. Large sods were lifted, without disturbing the plants, and brought into the warmth of the greenhouse, whereupon they responded promptly to the blander conditions. After a week, upon gently disentangling the individuals in order not to detach the semi- decayed strobili which were partially covered by the soil, I found several young sporophytes which had thrust their cotyle- dons and roots through the crevices of the megasporangia. From this it seems safe to assume that an embryo may have two periods of growth separated by one of quiescence, quite com- parable to those of seed plants. SELAGINELLA RUPESTRIS. There could be hardly a more striking contrast in the exter- nal appearance of two closely related plants than exists between these two species of Selaginella that are natives of New England. S. rupestris is to be found in the most exposed situations, grow- ing on the granite rocks on the mountain sides, wherever there has been enough weathering to insure the deposition of a little soil in the hollows. For six months of the year they endure alternate drought and drenching, and the frequent and rapid changes of temperature which are characteristic of this region, with no more protection than is afforded by their own structural adaptations. The midsummer conditions are even more trying. The plants that were studied for this work grew on bare rocks upon which the sun beats nearly all day from June to September. The fruiting spikes are to be found in profusion at all times of 126 BOTANICAL GAZETTE [AUGUST the year, but, as will develop later, the spores in the main are sterile. Material was fixed first about June 15, 1900, and at intervals of two weeks thereafter until February 1901. Sods were lifted from the rocks, and planted in shallow boxes which were kept in a warm place exposed to the direct sunlight. The spores were shed freely all summer. I collected other plants from a similar xerophytic situation at Starved rock, Illinois, in August, and for comparison material was sent me in Northamp- ton, Mass., from Austin, Texas, in November. These three regions are 800, 900, and 1600 miles apart. The plants collected in Massachusetts and Illinois, from June until February, almost without exception produced megasporangia only. The Texas material collected in November, on the other hand, was almost purely microsporangiate.? Expecting to find prothallia and young sporophytes developing in the spores that had been shed, these were picked out from the loam from which the plants were growing and killed at frequent intervals during the summer, with the purpose of getting all important stages. In all cases these proved to be barren. Late in August—as was the case with S. apus—the young sporophytes were discovered protruding from a withered strobilus that was nearly covered by the dirt. Further search produced many well-developed embryos, but it was apparently too late in the season to secure early stages (fig. 126). During the first week in January, a sudden thaw was suc- ceeded by several days of rain. Plants were collected from the mountain and gradually brought into a greenhouse temperature. The old strobili became greener at their tips, and new vegetative shoots started from the lower part of the stems. It is doubtful whether the increase in the number of individual plants in a given locality is due to any considerable extent to sexual repro- duction. A hollow in the rock, or a crevice, becomes filled with a closely compacted colony that is the result in great measure of vegetative reproduction, where prostrate branches, under Some of the Texas material was submitted to Professor L. M. Underwood, of Columbia University, for identification. He informs me that it is not S. rupestris, but a closely allied unnamed spec a aon eT. Is ae 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA E27 favorable conditions of moisture, have rooted, and later have severed their connection with the parent plant. In examining many colonies, very few young plants that originate from spores are in evidence, although the soil may be thickly beset with the spores that have been shed. Bits of the old plants that have been torn off by the action of the wind or rain are frequently caught in crevices along the precipices, and it is from these that new clumps are most frequently started. In fig. 124 I have endeavored to convey an idea of the struc- tural adaptation of the strobilus to its austere environment. The closely overlapping sporophylls form four rows, in whose axils the sporangia have little space to develop. The growing apex is protected by at least twelve and frequently sixteen sporo- phylls which envelop it. The epidermis is two or three layers of cells thick on both surfaces of the sporophyll, except in a shallow groove running lengthwise along the middle of the ventral surface. In this groove are the comparatively large crowded stomata, which are protected by the next older over- lapping sporophyll of the same row. While the leaves are yet very small the apex ceases to grow and becomes transformed into a branched spine.* The formation of the horny epidermis proceeds from the apex toward the base. Between the ligule and the sporangium the tissue retains its meristematic nature. The vascular bundle occupies the central shaft of the strobilus, and is augmented from the apex downward by the leaf traces that join it from the sporophylls. Four large communicating air Spaces, spanned by trabeculae, surround the entire system. These communicate with the exterior by certain larger chambers into which the stomata open. There is little closely compacted tissue in either sporophyll or stem. All the cells except the epidermis and the vessels contain chlorophyll. This structure is in marked contrast to the delicate unprotected sporophylls of S. apus. A single superficial cell, which uniformly is so close to the base of the subtending sporophyll that it is impossible to deter- mine whether it belongs to the stem or the leaf, is the origin of 138 BOTANICAL GAZETTE [AUGUST the sporangium (fig. zor). The sporophyll immediately above it starts simultaneously from an exactly similar cell. A wall separates the archesporial cell into an epidermal cap cell anda hypodermal cell (fig. 102). The sporangium wall develops from the former, the sporogenous tissue from the latter. A few sporogenous cells near the base originate from the stem tissue (fig. 105). The tapetum and sporangium development are quite like those of S. apus. The sporophylls and stem apex, on the other hand, have no apical cell. The ligule appears later in the his- tory of the sporophyll than in S. apws, and the megaspore mother cell is somewhat larger. Certain cytological features serve to distinguish these cells from the sterile sporogenous cells (jig- 106). The division into spores presents some curious varia- tions. Sometimes the spore mother cell divides once only, form- ing two megaspores; again, after the first division, one of the daughter nuclei may divide, or occasionally both. In any case, so far as observed, but one or two normal megaspores are ever formed; if there are others, they are dwarfed in size and never grow. That the exospore is a membrane common to both spores is. evident from figs. zz9, 720, r2z. The forthation of the mesospore and endospore is not preceded by any such deeply staining layer as is represented in fig.42. The protoplasmic vesicle expands and overtakes the spore wall. This stage terminates the develop- ment of the megaspore (fig. 723). Inthe older sporangia, nearer the base of the strobilus in the specimens that I examined, no further development had occurred beyond increase in size and. thickness of the spore coats—the contents showed that the spores were abnormal. This sequence of events, although followed step by step: repeatedly, had been rejected as probably not normal, until the sporophytes were discovered. As the number and shape of the megaspores in these sporangia which contained embryos agree with the stages described, in spite of the gap that remains to be. bridged between the megaspore and mature gametophyte, itseems. reasonable that the observations can be relied upon as far as 1901 | SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 139 they go (figs. roS-r20). Avery close series of stages from start to finish is necessary to produce confidence in any interpretation of phases of a plant which displays so many irregularities, and evidently has so nearly lost its power of sexual reproduction. The development of the microspores has not yet been fol- lowed in detail. Observations on this subject, together with the growth of the female gametophyte and the embryology, will be given in a future paper. One interesting feature that may be noted now is the incomplete septum which projects into the cavity of the microsporangium (fig. 725). This is formed in part from sterilized sporogenous tissue and in part from the cells of the pedicel. Since the above was written, a package of fresh material has been sent me from Texas. All the plants have begun their spring growth and the winter strobili are growing at the apex. The first sporophylls of the season subtend megasporangia. I find not more than eight or ten in each strobilus. Above these are microsporangia to the number of twenty or thirty. In this material it has been my good fortune to observe the method by which the spermatozoids obtain their entrance to the megaspo- rangia. A perfectly fresh, vigorous strobilus was cut from the plant and stripped of its sporophylls, thus exposing the sporan- gia zm situ. A megasporangium, which was observed gaping open and in close proximity to a microsporangium that had discharged its microspores, was separated from the strobilus and examined under the microscope. Six microspores were caught on the sculptured surface of the megasporangium near the edge of the valves, which were slightly separated from each other. One microspore, evidently mature from the fact that the exospore was split and the endospore protruding, suddenly discharged a current of slime which at first proceeded directly away from the megasporangium. Soon, however, as if acted upon by some attractive influence from the gametophytes within, the stream turned abruptly and entered the opening between the megaspo- rangium valves. The spermatozoids were swimming with char- acteristic rotary motion in this stream. With difficulty, it was 140 BOTANICAL GAZETTE [ AUGUST possible to make out that they were similar in shape to those of S. apus, but much smaller. To each was attached a vesicle. An attempt to stain them with osmic acid on the slide failed to demonstrate any more details. AppENDuM.— Since the foregoing was forwarded for publica- tion it has been possible to verify the main facts and to explain certain apparently contradictory features with regard to S. rupes- tris. The spring and early summer of 1901 were extremely wet as compared with those seasons inigoo. S. apus growing in low pastures made few strobili and aborted spores in contrast to the profusion of the year before, whereas S. rupestris throve in the unusual supply of moisture and developed strobili, which in turn produced embryos prodigally. The impotency of the mega- spores gathered in midsummer of 1900 was obviously due to the lack of water; for at the expiration of the rainy season this year, the last of June, and before the plants had suffered from drouth, they were lifted with all the underlying humus and placed in an open situation in the Botanic Garden. Profuse watering was continued and thorough draining secured. A large percentage of the strobili ripened after fertilization had occurred, and instead of few embryos found with much effort as in the season of 1900, hundreds of young sporophytes have been secured. The sequence of events in a propitious season, based upon the observation of two years, seems in brief to be this: Strobili are formed on the new vegetative shoots of the plant in late summer and autumn. Only megasporangia develop that season, and in these the gametophytes reached the stage bear- ing archegonia. In the spring, these strobili resume their api- cal growth, and first microsporangia appear. Thus each strobi- lus has a basal zone of megasporangia approximately six months old, and above it a narrow region of microsporangia. The num- ber of microsporangia appears to be strictly limited. I have found eight to twelve as a rule. Thenceforth so long as the strobilus continues to grow during the remainder of that season megasporangia only are developed. The production of these is checked in case embryos form in the lowermost zone of BOTANICAL GAZETTE, XXXII PLATE V (4 F \ \ ¥ Fg QO ce. ee << Ogi (Bao eee le | € a, PLATE VI BOTANICAL GAZETTE, XXXII . a ae PO fey oy oe ee A 2S Ss a = HELIOTYPE PRINTING OO. SAX Lit {¢ \ KI} Be * w ON a Ser my Ce yr rae S| ‘a ord an |e il iP 4 ; Be as tA ec [a Ni ZU; 4 Sie GBH oe AN \\ % \\ Sar (ia aoro. Oy, Maes delahits LINN a o> Peecs ra ge eg 5 a aoe 4 som (ungp, AN c\ HELIOTYPE PRINTING CO. Igor] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA I41 megasporangia. Immediately upon the cessation of growth in the strobilus and during its ripening, a vegetative lateral bud on the axis immediately below the strobilus is stimulated into activity, develops horizontal branches, and_roots which grow down into the humus. When this new growth is thus made independent, the axis bearing the strobilus and the germinating embryos decays, and at the period when cotyledons and roots of the young sporophytes are thrust out of the sporangia, the strobilus is lying in contact with the ground, shaded and otherwise protected by the vigorous vegetative growth that is spread above them. The strobilus decays much more slowly than the leafy axis beneath. This accounts for my finding strobili which appear to be shed from the plants. In case a strobilus fails to produce embryos, either through failure in fertilization, or probably more often because the dry season overtakes the gametophyte at a critical stage of its growth, the strobili continue to grow inde- terminately throughout the season, and apparently perform the vegetative work that, under more auspicious conditions, is taken up by the new growth from the lateral bud. Moreover, the spores rendered sterile by adverse conditions are shed from the sporangia. Thus not infrequently one finds greatly elongated strobili with a basal zone of empty gaping sporangia, surmounted by two or three whorls of empty microsporangia, then a zone of Sprouting embryos, and an apical region of degenerating young stages of megasporangia. In contrast with this are the strobili which have developed under continuous favorable conditions, in which the éasal zone displays embryos. Whether or not the strobilus may carry on its growth a third year in case of two suc- cessive failures to produce embryos, remains to be demonstrated. It will be a matter of interest to discover what variation in eco- logical conditions causes the strobili of the allied Texas species to pass the winter in a microsporangiate condition, whereas S. rupestris in Massachusetts appears to bear during the same sea- son only megasporangia. THE UNIVERSITY OF CHICAGO. [ Zo be continued.]} BRIEFER ARTICLES. THE PROBABLE FUNCTION OF CALCIUM OXALATE CRYSTALS IN PLANTS. A RATHER comprehensive study of vegetable histology (medicinal plants), extending over a period of six years, has brought to my attention more specifically the great abundance and wide distribution of crystals of calcium oxalate, and has led to the formulation of a theory as to their probable function in the plant economy. There seems to be no logically deducible reason for assuming that the crystals serve as a protection against herbivorous animals, though such a theory was promulgated by Stahl and others and is now widely accepted by teachers of botany. Calcium oxalate occurs in four predominating form types. Of these the least common is the crystal sand (Krystallsand, Krystallmehl, Krystallpulver) which occurs in the roots of belladonna, in the stem parenchyma of Solanum dulcamara, species of Atropa, Datura, Physalis; bark of Sambucus, Cinchona, and in some other plants. The prismatic and aggregate forms are perhaps about equally common and are very widely distributed. The needle-shaped or acicular crystals are also very common, but predominate in monocotyledonous plants. Leaving out of consideration the still undecided question of the chemical formation of the crystals in the plant and the causes which lead to the production of one or the other form type, we shall refer briefly to their probable function. G. Kraus in 1891 expressed it as his opinion that calcium oxalate was a reserve product to be redissolved by the plant and again utilized. This applies, however, only to a part of the crystals deposited during the previous season. Calcium oxalate dissolves with difficulty. It is insoluble in water, alcohol, ether, acetic acid, saliva, and other animal secretions. Being insoluble in these substances it is also tasteless ; hence taste cannot enter as a factor to guard against destruction by herbivorous animals. Nor is this substance poisonous. Some of it would no doubt be decomposed by the secretions of the digestive tract (as free hydro- chloric acid in the stomach), but not enough to produce poisoning due to 142 [auGusT i ' i es —”S t90r] _- BRIEFER ARTICLES 143 the oxalic acid liberated. It is generally admitted that the oxalic acid in various plants, as Rheum, Rumex, etc., serves as a protection against animals, not because of its highly poisonous nature, but because of the extremely sour and astringent taste. The theory that calcium oxalate serves to keep away animals through mechanical interference is highly improbable for several rea- sons. If this were the case the crystals would be peripherally located, as in this position they would soonest produce the desired effect. The crystals actually occur about uniformly distributed through the tissues of the various plant organs, and are in many instances especially abundant in the interior, as in the spongy tissue of leaves, the pith of stems, and the heart wood of steins. This mechanical interference can have application to small animals only, such as snails, insect larvae, etc. The crystals could not possibly injure or repel large animals capable of destroying the entire plant rapidly. Based upon observation, the conclusion is reached that the prime function of calcium oxalate in plants is that of mechanical support; secondarily it plays the part of a reserve product as stated by Kraus. The following are the chief reasons in favor of the mechanical support t oe . Cells containing prismatic crystals are quite generally associated ia, bast fibers. These crystal-bearing fibers consist of rows of rec- tangular, thin-walled cells, each cell bearing, as a rule, a single crystal. The cells surround the bast fibers or bast bundles. They are very abundant and distinct in the bark of Salix, Quercus, and Populus, for example. They enclose completely the single enormous bast cells of quebracho, and occur in the majority of bast-bearing barks and stems, and are associated with the bast tissue of vascular bundles. Bast cells are essentially non-elastic ; the crystal-bearing cells and fibers enclosing the bast give elasticity. This is shown to a remarkable degree in the inner bark of Quillaja, which contains an enormous quantity of large, elongated prismatic crystals of calcium oxalate distributed through the bark parenchyma. 2. In other instances the crystal-bearing cells are not merely an aid to mechanical tissues, but serve as a substitute therefor, function- ally taking the place of sclerenchyma. For example, in the seed of quince there is found a sclerenchymatous tissue below the layer of mucilaginous epidermal cells. In the white garden bean this scleren- chymatous tissue is replaced by a layer of cells carrying large prismatic 144 BOTANICAL GAZETTE [AUGUST crystals so constructed and placed as best to resist vertical and diago- nal pressure, as of soil particles and the weight of the bean. In the case of acicular crystals, so prevalent in monocotyls, it is evident that they give elasticity as well as support against crushing pressure. This applies especially to the very long and comparatively thick needles of calcium oxalate found in squill, iris, and other mem- bers of the lily family. These crystals (raphides) are especially com- mon in the parenchyma of roots, rhizomes, tuberous roots, stems, and leaves. It is generally admitted that the cystoliths of Ficus leaves, etc., perform a purely mechanical function. In plant organs subjected principally to a radial pressure, as are thick roots, tubers, thick rhi- zomes, etc., not specially supplied with mechanical elements, the aggregate crystals predominate, ¢. g., Rheum and Rumex. In such organs as potatoes, corms of Colchicum, etc., the necessary mechanical support is given by the starch which fills the cells. A potato free from starch would be crushed by the soil in which it grows. Further evidence in favor of the mechanical support theory of calcium oxalate is to be deduced from the fact that in many instances the crystals are imbedded in a gelatinous or mucilaginous substance which equalizes the pressure exerted, acting as a bumper between cell- wall and crystal. In other instances the cell-sap takes the place of the mucilage. It is very frequently found that cells bearing calcium oxalate take the place of mechanical cells in leaves. The crystals are especially common in the cells bounding the air chamber of stomata which cer- tainly require some mechanical support. It is also highly probable that calcium oxalate is merely accident- ally present in some plants and plant organs, but in the majority of instances its presence points toward a function of mechanical sup- port as indicated.— ALBERT SCHNEIDER, Worthwestern University School of Pharmacy, Chicago. | : | | | 3 a H i ; i i CURRENT LITERATURE. BOOK REVIEWS. Perception and propagation of stimuli in roots. A NEW FIELD oft research in plant eee has recently been opened stems. In the first of these* the author shows that there exist starch grains in the cells of the root caps of many roots, and in several other sensitive regions, These grains invariably occupy the physically lower side of the cells in which the lie. Most of the experiments were conducted upon the roots of seedlings and upon the coleoptiles of grasses. At the close of an experiment the organs were killed, and studied by staining and sectioning. The movement of the grains occupies a comparatively short time, a half hour being sufficient for their rearrangement in the cells of the root cap in Pisum, when this has been turned on one side. The grains are embedded in the plasma, and as they fall away from that part of the protoplasm against which they normally lie (z. ¢., when the organ is upright), the protoplasm on that side of the cell hides altered so that it stains much more deeply. If the organ has been so placed that when the grains come to rest they still lie against a part of the normally lower wall, only that part of the protoplasm which has been freed from them becomes altered. Cells which have been in an upright position, but have lost their starch, owing to the influence of a plaster jacket, show the same proptoplasmic thickening as those which have been turned so that the grains fall by gravitation away from their normal position. Roots so treated are found to have lost their power of geotropic curvature. The author suggests that this may be due to the loss of the starch. Also, when these roots are freed from the plaster and allowed to renew their growth, they regain their normal sensitiveness. The return of sensitiveness is accompanied by the formation of new starch grains in the root cap. Also, if the region containing the grains be cut away, geotropic reaction ceases for a time, and its return is simultaneous with the formation of new grains in the regenerated tissue. Némec is of the opinion that the stimulus for geotropic curvature is the change of pressure of the starch grains upon the protoplasm. These varia- tions in pressure are of such small magnitude, however, that it is well-nigh *NéEMEC, B.: Ueber die bears sprit des Schwerkraftreizes bei den Pflanzen. Jahrb. f. wiss. Bot. 36 : 80-178. Igor] 145 146 BOTANICAL GAZETTE | AUGUST inconceivable that they should produce an effect great enough to be propa- gated through many cells and produce a reaction. It seems much more probable to us that the sensitiveness is due rather to a chemical than to a physical change of condition within the cells. This seems the more probable on account of the fact that when the grains leave their normal position, the protoplasm here becomes changed. Possibly this may be due to the removal, not of the grain, but of the leucoplast in connection with it. The chemical effect of the leucoplast upon sugars passing through it is very great (since it produces in them condensation to form starch), and it may well be that it also has an effect upon the surrounding protoplasm. There may be other, as yet invisible, substances within the cell, which cannot diffuse readily, whose specific gravity differs from that of the protoplasm, and these might affect the protoplasm unsymmetrically, thus setting up a stimulus which could be prop- agated to the curving region. The second paper? treats of the conduction of the stimulus from the sensitive regions to the region of the curvature. Traumatropic responses in roots were chosen as field for experimentation. If the tip of an Allium root is wounded by a cut or needle thrust, the protoplasm of the meristematic cells bordering upon the wound heaps up and becomes more dense on the side toward the wound. The nucleus also migrates toward the wound, often coming to lie against the wall on that side of the cell. After a very short time these cells regain their normal condition. But in the meantimea second and third layer of cells, at the side of and behind the wound, have responded in the same manner. Thus in roots killed 15 minutes after wounding, the response has been propagated through 1.25 ™™ of tissue, but those cells within 1™™ of the wound have already regained their original condition. This propagation takes place most rapidly in a longitudinal direction, but only towards the base of the root. It also occurs in both directions laterally. A careful investigation was made to determine whether this difference in the rate of propagation of the response might correspond to any difference in structure. In the cells taking part in the traumatropic reaction, longitudinal strands of protoplasm can, by proper staining, be made visible with compara- tively low magnification. Sometimes there is a single strand, sometimes Sev- eral; sometimes they lie near the middle of the cell, and sometimes, especially in vacuolated cells, along the lateral walls. The always in contact with the nucleus, often dividing and enclosing this body between several branches which reunite beyond it. A differential stain for the branches has not been discovered, but they take the ordinary stains pusich more deeply than the surrounding protoplasm. The best results were obtaine by staining strongly with fuchsin S. 2 NEMEC, B.: Die Reizleitung und die reizleitenden Strukturen bei den Pflanzen. 8vo. pp. iv-+ 154. pls. 3. figs. zo. Jena: Gustav Fischer. 1901. Cf. note Bor: Gaz. 3x: 133. 1901. Igor | CURRENT LITERATURE 147 By use of high magnification the protoplasmic strands in Allium root tips can be shown to be bundles of fibrillae. These fibrillae have a definite sheath and lie embedded in a special plasma. In other plants the fibrillae could not be so well made out, but enough was accomplished to convince the author that, in general, the longitudinal strands are fascicles of smaller author supposes that the fibrillae are in contact, or perhaps continuous, through these walls. Transverse fibrils were found in certain cells, but never in bundles. The question whether or not these fibrillae have any connection with the transmission of traumatropic and other stimuli is a difficult one to answer. The influence of a number of changes in external conditions was determined, first with regard to the propagation of the traumatropic reaction, and then with regard to the fibrils. It was found that the conditions which cause a degen- eration of the fibrillae diminish the rate of propagation or cause this phenome- non to cease altogether. By a sudden change in temperature the fibrils may of this power to the cells is always accompanied by the regeneration of the fibrillae. Another line of evidence is furnished by the study of certain roots of Allium which exhibited an apparently spontaneous nutation, uninfluenced by gravity. In these the starch-bearing cells of the root cap were perfectly normal in appearance and behavior, but in almost all of these roots the bundles of fibrillae were disorganized. Still other evidence btained from Vicia faba. In roots of this plant the fibrillae are found only in the large plerome cells. If the plerome is severed bya knife-thrust the geotropic reac- tion either occurs not at all, or bending takes place only so far up the root as the wound. Némec concludes that the bundles of fibrils are the path of con- duction for traumatropic, geotropic, and other stimuli. The fibrils are strands of protoplasm specialized for conduction. The two pieces of research here reviewed are accompanied by figures which are certainly convincing with regard to the facts. To us it appears that the conclusions of the second paper are much better supported by experi- ment than those of the first. The bundles surely bear a close relation to the process of conduction, whether this relation be causal or not. The author compares the fibrils to the nerve fibers of animals, but it seems to us that there is little similarity--BurTON EDWARD LIVINGSTON. 148 BOTANICAL GAZETTE [ AUGUST Two new books on plant physiology.’ THE dearth of text-books on plant physiology seems about at an end. We record with pleasure the almost simultaneous publication of two books on this subject, one by Professor D. T. MacDougal, of the New York Botanical Garden, the other by Professor W. F. Ganong, of Smith College. It is sig- nificant that both these books are American. he first named volume comprises a very comprehensive, though necessa- sarily brief, account of the phenomena of plant physiology, together with explicit directions for laboratory experimentation. Discussion of principles and laboratory directions are so interwoven that the temptation for the stu- dent to work mechanically, without other end in view than to finish the experiment, must here be reduced toa minimum. For this reason the book may be found more useful on the laboratory table than in the reading room. The mere reader will often be disappointed by the absence of measurements, etc,, since it is intended that these shall be obtained in the laboratory. On the other hand, it will be well-nigh impossible for one to peruse any section without gaining a fair knowledge of the methods by which the principles there treated are established. After an introductory chapter on the nature and relations of an organism (in which many statements are unavoidably made which the student cannot understand until he has gone further) the author devotes seven chapters to the presentation of the subject of irritability. In each of these a group of external conditions and the influence of these upon the plant is taken up. These chapters are headed as follows: Relations of plants to mechanical forces, Influence of chemicals upon plants, Relation of plants to water, Rela- tion of plants to gravitation, Relation of plants to temperature, Relation of plants to electricity and other forms of energy (although there are no “other forms” mentioned here), and Relations of plants to light. The chapter on the influence of chemicals will be found especially valuable. The next chapter (1x) deals with the composition of the body, and consists of avery brief treatment of the different groups of compounds found in the plant, fol- lowed by methods for their extraction, separation, and identification. This discussion is so brief that the reader may be led into error by generalizations ; and students will need to be cautioned that the qualifying phrases in the chapter are very important. Following this are five chapters on the processes going on within the liv- ing plant. These are entitled: Exchange and movements of fluids, Nutri- tive metabolism, Respiration, Fermentation and digestion, Growth, and 3MacDoueat, D, T.: Practical text-book of plant physiology. 8v0- PP- xiv + gs. 159. New York: Longmans, Green & Co., 1901. GANONG, W.F.: A laboratory course in plant physiology, especially as ecology. 8vo. pp. vi+ 147. figs. 35. New York: Henry Holt & Co., 1901. 352. a basis for | 1901] CURRENT LITERATURE 149 Reproduction. The treatment of enzymes, which have become so important a consideration within the last few years, is comparatively full, both in the chapter on the composition of the plant and in that on nutrition, and will be especially useful. ollowing the chapters on processes within the plant, the last chapter in the book gives a very excellent exposition of the subject of vegetative propa- gation throughout the plant kingdom. Sexual reproduction is not treated. At the end is a valuable appendix containing tables of physical constants, such as the expansion of air at different temperatures from 0° to 35°, the density of oxygen and of carbon dioxid, the absorption of these gases by water, etc. It will be a great aid, in experiments where calculation is neces- sary, to have these tables in a convenient form for reference. Regarding the style, we wish it might have been clearer in places; there will surely arise difficulties of interpretation now and then. ‘The only impor- tant point of theory wherein the author may find others in disagreement with him is that which expresses itself here and there in the idea of some purpose underlying the whole field of plant phenomena. For instance, it is stated that there exist “reactions to shock . . . . which the most thorough investiga- tion has failed to invest with a purpose. New relations of the plant may be discovered, however, which will interpret these reactions.” The book is fully illustrated throughout, including many figures of appa- ratus. References to the most important recent publications on the different topics are given as footnotes, and both treatment and references are surpris- ingly up to date, articles being cited which appeared only a few months ago. The index is complete, and includes footnotes as well as text. The arrange- ment of the subject-matter is thoroughly scientific, which fact, together with its general completeness and reliability, should give the book a broad and constant use in laboratories where the subject is taught. Professor Ganong’s new book does not attempt to present any general dis- cussion of the subject, but is designed, as its title denotes, for laboratory use only. It is divided into two parts, the first being on methods of study and necessary equipment, while the second comprises the true outline of the course. Part I is written largely for the teacher, and every teacher of labor- atory science (whether it be physiology or not) will do well to read it. The first two sections of this part are especially noteworthy for their fund of ped- agogical ideas, which, it seems to us, are almost all philosophically and practically sound. The third and fourth sections deal with laboratory, greenhouse, and materials for the course. Section 5 gives practical direc- tions for a great number of laboratory manipulations, some of which often are a source of considerable trouble to the inexperienced. This will surely prove a very valuable part of the book. Part II, the outline of the course itself, consists of two divisions. Division sae? Se = et 159° BOTANICAL GAZETTE [AUGUST I treats of the structure and properties of protoplasm, including its composi- tion, its relation to external conditions, and its power of organism building. Division II deals with the physiological operations of plants. Here are included nutrition, growth, reproduction, irritability, locomotion, and protec- tion. The last two are not treated, however, the statement being made that they are almost purely ecological in their nature. The titles of the exercises throughout this entire part are put in the form of questions. The directions which follow are designed to aid the student in answering the question, but do not answer it for him. Just enough discus- sion is brought in with the laboratory directions to arouse the student's inter- est in the experiment in hand, and to make him appreciate what are its essential points. Ample references to the literature are constantly given, even to original articles; the author believes that direct contact with the sources of information is of great pedagogical value. Good half-tone repro- ductions of photographs show the student just how.the more complicated forms of apparatus are to be set up. With their aid he should be able to bring his experiment to completion with a minimum amount of aid from the teacher. Following this part are several pages of addenda in which are noted numerous improvements upon apparatus described in the body of the book. The course as outlined by Professor Ganong will doubtless occupy more time than many can give to an elementary course in plant physiology. For such teachers the book will still be useful, since it is so well arranged that one can easily strike out a topic here and there without materially affecting the course as a whole. The style is clear, vivid, and scholarly throughout. We can think of no book yet published which might better “serve as a guide to the acquisition of a general physiological education.” BURTON EDWARD LIVINGSTON. MINOR NOTICES. THE Transactions of the American Microscopical Society 21: 1900, contains 275 pages devoted largely to zoological papers and matters of gen- eral interest. The following are of special interest to botanists: C. A. Koroip, The plankton of Echo river, Mammoth cave; HENRY B. WARD, Comparative study of methods in plankton measurements ; GeorGE C. WHIPPLE, Chlamydomonas and its effect on water supplies ; CHARLES E. Bessey, The modern conception of the structure and classification of dialo ic with a division of the tribes and a rearrangement of the North, American genera. Professor Bessey accepts Miiller’s view that the filamentous condi- tion is the primitive one, and that diatoms should be regarded as typically filamentous rather than as unicellular forms. They should then be classed between the Peridiniales on the one hand, and the Desmidiaceae and Zygne maceae on the other. The Zygnemaceaeare regarded as the most primitive lee eee ee eee eee 1901] CURRENT LITERATURE 151 of the Conjugatae, while the desmids and diatoms are believed to represent two similar and somewhat parallel genetic lines in which the filaments tend to break up rather early into independent cells. The larger part of the paper is occupied by a key to the tribes and genera of the American forms.— CHARLES J. CHAMBERLAIN. NOTES FOR STUDENTS: EXPERIMENTS by D. Neljubow* have shown that the peculiar horizontal nutation of stems of seedling peas (Pisum sativum) grown in darkness at high temperature is due to the presence of small amounts of illuminating gas in the air of the laboratory. It is especially the acetylene and ethylene com- ponents of illuminating gas which are active. SO;, and vapors of CS, xylol, and benzol are very injurious. This peculiar reaction to gas seems to have been unknown before.— C. R. B. A FEw interesting points epsceaias the behavior of apples in cold storage are brought out in a bulletin’ by L. C. Corbett on this subject. ith the exception of York Imperial, the apples of all varieties tested lost less in weight when stored in the light than when kept in darkness. The acid content of was decreased in some varieties and increased in others. No explanation for these differences of behavior is attempted.— Ernst A. BESSEY. A NUMBER of experiments throwing light on the method of infection in pear blight are described in a bulletin® from the Delaware Experiment Sta- tion. The germs were introduced by means of needle punctures into termi- nal shoots, one year old wood, three year old wood, buds, leaves, and fruit. Leaves and young twigs were smeared with cultures of the germ and kept moist fora time. The latter, however did not cause the disease to develop, for it appeared only where the germs had actually been introduced into the tissue, as for example, where needle punctures had been made into the pres- ent year’s terminal shoots, into buds, leaves, and into the fruit. The pointed out by Waite, by transference of the germs by honey-seeking insects from nectary to nectary, and into the fruits, young succulent shoots, and leaves by the introduction through bites and punctures of insects of germs clinging to their mouth parts. The same bulletin reports the occurrence in Delaware of pear canker, which seems to be caused by Sphaeropsis malorum. 4 Beihefte z. Bot. Cent. 10: 128-138. figs. 2. 1901. _5CorsBett, L. C.: Cold storage. Bull. West Virginia Agr. Expt. Sta.) 74: 51-80. figs. 3. March 1901. Morgantown. 6CuEsTeER, F. D.: Pear blight and pear canker. Bull. Delaware Agr. Expt. Sta. 52: 1-8. figs. 1-7. April 1901. Newark. L52 BOTANICAL GAZETTE [AUGUST Spraying with Bordeaux mixture is recommended as a preventive measure for this disease.— ERNsT A. BESSEY. Dr. BRADLEY M. Davis has recently published’ some interesting studies. on the nucleus of Pellia. These studies were undertaken with the object of extending our knowledge of the cytology of the Hepaticae, and with the hope of throwing some light on the relationships of the various morphological manifestations of kinoplasm. To accomplish this three phases in the life history of the plant were examined, namely ee the germination of the spore, and the vegetative activities in the seta. In the spore mother cell the spindles are organized in the same manner as that characteristic of the asters with centrospheres were found during the prophase. These appear to- be transitory structures, however, as they disappear before the daughter nuclei are formed. In the vegetative cells the spindles are formed in essen-- tially the same fashion as that described by Hof and Nemec for the vegeta- tive cells of many spermatophytes. The kinoplasm here forms two caps. fitting closely over the ends of the nucleus which has become elongated, and these caps are changed into the poles of the spindle. The writer concludes, however, that the kinoplasmic fibrillae, the centrospheres, and kinoplasmic caps are all secondary developments from the primal granular protoplasm which is the only form of kinoplasm in any sense permanent in the cell. The paper forms an extremely interesting contribution and contains much that is of importance on the cytology of the Hepaticae.—A. A. LAWSON. CENTROSOMES in flowering plants are described by Bernard® in a series- of short papers. Froma study of Lilium candidum, L. Martagon and Helosis Guayanensts the writer convinced himself of the presence of the much dis-- cussed structures. Material was fixed in alcohol and in Flemming’s solution and was stained in a mixture of fuchsin and iodine green (1 per cent. aqueous- solution of fuchsin, 2 parts; 1 per cent. aqueous solution of iodin green, 2 parts and water 40 parts). The safranin gentian-violet orange combination: did not give as good results. _ In L. candidum the centrosomes were found quite regularly during vari- ous phases in the germination of the megaspore. They resemble the structures- described by Guignard, but are not so sharply defined. The centrosome was: also identified in the gametophytes of Helosis. In Z. Martagon centrosomes were found in the female gametophyte, in the vegetative cells of the ovule,. but could not be positively identified in the endosperm. The centrosome | is- cytoplasmic in origin. 7 Nuclear studies on Pellia. Ann. Bot. 15: 147-180. pls. ro-r7. IgOT. ® BERNARD, C. H.: Recherches sur les sphéres attractives chez Lilium ene: Helosis Guayanensis, etc. Jour. de Botanique 14: 118-124, 177-188, 206-212. pis. ¢-5- 1900 4901] CURRENT LITERATURE 153 Incidentally, it is noted that there are sometimes two embryo sacs in LL. candidum. In these species a very large vacuole develops between the two polar nuclei preventing the nuclei from fusing. The writer suggests that this may account for the sterility of this species. It is also noted that the ‘upper polar nucleus and the nuclei of the egg and synergids are erythrophil- ous, while the four nuclei at the antipodal end of the sac are cyanophilous. This difference in chromatophily is attributed to chemical differences due to ‘sexuality, the nuclei at the antipodal end of the sac having lost all sexual character.— CHARLES J. CHAMBERLAIN. ITEMS OF TAXONOMIC INTEREST are as follows: H. J. BANKER (Bull. Torr. Bot. Club 28: 199-222. 1901) has published a synopsis of the species of Hydnum, 40 being recognized, and one described as new.—AVEN NELSON (idem 223-235), in his 13th paper entitled ‘‘ New plants from Wyoming,” has described 18 new species and varieties belonging to various families.— DaviD GRIFFITHS (ddem 236-241) has described a new ergot growing on species of Hilaria in Arizona~—-MAXWELL T. MAsTERs (Bot. Jahrb. Beibl. a number na new species ae Seen vere Bek New G inea.— In Torreya (1: 41-43. 1901) N. L. Brirron has described a new Hieracium from Florida, E. L. GREENE a new Arnica from regon, J. BEAL a new d ee ae from Colorado.— B. SHIMEK (Bull. Lab. Nat. Hist. lowa Univ. : 139-170. Igor) has published a list of the pteridophytes of Iowa, with sta- Rae MAKINO and K. SuH1paTa (Bot. Mag. Tokyo 15: 1-14. f/. 1. 1901) have described a new genus (Sasa) of Bambuseae, including eight species which have heretofore been referred to Bambusa. The name is a common Japanese name for “small bamboos.”—Among some recently published notes on algae (Zoe 5: 121-129. 1901) W. A. SETCHELL has described new genera as follows: Hedophyllum ss Pleurophycus Laminariaceae), and Weeksia (Dumontiaceae).— B. L. RoBINsoN (Proc. Am. Acad. 36: 455-488. 1901), in a recent ae from the Gray dota has published as follows: A synopsis of the genus Melampodium, 36 species being recognized, 7 of which are new; a synopsis of the genus Nocca (Lagascea), 14 species being recognized, 3 of which are new ; new species and newly noted synonymy among the spermatophytes of Mexico and Central America, the genus Eupa- torium receiving 15 new species.— M. L. FERNALD (dem 491-506) has described a fascicle of new seed-plants from Mexico and Central America. Among them is a new genus of fee (Hippomaneae), A/coceria by name.— WILLIAM TRELEASE (Rept. Bot. Gard. 12: 79-80. f4. sé se Igo!) has published a new palmetto ‘Sabet Uresana) from Sonora.— J. NEWS. PROFESSOR MAXIME CoRNU, of the Jardin des Plantes, Paris, died on April 3, in the fifty-ninth year of his age. Dr. DuncaN S. JOHNSON, associate in botany at Johns Hopkins Univer- sity, has been promoted to an associate professorship. Dr. ALEXANDER P, ANDERSON has been appointed assistant in botany in Columbia University to succeed Dr. M. A. Howe. Dr. Anderson will also carry on physiological studies at the New York Botanical Garden. Mr. GeorGE P. CLINTON, of the botanical department of the Univer- sity of Illinois, spent last year at Harvard University on leave of absence. His leave has been extended for another year. He is at work upon the Ustilagineae. Dr. MARSHALL A. Howe has been appointed assistant in the New York Botanical Garden and will devote himself especially to the study of the red algae. He is spending the summer in the eastern provinces of Nova Scotia, Cape Breton, and Newfoundland. FRoM Science we learn that Dr. H. W. Harkness, well known as a botanist of the Pacific coast, and especially for his recent studies on the Tuberaceae, died in San Francisco on May 10. His collections become the property of the California Academy of Sciences. Dr. F. L. Stevens, who has spent the past year in Europe as an hon- orary fellow of the University of Chicago, has been appointed instructor 1p biology at the College of Agriculture and Mechanic Arts, Raleigh, North Carolina. He will have entire charge of the newly-established department. Dr. L. M. UNDERWooD spent the months of June and July in botanical exploration in Porto Rico in connection with a party from the Department - Agriculture at Washington, consisting of O. F. Cook, special agent for trop!- cal agriculture, Guy N. Collins, plant photographer, and Robert F. Griggs, botanical collector. Mr. H. HASSELBRING, a graduate of Cornell University and later assist- ant in botany at the New York Agricultural Experiment Station, Geneva, has been appointed assistant in botany in the Agricultural Experiment Station of the University of Illinois. Mr. Hasselbring will devote his attention chiefly to plant pathology. : CHARLES F. Hotrasg, Ph.D., formerly an assistant in the botanical labo- ratory of the University of Illinois, has just returned from three years study 154 [aucusT 1901] NEWS 155 at the University of Bonn. He is now promoted to an instructorsbip in botany, and is given charge of vegetable physiology. While abroad he gave chief attention to plant physiology and cytology. Some researches will be published shortly. THE OPENING of the Minnesota Seaside Station at Point Renfrew, Van- couver island, took place June 21. Thirty students are in attendance. Even- Yendo, of the University of Tokio, Professor Conway MacMillan and Mr. Harold Lyon, of the University of Minnesota, and Professor Francis Ramaley, of the University of Colorado. The shore is exceedingly rich both in fauna and flora, and large collections are being made by some members of the party. IN THE department of botany of the University of California Dr. W. J. V.-Osterhout has been promoted to an assistant professorship. Professor W. L. Jepson is this summer exploring the western portion of the Colorado des- . ert about San Jacinto mountain, in company with Mr. H. M. Hall, assistant in the herbarium. They will also make an exploration in Humboldt an Del Norte counties. Mr. N. L. Gardner, assistant, is making further studies of the algae of the Puget sound region. Dr. W. A. Setchell is at Pacific Grove, California, investigating the ecology and zonal distribution of the marine algae BOTANISTS will be interested in the organization of the new Bureau of Plant Industry of United States Department of Agriculture. The old divi- sional lines have been eliminated and instead the work has been divided into various related groups, as follows: Vegetable pathological and physio- logical investigations; Botanical investigations and experiments: Pomologi- cal investigations; Grass and forage plant investigations; Experimental gardens and grounds ; Arlington experimental farm; Congressional seed dis- tribution ; Seed and plant introduction; and Tea culture experiments. The executive officers of the bureau are: Beverly T. Galloway, physiologist and pathologist, and chief of bureau. Albert F. Woods, pathologist and physiologist in charge of vegetable pathological and physiological investigations, and acting chief of the bureau in the absence of the chief. Frederick V. Coville, botanist, in charge of botanical investigations and experiments. - Lamson-Scribner, agrostologist, in charge of grass and forage plant investigations. Gustavus B. Brackett, pomologist, in charge of pomological investiga- tions, The pathological investigations are conducted by the following staff : Erwin F. Smith, pathologist, in charge of laboratory of plant pathology; Walter T. Swingle, physiologist, in charge of laboratory of plant physiology; 156 BOTANICAL GAZETTE [AUGUST Herbert J. Webber, physiologist, in charge of laboratory of plant breeding ; Newton B. Pierce, pathologist, in charge of Pacific coast laboratory; Her- mann von Schrenk, special agent, in charge of Mississippi valley laboratory; Peter H. Rolfs, pathologist, in charge of tropical laboratory; Merton B. Waite, assistant pathologist, diseases of orchard fruits; Mark Alfred Carle- ton, cerealist ; C. O. Townsend, pathologist ; George T. Moore, physiologist ; M. Duggar, physiologist; Rodney H. True, physiologist; William A. Orton, assistant pathologist; Joseph H. Chamberlain, expert in physiological chemistry; Thomas H. Kearney, assistant physiologist ; Cornelius F. Shear assistant pathologist ; Flora W. Patterson, mycologist. The botanical investigations and experiments are conducted by the fol- lowing sinkee , ook, botanist, in charge of tropical agriculture; A. J. Pieters, botanist, in eee of seed testing laboratory; V. K. Chestnut, botan- ist, in charge of investigations of poisonous weeds; Lyster F. Dewey, assis- tant botanist ; Carl S. Schofield, expert on cereals. he following have charge of the grass and forage plant investigations : A. S. Hitchcock, assistant agrostologist, in charge of field work ; David Grif- fiths, assistant agrostologist, in charge of field management ; Elmer D. Mer- rill, assistant agrostologist, in charge of collections; C. R. Ball, assistant agrostologist. The officers of the pomological investigations are: William A. Taylor, pomologist, in charge of field investigations; H. P. Gould, pomologist, in charge of fruit district investigations ; George C. Husmann, expert, in charge of grape investigations. The following branches are conducted under the direction of the chief of the bureau, but the officers directly in charge are given under each branch: Experimental Gardens and Grounds: L. C. Corbett, puesrres E. M. Byrnes, head gardener; George W. Oliver, expert plant propagat Congressional Seed Distribution: Robert J. Whittleton, «penned of weighing and mailing section ; James Morrison, superintendent of records. Seed and Plant jatroduction: Ernst A. Bessey, assistant in charge; David G, Fairchild, permanent agricultural explorer. Arlington Experimental Farm: L. C. Corbett, horticulturist, in charge. Tea Culture Experiments: Dr. Charles U. Shepard, in charge. 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Univ., now run x 2 e . Applied Microscopy < W id > : Laboratory Methods Ww ~ OL Vol. IV January, 19 No. If ° ° . LEADING SUBJECTS Aan Moses C. White. e . GAGE, Cornell University, ..-....---6--: 1109 Fire in the Veterinary ‘Catege at Cornell. ' iit Labor Phot z “Siero = pee eeeeee 1113 re) The N Botanical Garden. D. DOUGAL, ating da Ceule Se mt 1115 a. Pretiminary Study of Mycerox CLAR ENBECK, Wells College, - 5: +. +555 1119 - Micro-Chemical Analysi: ‘otassium. o E. M. CHAMOT, Cornell University, . ....... - 2 Easy Method of Mounting and Preserv' osquit “a VA RAVHMMLMG Bee sae s a a ace eas OOS 129 Current Botanical Literature. c CHAS, J. CHAMBERLAIN, University of Chicago, ... . « 1131 Cytology, Embryology and Microscopical Methods. oO . 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Terms Liberal Send for Catalogue WEBER WAREROOMS: 108 Fifth ge New Yor 66 Wa oy Aaa Chicag 1 Tremont Seeis, Boston Vol. XXXII SEPTEN g, 1901 vis No. 3h" % _ BOTANICAL GAZETTE | _ JOHN M, COULTER axp CHARLES R. BARNES alten | WITH OTHER MEMBERS OF THE BOTANICAL STAFF OF THE UNIVERSITY OF CHICAGO. ats tema : oe reateraeen ae _— _ eS - | ees see — ee eee 4 Wis oe a | gomarcanas | nadia * Bg paren ata | Sa (= fee an Soap made by PEARS, in Great Britain, is incom- parably the purest and best for the toilet and bath. All sorts ores : secured. 3 . sell it, alj Sorts of people use it. All rights otanical Gazette HA Montbly Foutnal Embracing all Departments of Botanical dacserter Subscription per year, $4.00 Single Numbers, 40 Cents Thes subscription price must be ~~ in advance. No numbers are sent wens Ae expiration the time paid for. No reduction is made to dealers or agent FOREIGN AGENTS: Great Britain — Wm. 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Under the Shelter of a Great Rock are the holders of Four Million Policies, in force in THE PRUDENTIAL Protecting their homes and home interests, a convincing proof of the popularity of this abe gg act Insurance Com ans 298 to liberality to policy-holders, absolute safety, prompt payment te economical managem Write for Information Dept. 25 Profit. sharing Policies, $100,000 to S15. ‘ YDEN, Presi The Prudential Insurance Company of America HonzOFFICE: NewN nt J, VOLUME XXXII NUMBER 3 DOLANIGAL. (ter LE SEPTEMBER, ro0r GAMETOGENESIS AND FERTILIZATION IN ALBUGO. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. XXIX. FRANK LINCOLN STEVENS. (WITH PLATES I-IV) [Continued from p. 98.] Il. PHYLOGENETIC. Tuis study of Albugo has established a series of forms showing advances in certain respects and a marked reduction in others. The species are closely related, and the difference between two consecutive members of the series is not great, although the extremes are widely divergent. There are four prominent structures or conditions connected with the series, namely, the coenocentrum, the receptive papilla, the mode of zonation, and the number of functional nuclei. Arranged in the order of these characters the species may be listed as A. Portu- lacae, A. Bliti, A. Tragopogonis, and A. candida. In this series the coenocentrum constantly increases in complexity of structure and perfection of function, the receptive papilla becomes less conspicuous, the number of functional nuclei decreases, and the mode of zonation varies in the last two species. These condi- tions are graphically represented in the accompanying figure which is self explanatory. We may consider the series as a possible expression of the phylogeny of the genus. Which conditions are likely to be primitive ? 157 158 BOTANICAL GAZETTE [SEPTEMBER The coenocentrum increases as the receptive papilla decreases; either one or the other must be retrograding. From a study of these two structures alone it might be difficult to determine the true history, but a consideration of the manner in which the A. Butiand A. Portulacae. A. Tragopogonts. A. candida. Many Many potential and Several poeres and functional, functional. one functional. NUMBER OF FUNCTIONAL NUCLEI DIMINISHING A. Bliti and A. Portulacae. A. Tragopogonis. A. candida. Large. Small. Smaller, RECEPTIVE PAPILLA DIMINISHING. A. Blitiand A. Portulacae. A. Tragopogonis. A, candida. as Larger, nutritiv Small, Large, nutritive. ren tie ngly pe sn COENOCENTRUM ADVANCING IN STRUCTURE AND FUNCTION. oosphere of A. Tragopogonis attains the uninucleate condition shows that this structure in its ontogeny passes through a stage identical with that reached by A. Portulacae and A, Buti in their full maturity, thus giving an excellent illustration of the biogenetic law. Both the antheridia and oospheres of A. 7ragopogonts are multinucleate in their earlier stages, and the development is Igor] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 159 strictly comparable to that of A. Portulacae and A. Bliti, stage for stage. Later the supernumerary oospheric nuclei are elimi- nated, leaving only one female pronucleus to function. It seems necessary, therefore, to regard the supernumerary oospheric nuclei in this species as potential gameto-nuclei, and to regard the uninucleate condition in A. Tragopogonis as due to the sup- pression of the many and the survival of only one gameto-nucleus. In the light of this behavior A. Portulacae must be regarded as the primitive form, and A. candida the most highly developed. In view of the great functional importance of the coenocentrum in A. candida and A. Tragopogonis, it is also rational to regard the coenocentrum as an advancing structure. The reverse is true, however, in reference to the receptive papilla, as there is no indication of its present utility, and everything indicates that it is a vestigial structure. The difference in the mode of zona- tion between A. Tragopogonis, where the protoplasm aggregates into preliminary centers which finally coalesce into one, and A. candida, where one central region is formed directly by a reces- sion of the protoplasm from the oogonial wall, may well be regarded as an outcome of the reduction of A. candida to a uninucleate condition of the oosphere, and may be taken as further evidence indicative of the highly developed rather than the primitive condition of A. candida. It is hardly conceiv- able that evolution could have proceeded in the reverse direc- tion, that is from the type of oogenesis in A. candida to that of A. Tragopogonis, A. Bliti, and A. Portulacae. The serial evi- dence from the coenocentrum, the receptive papilla, the mode of zonation, and the number of functional nuclei concurs in point- ing to the multinucleate conditions shown in A. Portulacae and A. litt as primitive, and the uninucleate oosphere of A. candida as derived. The conditions are almost precisely the same as those in the Fucaceae studied by Oltmanns (1889), with the exception that in Albuginaceae and Peronosporaceae a coenocentrum plays an important réle in the reduction of the number of functional nuclei to its ultimate expression. It is not difficult to imagine 160 BOTANICAL GAZETTE [SEPTEMBER that the process of evolution of one form from another may have been very rapid after the development of the nutritive region, the coenocentrum. In 4. Portulacae and A. Bltt the struggle between sister gameto-nuclei is possibly close, but the conditions are quite different in forms possessing a highly devel- oped trophoplasmic area in the center of the oosphere, for it becomes a matter of much importance which nucleus is the first to reach and avail itself of the nutrition. The determining factors may be the position of the nucleus, its orientation, or its sensitiveness to chemotactic influences. In any event, a nucleus which gains more nutrition will contribute its substances to a larger fusion nucleus, and consequently leave stronger descen- dants to the next generation. Moreover, this action of natural selection, aside from furnishing stronger descendants, will foster exactly those characters which enabled the parent nucleus to prevail in the mother oosphere. In short, a condition obtains here which can easily be conceived as one that would conduce to a rapid evolution from a multinucleate to a uninucleate oosphere through the action of natural selection. It is a manifestation of what Klebahn (1899-1900) has termed “‘ Streben nach Einkernig- keit der Sexualzellen,” under such conditions that it is possible to recognize the cause of the ‘ tendency.” A peculiarity, probably a consequence, of the uninucleate condition of the oospore of A. Tragopogonis and A. candida is the division of its fusion nucleus before the spore passes into the winter condition. A series of mitoses rapidly con- verts the spore from a uninucleate to a multinucleate struc- ture. The division here, as in many of the higher plants, may be regarded as the initial step in germination. In the species with multinucleate oospores no mitoses occur until the long resting period, normally in this case the winter, is passed. In such forms, presumably the primitive ones, the intrasporal evidence of germination appears when the spore ruptures its coat and manifests externally the new activities. It is clear that the inception of division in one case is homologous with that in the other, and in both it constitutes the commencement {901 | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 161 of germination. It matters not that in one instance it is before and in the other after the resting period. The divisions which in primitive multinucleate forms come after the resting period have in A. candida and A. Tragopogonis been transferred to a time before the winter rest. The condition is very similar to that presented in the spermatophytes, where intraseminal ger- mination immediately following fertilization is the rule, although many orchids retain their one-celled condition indefinitely. Although the division of the fusion nucleus is the first step in germination, the time of this mitosis is subject to variation in different types. The division is delayed in the forms of Albugo having multinucleate oospheres, in Peronospora parasitica (Wager 1900), in Saprolegnia and Achlya (Trow 1895 and 1899). The division occurs soon after fusion in species of Albugo having uninucleate oospheres, and in the Peronospo- raceae (Berlese 1898). It appears from the consideration of these species that it is not simply a disproportion between the volume of the cytoplasm and the number of nuclei that induces division, for this disproportion is as great in the case of Perono- spora parasitica, Achlya, and Saprolegnia, as in A. 77 ragopogonis. Wager (1900) has already indicated that in these species, in which the retardation of the fusion is most marked, the incep- tion of division is also most delayed. He says (pp. 275-276): Peronospora parasitica is at St the only member of the group with retarded nuclear fusion. . . . - this species it is delayed until the thick zygote membrane has been oe formed. The ripe oospore of P. parasitica is uninucleate. . .. . In all the other species of Peronospora which have been examined the ripe zygote is multinucleate. Similarly, Trow (1895, p. 648, and 1899, p. 175) finds that fusion is slow in Achlya and Saprolegnia, and likewise the division of the fusion nucleus does not occur until after the oospore has passed the resting period. This association of retardation in fusion with delay in division suggests that after the act of fusion is completed, so far as the microscope can give evidence, there may be further changes which the elements of the pronuclei must undergo before fusion is really completed. 162 BOTANICAL GAZETTE [SEPTEMBER The delay of division in species like Saprolegnia, P. parasitica, and Spirogyra (Chimelewsky 1888) may be due to a delay on the part of the nuclei in completely preparing for fusion, and to slowness in the act of fusion itself, an act which is not com- pleted when the mere fusion made visible by our present powers of magnification is effected. In this connection Wager (1899, p. 578) makes this suggestion: This difference in the behavior of the nucleus during the maturation of the oospore is probably connected with the mode of germination, DeBary has already pointed out that in Cystopus and some other species the oospore on germination produces at once a mass of zoospores. In Peronospora Valerianellae and others the oospore at once develops a germ-tube. It may be, therefore, that the uninucleate condition of the zygote indicates germina- tion by a germ-tube, and the multinucleate condition germination by the formation of zoospores. Apart from the relationships within the group which stand out with more or less clearness, the cytological phenomena emphasize strongly the affinity between the Albuginaceae, Peronosporaceae, and Saprolegniaceae. Peronospora parasitica, according to the research of Wager (1900), has a uninucleate oosphere, which is fertilized by the entrance of one antheridial nucleus. In oogenesis the nuclei divide simultaneously while passing to the periphery, as in Albugo, and then one returns to the ooplasm. A highly developed coenocentrum is present, and exerts an attractive influence upon the nuclei even as they lie in the periplasm. All of these conditions, together with the presence of the receptive papilla, clearly attest a relationship between the Albuginaceae and Peronosporaceae. The highly developed coenocentrum and the presence of a uninucleate oosphere indicate that the Peronosporaceae are the more highly developed group. Their ancestors may have been some forms possessing a uninucleate oosphere, similar to that of A. candida, from which one of the mitoses has later been suppressed as supet- fluous. Merely from the evidence of oogenesis, the Peronospo- raceae might be regarded as an offshoot from the Albuginaceae after the uninucleate condition had been attained; that is, they may represent a further development of the condition illustrated 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 163 in A. candida. But certain peculiarities of the asexual organs make such a view improbable. The Albuginaceae and Perono- sporaceae may both have attained the uninucleate condition of oosphere independently, each being derived from ancestors having multinucleate oospheres. The probability of the multi- nucleate condition being the primitive one discredits the validity of Fischer’s position (1892, pp. 223, 224) regarding the deriva- tion of the Peronosporaceae and Saprolegniaceae from the Chytridiaceae. Oogenesis in the Saprolegniaceae resembles that in the Albuginaceae in having a multiplication of nuclei through a mitosis, followed by a degeneration of the superfluous nuclei, thus presenting very closely the condition seen in A. Trago- pogons. Trow, Hartog, and Humphrey (1892) failed to report a coenocentrum, but Dangeard (1890) has described a structure in both Saprolegnia and Peronospora which may prove to be the same (cf Wager 1896, pp. 308 and 322). From my own studies, Pythium closely resembles Albugo in its oogenesis. There is a withdrawal of the protoplasm from the oogonial wall, differentiating a vacuolate periplasm, clearly homologous with that of Albugo. Cytological knowledge of the events of oogenesis in this genus will have great value in determining whether its relationship is closest to the Peronosporaceae or Albuginaceae. The present indications favor the Peronosporaceae, where it was placed by DeBary (1881). Two diametrically opposed processes of oogenesis occur in the single-egged (eineiig) Albuginaceae and the many-egged (vieleiig) Saprolegniaceae. In the first there is a massing of protoplasm in the center, forming the rudimentary oosphere; in the second a peripheral accumulation of protoplasm leaves the central region comparatively free. There is in Peronospora, however, a movement toward the periphery rather than away from it, resembling in this respect Saprolegniaceae. It is con- ceivable that Pythium may represent a transitional condition between oogonia with single eggs and those with several, which 164 BOTANICAL GAZETTE | SEPTEMBER would accord with the theory of DeBary that the Saprolegni- aceae evolved through the Peronosporaceae. The origin of the Peronosporaceae and Saprolegniaceae from lower forms is far from clear. Before the discovery of the multinucleate oosphere in 4. But, a relation to some type like Vaucheria seemed probable. In -Vaucheria many nuclei pass into the rudimentary oogonium, after which, as in A. candida, the superfluous nuclei retreat from the cytoplasmic region which is destined to become the oosphere. The essential difference is that in Vaucheria the formation of the wall at the base of the oogonium is delayed until the superfluous nuclei pass back into the parent branch. In the Peronosporaceae the wall is formed when the nuclei are still in the oogonium, thus prohibiting their retreat. The result must be either a multinucleate oosphere or a degeneration of superfluous nuclei. Oltmanns (1895, p. 414) has justly remarked that the condition in Vaucheria is very like that in Fucaceae, presenting only such differences as are neces- sitated by the presence of cell walls between the oogonium and the parent branch. The remark applies with equal force to the Albuginaceae, The probability that the ancestors of the Peronosporaceae and Saprolegniaceae had multinucleate oospheres removes these groups from Vaucheria-like forms. If there is any relationship the connection must have been at a time in the ancestry of Vaucheria before the abandonment of the multinucleate condi- tion of its oogonium. As Oltmanns (1895) says, the passage of many nuclei into the rudimentary oogonium in Vaucheria is most readily explicable on the assumption that it once produced many gametes instead of the one that is now habitually found, a view that must meet quite general acceptance, as may be seen from the recent paper of Blackman (1900) and the literature there cited. Mycologists generally acknowledge the close relationship between the Peronosporaceae, Saprolegniaceae, and Mucorineae. The presence of a multinucleate oosphere and antheridium, and " the fusion of two multinucleate masses of protoplasm, quite ES QSOS Igor] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 165 naturally suggest a possible derivation of the two former groups through the latter. However, as Schréter (1893) has remarked, * Durch das vollstandige Fehlen von Schwarmsporenbildung, das oft rein fadige Mycel, die fast nur an der Luft terminal gebildeten Sporen entfernen sich die M. immer weiter von den Algen;” and it is difficult to regard the Mucorineae ancestral to these other Phycomycetes. The manner in which the Mucorineae may have arisen from ancestral algae has been discussed by Davis (1900, p. 308), who has indicated the similarity between the coenogametes of Mucorineae and those of A. Bit. Not- withstanding this similarity it does not seem advisable to regard the Mucorineae as a line productive of such forms as the Per- onosporaceae and Saprolegniaceae for the reasons expressed above. The similarity of the vegetative body, however, is sufficient to indicate the possibility, even probability, that the Siphoneae, Mucorineae, Peronosporaceae, and Saprolegniaceae constitute three distinct lines of development from a common parent stock. Since transition stages in the evolution of these groups are not known, the phylogeny of the coenogamete is little more than a matter for speculation. Two modes of origin are con- ceivable; either the coenogamete arose from a gametangium producing numerous gametes through the failure of the gametes to separate and become completely individualized; or it may be regarded as a structure, originally multinucleate, which arose and attained sexual differentiation through a line of ancestry composed of multinucleate zoospores. According to the first view the coenogamete is ‘morpholog- ically a gametangium or physiologically a compound gamete. Each nucleus is the nucleus of a gamete. According to the latter view the coenogamete itself is homologous with a gamete, being a multinucleate gamete. Arguments in support of either view must be based upon such fragmentary evidence as is afforded by the present existing species (which may be made to stand for Stages in phylogeny), or the partial repetition of phylogeny through ontogeny. Unfortunately, very little evidence can be 166 BOTANICAL GAZETTE | SEPTEMBER expected from ontogeny, since the conditions prevailing during oogenesis are such that any vestigial attempt to individualize gametes from the nuclei would be at once obliterated. Naturally the individualization of gametes would without doubt become quickly eliminated from ontogeny after fertilization en masse became a fixed habit. Perhaps some one of the other eight species of Albugo may exhibit in the oogonium vestigial traces of older conditions and a more perfect individualization of gametes, as the conidia in many Peronosporineae develop zoospores which later merge their individuality into a common mass of protoplasm when the conidium germinates by means of atube. If the multinucleate sex cells were primitively multi- nucleate, if there has been an independent line of development starting with multinucleate zoospores and leading through multi- nucleate isogamous gametes to the coenogamete, it might be expected that some structures indicative of such evolution would now be found among living plants. But there are no stages of such a character known, nor is there any group among the algae or fungi which promises to supply them. Possibly the most sug- gestive group is the Monoblepharadineae, which has recently been investigated by Lagerheim (1899), who finds only one nucleus in the rudimentary oogonium, and as many nuclei in the antherid- ium as there are sperms to be formed. There is much evi- dence, however, to support the hypothesis that the coenogamete is homologous with many gametes, which have failed to separate. This hypothesis postulates an origin from uninucleate, swarming gametes, generally present in coenocytic algae, as Hydrodictyon (Artari 1890), Dasycladus (Berthold 1880), Acetabularia (DeBary and Strasburger 1877), Protosiphon (Klebs 1896), not Botrydium (Rostafinski and Woronin 1877) as is so often erroneously cited,’ thus showing the prevalence of this mode of sexuality among the algae that are usually looked to for the ancestry of the Phycomycetes. The elongation of the sperm nuclei as they lie in the antheridial tube may also be regarded as indicative of an ancestry in which each nucleus was the * For the most recent work on these species see Iwanoff, 1898. een pene paar Igor | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 167 nucleus of an individual gamete. The fact that Phycomycetes, in conidia and oospores, sometimes lose the ability to individ- ualize their units has an important bearing on the problem. It is well known that the contents of conidia and oospores ordinarily separate into as many regions as there are nuclei, and that these develop into zoospores, each unquestionably an individual. However, if these spores, by a change of external conditions, are induced to germinate by means of a tube, the zoospores do not emerge as individuals. Not merely is there a failure to resolve the protoplasm into individuals, but there is an actual surrender of individuality after it has been attained. Not only does this occur in ontogeny, but it is a very generally accepted belief that it likewise occurred in the phylogeny of this group. In the more primitive forms which germinate by zoospores individualization is not abandoned but is merely delayed, since in germination each fusion nucleus or each of its immediate descendants develops its own plasmoderma and begins independent life. In such form the phenomenon is comparable to delayed wall formation in endosperm, with the remarkable exception that in Albugo fertilization occurs during the period of delay. Pyronema (Harper 1900) may be regarded as an excellent illustration of a condition .in which the individuality of the sexual nuclei habitually finds expression only through their behavior in the act of fertilization. The manner in which the fusion nuclei wander away immediately after fertilization, and the absence of units in the cytoplasm, are strikingly similar to Albugo, as are also the mitoses of the nuclei during oogenesis, the marshaling of the nuclei into a hollow sphere, the participa- tion of the oogonium in the dissolution of the wall adjacent to the antheridium; even the trichogyne resembles in many respects the receptive papilla of Albugo. While certain Ascomycetes resemble the Florideae more than they do the Phycomycetes, the points of similarity men- tioned above between the Phycomycetes, Albugo, and Pyronema, are sufficient to justify careful scrutiny of all features before 168 BOTANICAL GAZETTE | SEPTEMBER abandoning the view held by DeBary (1884) concerning the relationship of these two latter groups. Zoosporangia have not yet been studied cytologically with the completeness they deserve, and it is unwise to deny the possibility of relationship ‘between the ascus and the sporangium until both cell and cilia formation in the Phycomycete zoosporangium is understood. As the conidia in phylogeny and ontogeny stand for zoospo- rangia, it is clear that they represent masses of potential zoo- spores, and are consequently synplasts in the sense in which that word was used by Hanstein (1880). The vegetative coeno- cyte that develops from the spore is also a synplast, which in phylogeny (and often in ontogeny) goes back to a time when each nucleus governed a definite region of cytoplasm. The multi- nucleate condition of this vegetative body results from a retarda- tion of cell division, similar to that in the embryo sacs, particularly well illustrated in the gymnosperms, and in the eggs of insects (Hertwig 1892). In the latter forms the delay is not maintained long in ontogeny. Cell division follows soon after nuclear division. In the coenocytic algae there is failure to individual- ize during a whole generation. Existing forms show that the vegetative body was coenocytic long before the gamete became so. In Albugo, the multinucleate condition of the female sex- ual cell may be regarded as the result of pushing the synplast habit one step further in ontogeny. The sexual cells were the last to give up their individuality, the vegetative cells the first. The synplast in the Phycomycetes differs from that of most spermatophytes in many ways, most strikingly, however, in the mobility of its potential units. While it is phylogenetically equivalent to many cells, a compound of potential cells, in the sense that Hertwig (1892) uses the expression, the phycomycete Ssynplast, both in sexual and asexual parts, must be regarded as a morphological unit. The potential units have lost their definite limitations and so their morphological value. Discus- sion regarding the nature of the vegetative coenocyte may be found in numerous texts, as Strasburger (1880, pp. 372; and 1893): Zimmermann (1896, p. 10), Haberlandt (1896, pp. 12-62), Igo1 | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 169 Verworn (1897, pp. 74-78), Pfeffer (1897, pp. 49-51). It is unnecessary to quote these authors or repeat familiar discussion here, further than to state that they agree in general with the statement of Pfeffer (1897, p. 51), ‘“‘Auch derartige Erwag- ungen zwingen dazu, zunachst den veilkernigen Protoplasten als eine morphologische und physiologische Einheit anzusehen.”’ There is but little in common between the coenogamete, which is a structure of high physiological efficiency, and such structures as have been described by Golenkin (1900) and Klebahn (1899) in Sphaeroplea, which, as Klebahn remarks, resemble cases of polyspermy. They seem to indicate a patho- logical rather than a normal condition. The strict maintenance of the individuality of the nuclei and their characteristic behavior in fertilization adds another strong argument to the evidence, which is becoming cumulatively great, that these structures are the bearers of hereditary characters. The apparent. ease with which one nucleus can usurp the cyto- plasm of many is an argument against the energid theory of Sachs. [ Zo be concluded.) BoTANICAL INSTITUTE, BONN. A STUDY OF THE SPORANGIA AND GAMETOPHYTES OF SELAGINELLA APUS AND SELAGINELLA RUPESTRIS. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. XXXII. FLORENCE May LYON. (WITH PLATES V—IX) [ Concluded from p. 141.) GENERAL DISCUSSION. CONSIDERING the phylogenetic importance of the group, the literature on the Selaginellaceae is surprisingly scanty. More especially is this true of that which treats of the development of spore and gametophyte. Some marked discrepancies in essen- tial features have appeared in these papers, however, and show how much detailed life histories of more species are needed before further inferences are drawn as regards the relation of this group to others. Only one comparatively full account of a single species has yet appeared, Bruchmann’s monograph on S. spinulosa, published in 1897, although he gives no account whatever of the development of the male gametophyte nor of the sporangia. He had much difficulty in obtaining normal spores,” and therefore was unable to follow the earliest stages of the prothallium in detail. In general, however, he agrees with Heinsen and Arnoldi that there is free cell formation within the spore, followed by periclinal and anticlinal cell walls, thus making a disk-shaped mass of cells in the apical portion of the *“Uber die Keimung der Sporen habe ich zu zwei verschiedenen Malen Ver- suche angestellt. Das erste Mal sate ich sie gleich nach ihrer Reife (im August) auf Torf aus, hielt sie bestandig feucht und schutzte sie im Winter gegen Frost. Die erste Keimung weniger Sporen dieser Aussaat bemerkte ich erst nach zwei Jahren, weitere keimten dann in dritten Jahre, aber der grosste Teil zeigte selbst nach solcher Zeit keine Keimung.” 170 [SEPTEMBER oes entmemesns meee tan 1901] SPORANGIA AND GAMETOPAYTES OF SELAGINELLA apt spore. The upper layers of cells are much smaller and irregu- larly polyhedral in shape. No diaphragm is formed. Three tufts of trichomes grow from the upper surface of the prothal- lium, rend the exospore into three flaps along the ridges at the apex, and push them up out of the way of the developing archegonia. Only a limited number of the latter develop, some five to ten as compared with Pfeffer’s report of thirty in S. Martensii. There are four rows of neck cells, with three cells in each row. Pfeffer reports but two cells in each of the four rows in S. Martensii; Campbell reports two in S. Kraussiana. Heinsen gives a list of eleven species: S. Martensii, S. lepidophylla, S Willdenoviana, S. denticulata, S. apus, S. erythropus, S. Helvetica, S. serpens, S. Douglasi, S. glauca, S. pilifera, and says that he agrees with Pfeffer on this point. Bruchmann finds that although sev- eral embryos may start to grow, but one comes to maturity. The first division of the oospore of S. spinulosa is transverse to the axis of the archegonium, and the cell nearer the neck becomes a suspensor. . The most notable fact, however, is that this embryo has no | : foot. ‘Ein Fuss in dem Sinne, sie ihn z. B. S. Martensii, S. Apo Kraussiana und andere Arten besitzen, fehlt.” Whereas, the ryo of S. apus; as 1 have found, has (no suspensor, After tupesives ete perme the cover cells of the archegonium close together, ov cfenozs the walls of the neck cells thicken and turn brown. On the other hand, Pfeffer maintains that the neck of the archego- nium of S. Martensii gapes widely during the entire embryonic development; and he represents the suspensor pushing up into the neck canal like a wedge. This latter method is the rule in most pteridophytes, and hence S. spinulosa together with S7 apus-and S. rupestris are exceptions. In the last twenty years, as far as I know, but four other con- tributions have been made to a knowledge of these phases of the history of the Selaginellas. In 1871, Pfeffer published his paper ‘Die Entwickelung des Keimes der Gattung Selaginella.”’ The extreme difficulty of interpreting the spore and prothallial Structures, even with the aid of all the modern technique, makes 172 BOTANICAL GAZETTE [SEPTEMBER the description given by this author seem an extraordinary per- formance, and none the less that apparent discrepancies with other species may disappear upon reexamination. His account of S. Martensii differs from the foregoing in the following par- ticulars: The megaspore when about one fourth its final size pos- sesses two coats, exine and intine. Against the inner surface of the latter lies a thick layer of protoplasm inclosing a transpar- ent fluid in which floats a large ‘‘nucleus.”” Near the apex “the protoplasm has a different appearance,” but he was unable to interpret it. Obviously, he misinterpreted the so-called nucleus, which is the protoplasmic vesicle, and in all probability the peculiar appearance near the apex was due to the presence of the real nucleus. A dome-shaped diaphragm separates the gametophyte into two regions. This he thinks arises at the first division of the spore into twocells. The one toward the apex of the spore becomes subdivided into a tissue three layers thick in the middle, one layer at the periphery. The continuous division of the lower cell fills the basal portion of the spore with larger cells. The hollow spherical portion of the gametophyte below the disk he describes as becoming filled with ‘ frei- gebildete Primordialzellen.” He could not demonstrate the presence of nuclei in these cells, but regards it as probable. According to his statement, the growth of the prothallium occurs at two periods. The disk-shaped mass of cells appears before the spores are shed, the tissue below the diaphragm, the “ sec- ondary endosperm,” afterward. Archegonia do not develop until the spores fall. In 1894 Heinsen reexamined the species studied by Pfeffer. He fell into the same error as regards the interpretation of the protoplasmic vesicle and nucleus in the young megaspore. He was unable to determine the origin of the megaspores, other than that a tetrad arose endogenously in the spore mother cell. He denied the existence of a primary and secondary endosperm, and of a diaphragm, and corrected Pfeffer’s misinterpretation of the food balls in the gametophyte cavity “as freely formed cells.” Infrequently he found the archegonia formed in unshed spores- t i 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 173 The diaphragm which Pfeffer thought was the wall of the first division of the spore Heinsen regards as the plane of separation between the small cells at the apex of the gametophyte and the larger ones below, and not a wall. He also refutes Pfeffer’s state- ment that ultimately the cell divisions completely occupy the basal region of the spore. Heinsen lays great stress on the supposed fact that the nuclei of the Selaginellaceae increase solely by direct division. He investigated material killed at all hours of the day and night, not only spores but vegetative tips, with special reference to establishing this point. He found a total absence of karyokinetic figures. As this statement, which if true would be most surprising, has not been refuted by later writers, it may be of interest to note a possible explanation of the error. In describing his methods of imbedding, Heinsen says that he killed his material by immersing some specimens for ten minutes in Flemming’s fluid, others three minutes in aI per cent. corrosive sublimate solution, still others ‘mit gleich gutem Erfolge wante ich ein zweimaliges schnelles Eintauchen in kochendes Wasser an.” It is doubtful whether any of the kill- ing fluids thus employed had opportunity to penetrate the sporangium wall in so brief a contact; in which case his mate- rial died a lingering death through the washing and dehydrating processes. Under such conditions naturally there would be no traces of karyokinesis. The statement that he had as good results from two quick dips in boiling water as from the Flem- ming fluid is otherwise inexplicable. It is somewhat difficult to determine whether Heinsen means his statements to be general with regard to all the species he enumerates or not. His sum- mary leads one to assume that the sequence of events there set down is true of the eleven species above named. There are cer- tainly inaccuracies in many details if applied to S. apus. Arnoldi in the Botanische Zeitung of 1896 followed in a brief article on “Die Entwickelung des weiblichen Vorkeimes bei den heterosporen Lycopodiaceen.”’ He investigated S. cuspidata elon- ata, and gives a résumé of Heinsen’s paper, He agrees with Heinsen on all points, thus reiterating the misinterpretation of | 174 BOTANICAL GAZETTE [SEPTEMBER the spore content. Campbell in his Mosses and Ferns, published in 1895, gives an account of his investigation of S. Krvaussiana. He too describes “the single large globular nucleus’’ (z.e¢., the proto- plasmic vesicle) of the megaspore. The diaphragm which exists in this species he explains as arising, not as Pfeffer thought from the first division of the spore, but from the thickening of the walls of the lowermost layer of cells of the disk at the spore apex. He finds “numerous small nuclei” scattered through the protoplasm of the spore cavity, and the protoplasmic layer thickens until it ‘completely fills the cavity of the spore.” I have examined 2 Kraussiana with reference to these points. The diaphragm is very evident and obviously formed in the manner described by Campbell. I can demonstrate no nuclei at any period in the spore cavity, nor any protoplasm. The protoplasm forms a layer next the wall, as in S. apus, and the vacuole within is full of food matter, at first a fluid, then an emulsion, and finally filled with granules and small balls of matter which stain like nuclei. The protoplasmic layer grows thicker, but never fully occupies the interior space as Campbell describes. The latest contribution is that of Fitting (1900), ‘‘ Bau und Entwickelungsgeschichte der Makrosporen von Isoétes und Selaginella und ihre Bedeutung fur die Kenntnis des Wachstums pflanzlicher Zellmembranen.” He corrects the error made by each of the above named writers in interpreting the parts of the megaspore. An attempt to discover the exact origin of the megaspores failed, due to the fact that the spore mother cell stained so deeply in all his preparations that the details of the evolution of the four spores could not be made out. He was unable to avoid shrinkage of cells. There follows a very detailed account of the development of the spore coats, in which he differs from Heinsen as regards their origin. He drew con- clusions largely from living material examined in a salt solution. As I have said before, the interpretation of the origin of the spore coats seems to me largely a matter of theory, and one who has examined the young spores of S. apus, while inclosed in the sporangium and surrounded by sterile mother cells, is 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 175 skeptical of the results obtained by this method of investigation. He fails to comment on Heinsen’s statement with regard to the method of cell division. Both Arnoldi’s and Fitting’s articles have been comparative studies of certain phases of the Selagi- nella life history with the same stages of Isoétes. Thus, until the year 1900 we have not had a correct interpretation even of the parts of the megaspore. MALE GAMETOPHYTE.— We are indebted to Millardet's memoir for our earliest knowledge of the male gametophyte. This work appeared in 1869, and almost no detail of importance has been added by later workers. In 1885 Belajeff repeated Millardet’s work on the same species and corroborated all essential details. There has been no other during the last sixteen years until the present account given in this paper of S. agus. How much the discrepancies between the two are due to differences in methods of technique, rather than to specific characteristics, remains to be demonstrated. Millardet and Belajeff both examined the microspores of several species of Selaginella (S. Avaussiana, stolonifera, and cuspidata) in living condition, then added various reagents to the microscope slide, focusing through the spore coats or crushing them by pressure of the cover glass, to deter- mine the phenomena taking place within. The material for the account of the male gametophyte of S. apus was killed, imbedded, sectioned, and stained without removing the spores from the strobilus. It will be seen that the main difference lies in the fact that in the earlier accounts there are eight cells described which constitute an antheridial wall; that later these cells dis- appear, and the sperm cell complex floats free in the cavity thus formed. Both authors state that they were unable to secure a cellulose reaction for the cell walls. On the other hand, I could demonstrate the presence of no such walls in S. apus. The first division of the spore results in two free cells. The first, accord- ing to Pfeffer’s and Millardet’s view, is the reduced vegetative part of the prothallium. The latter, which I have called the generative cell, divides at once into a complex of sperm cells. Is it possible that the protoplasmic films surrounding the vacuoles 176 BOTANICAL GAZETTE | SEPTEMBER containing food bodies were mistaken by these authors for cell walls? It is not quite certain that the species of Selaginella called Kraussiana in America is the identical one known by that name in Europe. The writer investigated the American form and found the sequence of events identical with those that obtain in S. apus. In attempting to repeat the methods employed by Millardet and Belajeff, it was impossible to distinguish through the spore wall the nuclei from the food granules, as both stained alike. Removing the exospore by shoving the cover glass to and fro was a tedious performance, whose results hardly repaid the effort. Moreover, it was impossible to manipulate the stains with enough precision to differentiate the nucleoli. As regards the spermatozoids, explicit statements are few. Belajeff figures those of S. cuspidata as biciliate bodies, somewhat ellip- tical in shape, pointed at the ends, with a slight spiral twist. Millardet gives substantially the same description, but no figures. In connection with his account he makes this reference to the work of M. Roze, who had published his observations in 1864: Ainsi qu’on le voit, mes observations sur la forme des anthérozoides du Selaginella Kraussiana sont loin de s’accorder avec celles de M. Roze sur es anthérozoides des S. Martensii et S. Galeottii. Toutefois, comme il a en affaire 4 des espéces differentes de celle que j'ai étudiée, je m’abstiens de toute discussion sur ce sujet. That there existed little information on the subject at this time (1869) is evidenced by the following statement : Les seules observations que je connaisse sur la germination des micro- spores du genre Selaginella sont celles de M. Hofmeister et de M. Roze. Ces deux auteurs se sont bornés A mentionner le fait de la production, dans la spore, de cellules contenant chacune un anthérozoide. C'est M. Hof- meister qui a constaté le premier l’existence, et la de ces animalcules; M. Roze a montré qu’ils sont biciliés. So far as I can determine, the authors who have described spermatozoids of named species of Selaginella that they them- selves have seen are as follows: 1862. Hofmeister. : . 1864. Roze first demonstrated two cilia in S. Martensii and S. Galeotti. | | | | 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 177 1869. Millardet thus describes the spermatozoids of S. Krausstana: Il m’a semblé qu’elle se colore en violet par le chlorure de zinc iodé; je n’ose toutefois l’affirmer. On voit en dedans d’elle un filament roulé en hélice, soit 4 droite, soit A gauche, de facon a faire environ deux tours com- plets; l’une de ses extrémités est occupée par le granule réfringent d’amidon que je viens de signaler et par suite extrémement visible; l'autre est a peine distincte. La premiére constitue la partie postérieure du corps de |'anthéro- zoide, elle décrit un cercle plus étroit que le reste de l’animalcule ; on voit d@habitude a cété d’elle quelques granulations a peine appréciables. Les cils semblent placés l’un a cété de l'autre; on les distingue difficilement du corps. . Ces détails serviront a l’intelligence des différentes formes des pdwatsotles, Dans leur état de développement complet, ils sont entiére- ment débarrassés de leur vésicule et de la membrane de leur cellule mére, et présentent a l’extrémité postérieure un corpuscule d’amidon; il est resté dans la vésicule ow on le eigpniben c’est 1a une seconde forme. Une troisiéme est constituée par les l i portent avec eux une vésicule; une quatri- éme, par ceux qui ne sont qu a dégagés de leur cellule mére. Dans la forme la plus dévéloppée, le corps est presque droit et décrit une Spirale 4 peine sensible. Sa longueur dans cet état est 0.018 ™™ environ. II S'amincit graduellement depuis l’extrémité postérieure jusqu’a la naissance des cils: son épaisseur maximum ne dépassé pas 0.0007 ™ envir avant il se bifurque et se termine ainsi par deux cils trés-tenus, fee fois aussi longs le corps. 1871. Pfeffer described the male gametes of S. Martensit, and S. caulescens. 1885. Belajeff examined S. cuspidata, S laetevirens, S. Mar- tensit, S. caulescens, S. stolonifera, S. Kraussiana, and S. Poulteri. Of these he gives figures only of the spermatozoids of S. stolonifera. Thus, in the thirty-nine years since the speramatozoids of the Selaginellaceae were first described, there have been only four contributions upon the subject in which the spermato- zoids of but eight species are described from direct observa- tion crea for by the writer.3 The difficulty of determining S. Mart. : Roze, 1864; Pfeffer, 1871; Belajeff, 1885. Zs uk Calis Roze, ¥ 3. S. Kraussiana; Millardet t, 1869 ; Sapien 1885. 4. S. caulescens: Pfe fer, aes Belajeff, 1885. cuspt 1885. — a: fi as 178 BOTANICAL GAZETTE | SEPTEMBER the structure of bodies so extremely small can hardly be over- estimated. With either a Bausch & Lomb +, oil immersion objective and 3 ocular, or a Zeiss oil immersion 2™" and ocular 3, I was able to make out the spermatozoids only by their characteristic rotary movements as they left the microspore of S. rupestris. Their spiral form and attached vesicles were facts determined rather by interpreting appearances by those I had definitely seen in apparently similar but larger bodies, than by actual observa- tions. The spermatozoids of S. apus are somewhat larger, and I feel that there is less likelihood of error in describing them. It is obvious that a more critical examination of many species is needed before much weight be placed upon the so-called aberrant forms of the Selaginella spermatozoids in tracing the phylogeny of the group. THE DEVELOPMENT OF THE SPORANGIA.—The two most recent and important papers on the development of the sporangium are Goebel’s (1880) and Bower’s (1894). I shall quote verbatim Bower’s summary of results from Selaginella: 1. The sporangium is eusporangiate, and arises from the tissue of the axis, above the subtending leaf; the position varies in different species. 2. The origin of the sporangium is similar to that of Lycopodium, and especially resembles ZL. zxundatum, to which species the mature sporangium also is similar in form. 3. Two primary archesporial cells are usually present in each radial sec- tion, and these are derived, as in ZL. inundatum, from segmentation of two distinct cell-rows; as seen in tangential section, the archesporium is refer- able to three or four such cell-rows. 4. The first periclinal divisions in these cell-rows do not always define the archesporium finally; subsequent periclinal divisions may result in addi- tion to the central mass, as has been proved for Equisetum ; but here the addition is less regular. 5. The tapetum results from tangential division of the outermost cells of the central mass; the greater part of it originates as described by Goebel _ The tapetum is thus a sterilized part of the potential sporogenous tissue; a further example of sterilization is seen in the megasporangium, where all the sporogenous cells are disorganized, excepting the one mother cell of the megaspores. | | | | 1901] SPOXANGIA AND GAMETOPHYTES OF SELAGINELLA 179 7. Abortive sporangia are to be found at the base of the strobilus as in many species of Lycopodium. With regard to the derivation of the sporogenous tissue, repeatedly radial sections show a distinct plane of segmentation separating the sporogenous tissue into two such definite regions that it is difficult to avoid the conviction that each complex is the progeny of two independent cells. The term ‘‘archesporium ” is used by Bower to signify the lower cell or cells. I have used the term to indicate the superficial cell itself. With regard to the fourth statement, my observations on S. apus do not agree. With the fifth and sixth statements, as regards the origin of the tapetum, my observations are in accord. The seventh statement I find true of S. rupestris, but not of S. apus. In no particular do my preparations agree with Campbell’s figures of the sporangial development of S. Kraussiana. Of no little interest is the incomplete septum in the micro- sporangium of S. rupestris, which arises above the pedicel. In vertical section, the resemblance to the celebrated fossil Lepzdo- dendron Braunti, as figured by Bower (see his plate 48, fig. 100), is very marked. He discusses in the accompanying text the possible function of this region of the sporangium. ‘ Possibly,” he says, ‘‘ these extensions of sterile tissue may have facilitated the nutrition of the developing spores, or they may perchance have contributed to the mechanical support of the sporangial wall.”” In connection with the former theory, it is noteworthy that this septum is found only in the microsporangium, where some of the spores are remote from the tapetum. A feature that may have some bearing on this point is the fact that the megaspore mother cell, so far as I have observed, is always near the periphery of the sporogenous mass and never at the center, a favorable position to secure nutrition from the tapetum. Scott in his Studies in fossil botany, 1900, says, & propos of this feature in Lepidostrobus Veltheimianus, ‘this structure may be compared with the trabeculae of Isoétes. It is best shown in the miicro- sporangium, but may have originally been present in the mega- sporangia also.” 180 BOTANICAL GAZETTE [ SEPTEMBER Of even greater interest is the simultaneous discovery of a paleozoic lycopod each of whose sporangia contains a single megaspore or embryo sac, which presumably was fertilized while still attached to the plant, with the condition that I have described above that obtains in S. rupestris and S. apus. Of the integument that grows up about the sporangium of the Cardto- carpon anomalum, described by Scott, leaving at the top an open slit-like micropyle, there is no trace I think, except that in S. rupestris, quite late in the development, after the embryo has formed, the megasporangium becomes sunken in a shallow pit formed by the. cushion-like upgrowth of the sporophyll around the pedicel. This outgrowth hedges in the ligule with the sporangium, and may be homologous with the Lepidostrobus integument. Scott’s statement, therefore, that ‘the recent dis- covery of Lepidostrobi with integumented, seed-like sporangia, in which only one megaspore came to perfection, shows that some paleozoic members of the group went far beyond any of the living representatives in the differentiation of their repro- ductive organs,” needs modification in view of the fact that S. rupestris normally at the present day produces seed-like sporangia with well developed embryos. SUMMARY. 1. In both S. apus and S. rupestris the sporangium frequently, if not always, may be traced to a single superficial cell, the archesporium. 2. The sporogenous tissue may arise in two ways. First (S. rupestris), from the single hypodermal cell formed by the archesporium being divided by a periclinal wall, thus producing a wall and a sporogenous cell. Second, by the archesporium (which in this case is assumed to consist of two independent superficial cells) dividing into four cells by a periclinal wall, the two hypodermal cells thus formed developing the sporogenous complex. In each case the epidermal cells form the sporangium wall. It is possible but not demonstrated that the second case may be a phase of the first wherein the original superficial archesporial cell divided by an anticlinal wall. Igo1] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 181 3. The tapetum is formed in part from the sporogenous cells near the exterior of the mass, and in part from adjacent vegeta- tive cells which come to form a more or less regular layer. It is defined very early, and is active and glandular up to the period that the spores have attained their full size, then becomes reduced to a thin epithelial-like layer lying against the spo- rangium wall. 4. Microsporangia and megasporangia are indistinguishable before the spore mother cells are differentiated. At this stage, in the case of the megasporangium, one or rarely two cells become more conspicuous and divide into spores. If micro- Spores are to be formed, the majority of the cells continue dividing. After the megaspore mother cell is differentiated, all other cells in the megasporangium cease dividing. 5. In S. apus four megaspores arranged tetrahedrally are formed within the spore mother cell. In S. rupestris four spores may develop in the same manner, only one or two of which may come to maturity; or, most commonly, there may be but a single division of the mother cell nucleus, in which case there are but two spores formed; or again, there may be a redivision of one only of the two daughter nuclei, resulting in three spores of which only one or two attain maturity. Isolated cases have been met with where but one megaspore was formed. As there were no signs of other aborted spores, presumably the mega- Spore mother cell never divided, but became directly the single megaspore. In no case have more than two spores in a single megasporangium been found which were fertile, and very fre- quently only one. 6. The megaspore of each species has three distinct coats, the €xospore, mesospore, and endospore. The former originates on the inner face of the spore mother cell membrane, and when first distinguishable is a film of unequal thickness. This either directly or indirectly gives rise to the exospore. A thick layer develops between the exospore and the protoplasmic vesicle, which later in its history separates into two layers, the meso- Spore and the endospore. 182 BOTANICAL GAZETTE [SEPTEMBER 7. The female gametophyte is formed by free cell division of the megaspore, the nuclei dividing by indirect division. These nuclei are confined to the apical portion of the spore. Several layers of nuclei are formed by repeated tangential and radial divisions so that there are six or seven in the apical region and one in the basal. Areas of cells are blocked off by proto- plasmic radiations passing from the apex outward and inward. The walls of the cells are produced by nuclear plates in the final division. There is no diaphragm, and at no stage of its develop- ment are there nuclei in the lower or the central portions of the gametophyte, which at first contains liquid, and finally a semi- solid mass of granular matter. 8. A cushion of cells protrudes through the tripartite cleft in the exospore at the apex. From cells in the upper row of this cushion a limited number of archegonia develop. The cells of this region are markedly smaller than those in the other regions of the gametophyte. No part of the archegonium pro- trudes from the general level except the cover cells. g. The megaspores and gametophytes are nourished by matter secreted by the tapetum and passed through the spore mother cell membrane, which persists until the spores are nearly half grown. 10. The microspores develop in a fashion analogous to the megaspore. A large percentage of the mother cells form tetrads, the largest proportion of which are aborted at this stage. 11. The male gametophyte of S. apus consists of a single prothallial cell and an ovoid naked mass of potential sperm cells arranged in two groups. There is no antheridium, nor is there a wall which separates the prothallial from the sperm cells. 12. The spermatozoid of S. apus is a spirally coiled body; that of S. rupestris is of similar shape but much smaller. The presence of cilia in either was not demonstrated. 13. The megasporangia and microsporangia of both species open by definite lines of dehiscence. 14. Fertilization in both species occurs while the spores are LS Sa Igor ] SPORANG/JA AND GAMETOPHYTES OF SELAGINELLA 183 unshed and the sporangia are still attached to the strobilus. At this period the strobilus ceases to form new sporangia. The strobilus of S. rupestris retains its physiological connection with the plant until the embryo has produced cotyledons and a root. 15. In the early autumn S. apus sheds all strobili whether fer- tilization has occurred or not. S. rupestris retains its stroblii throughout the winter and fertilization occurs in the spring. 16. S. apus forms twelve to fifteen megasporangia in each strobilus, all of whose spores are generally fertile. The com- paratively limited number of embryos formed is due probably either to the limited number of fertile microspores, or to mechanical! difficulties in the way of fertilization, possibly both. S. rupestris produces strobili far in excess of the number of purely vegetative branches, the majority of which develop only sterile spores except under very favorable conditions. The sterile spores are shed profusely during the summer, and the strobili which remain on the plant throughout the winter retain their power of apical growth. The first sporangia formed in the early spring contain microspores. There is but a limited number of these ; then follows under favorable conditions either fertiliza- tion or the development of megasporangia, which continues until checked by the development of embryos in the older regions of the strobilus, or in case fertilization fails by the approach of winter. TECHNIQUE. The tender tips of the strobili offer no special difficulties in preparation for the microtome, but the older sporangia, espe- cially the closely compacted strobili of S. rupestris, are exceed- ingly resistent to the entrance of reagents, and the hardened epidermis of the sporophyll, combined with the thick walls of sporangia and spores, render sectioning difficult. Certain stages of the spores are peculiarly liable to collapse. These technical difficulties probably explain many gaps in our knowledge. The following method was followed in preparing the material for this investigation. 184 BOTANICAL GAZETTE [SEPTEMBER KILLING AGENTS.— Flemming’s weaker fluid, Hermann’s chromacetic acid, and bichromate of potassium combined with acetic acid, gave the best results. Corrosive sublimate, absolute alcohol, picric acid, Merkel’s and Perenyi’s fluid were tried, but the results were unsatisfactory in that I failed to get such suc- cessful staining after their use. These fixing fluids were used boiling hot. In the case of the Flemming, the chromic acid and water were brought to the boiling point, the osmic and acetic acids quickly added and the mixture poured over the strobili which had previously been removed from the plants. The vessel was covered tightly and the contents allowed to cool gradually to a temperature of 30° C. The spores were fixed in twenty four hours, but often were left in the killing fluid two or three days with no deleteri- ous effects. Water of approximately the same temperature (30° C.) was used for washing, in very large quantities, and changed frequently for two days. Sometimes it was more con- venient to use cold running water, in which case three days’ washing was necessary to remove all acid. The dehydrating process was equally gradual: twelve hours each in Io per cent., 20 per cent., 30 per cent., 50 per cent., 75 per cent., and 80 per cent. alcohol respectively. In g5 and 100 per cent. three days each, and the alcohol changed each day. Xylol was added to the specimens with even more caution, as an examination of material from day to day during these manipulations disclosed the fact that at this point danger of collapse was greatest. Six intermediate grades between absolute alcohol and pure xylol were employed, each for twenty-four hours. The strobili remained in pure xylol until they became transparent, which frequently was not for a week, dependent upon the age of the spores. When this condition was obtained, the xylol was replaced by fresh, and small bits of soft paraffin (melting point 30° C.) were added gradually as long as they would dissolve at the temperature of the room. This usually required two days. Thence the vessel was removed to the bath (temperature 40° C.), which was raised slowly to 54° C., soft paraffin being added 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 185 during the time up to the point of saturation. At this time the cover was removed from the vessel to facilitate the evaporation of the xylol, and harder paraffin added in small quantities at intervals. At the end of two or three days the temperature was raised to 65° C., and maintained for one week, after which time the xylol had evaporated, and the strobili were infil- trated with paraffin. At first, on the supposition that such protracted exposure to reagents and heat would injure the more delicate very young sporangia, the tips of strobili were removed, carried through the various media in much less time, and subjected to a temperature of 50°C. for about ten hours only. Comparisons later showed that these precautions were unnecessary, and that the longer exposures produced better results, even in the apical cells. More- over, having a strobilus cut zm /ofo was of the greatest value in interpretation. Further experiments prove that considerable latitude in the direction of longer periods of immersion in the various fluids are not injurious, but all attempts to hasten results by shorter exposure than the period stated above were unfortu- nate. By an oversight, at one time the temperature rose in the bath three successive nights to 75°C., but with no injurious effects. Some difficulty was experienced in imbedding. The paraffin was inclined to shrink away from the rough surface of the spore wall, which caused the sections to drop out of the paraffin ribbon when transferred to the slide. The difficulty was overcome by removing the strobili to the imbedding dish from the bath, letting them cool off slightly (to about 40° C.), pour- ing on paraffin at a temperature of 80° C., then cooling as rapidly as possible in ice water. No further trouble was experienced and the strobili could be sectioned without diffi- culty. Stains.—The best stains for the youngest stages of sporan- gia are iron haematoxylin (Haidenhain method), and the so- called Flemming triple stain—saffranin, gentian-violet, and orange G. Frequently gentian-violet and orange G were employed without saffranin; for the gametophyte development, 186 BOTANICAL GAZETTE [SEPTEMBER after the appearance of the proteid matter in quantities that obscured the other cell features, cyanin and erythrosin, after treating the sections on the slide with dilute sulfuric acid and chloroform, produced very satisfactory results. The power to take up the stains was retarded in the case of the proteid granules, whereas the nuclei and cell walls were more readily and more brilliantly dyed after this treatment. Karyokinetic figures were especially clear. It remains my pleasant duty to express my thanks to Dr. John M. Coulter, and to other members of the Botanical Depart- ment of the University of Chicago, for many suggestions offered in the course of this investigation. To Mr. E. J. Can- ning, Head Gardener of the Botanic Gardens of Smith College, I am deeply indebted for aid in collecting and growing material. BIBLIOGRAPHY. 1846. METTENIUS, G.: Beitrage zur Botanik, I 1851. HOFMEISTER, W.: Vergleichende Untersuchungen der Keimung, Entfaltung und Fruchtbildung héherer Kryptogamen, etc. Leipzig. pp. 111-123 1867. Rozx, E.: Les anthérozoides des Cryptogames. Ann. des. sc. nat. 7: 87-103. 1869. MILLARDET, A.: Le prothallium male des cryptogames vasculaires. Strasbourg. 1871. PFEFFER, W.: Die Entwickelung des Keimes der Gattung Selagi- nella. Botan. Abhandl. herausgeg. von Hanstein 1: 1-80. 7 #7. Bonn. 1873. STRASBURGER, E,: Einige Bemerkungen iiber Lycopodiaceen. Bot. ees 1874. HEGELMAIER, F.; Zur Kenntniss der Lycopodiaceen. Bot. Zeit. 32: — 1874. SACHS, J.: Lehrbuch der Botanik. 4 ed. Leipzig. pp. 454-475- 1877. DyER, W. THISELTON: Morphology of Selaginella. Nature 15: 489- 1879-80. SADEBECK, R.: Die Gefasskryptogamen, in Schenk’s Handbuch der Botanik 1 : 300-30 1879. BuLastus: Ueber ‘Statgcthe lepidophylla Spring. Jahresber. d. Vereins f. Naturwissensch. zu Braunschwei 1880. GOEBEL, K.: Beitrige zur Eutwickelunsgeschichte der poranie. Verhandl. d. sai med. Gesellsch. zu Wurzburg N. F. trage zur vergleichenden senior eam eat ee eae! eaecuas II. Bot. Zeit. 39 :681-694, 697-706, 714-720. 1 pl. Z 1881. BS 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 187 1885. BELAJEFF, W.: Antheridien und Spermatozoiden der heterosporen Lycopodiaceen. Bot. Zeit. 43 : 793-802, 809-819. 7 pi 1885. ————: Antheridien und Antherozoiden der heterosporen Lycopodia- ceen. Moskau. p. 71. ¢ pls. Ueber Bau und Entwickelung der Spermatozoiden der Gefiss- kryptogamen. Ber. der deutsch. bot. Gesells. 7: 122-125 1890. WojinowIc, W. P.: Beitrige zur Morphologie, Anatomie und Biologie der Selaginella lepidophylla Spring. Inaugural-dissertation. Breslau. p. 36. ¢ pis. 1894. Bower, F. O.: Studies in the morphology of spore-producing mem- bers. Part I. Equisetineae and Lycopodineae. Proc. Roy. Soc London 54:172-176; Nature 48 : 598-599 1894. ————: Studies in the morphology of ice bindaciene members of Equisetineae and Lycopodineae. Phil. Trans. Roy. Soc. London. 185 :473-572. pls. 42-52. 1894. HEINSEN, E.: Die Makrosporen und das weibliche Prothallium von Selaginella. Flora 78 : 466-496. 7 pi. 1895. CAMPBELL, D. H. Mosses and ferns. pp. 485-504 1896. ARNOLDI, W.: Die Entwickelung des weiblichen Vorkeimes bei den heterosporen Lycopodiaceen. Bot. Zeit. 54: 159-168. 1897. BRUCHMANN, H.: Untersuchungen itiber Se/aginella spinulosa A. Br. . Gotha. p. 64. 3 pls. 1900. GOEBEL, K.: Organographie der Pflanzen, inbesonderer Archegoni- aten und Samenpflanzen. I. Teil. Allgemeine Organographie, Jena. 1898; II. Teil, ee Organographie. 2. Heft. Pteridophyten und Samenpflanzen. Jen 1g00. FitrinG, H.: Bau u “ Entwickclanmspeschichte der Makrosporen von Isodtes und Selaginella und ihre Bedeutung fur die peeipes des Wachstums pflanzlicher Zellmembranen. Bot. Zeit. 58: 107-165. 2 1889. EXPLANATION OF PLATES V-IX. All figures were drawn with the aid of a camera lucidaand a Bausch and Lomb microscope, and were reduced to one fourth their original size in reproduction. PLATE V, Selaginella apus. Oc. ¥%, otl immersion +3. Fig. 1. Vertical median section of tip of strobilus showing the apical cell, a young sporophyll whose apical cell is just established, an archesporial cell (shaded), and a two-celled sporophyll (istingsed by black nucleoli). FIG. 2. Radial section of portion of strobilus showing sporophyll and two superposed cells (shaded), resulting from a transverse division of the archesporial cell. The superficial cell by repeated division in one plane Sa a a Tak 188 BOTANICAL GAZETTE | SEPTEMBER eventually forms the initial sporangium wall. The hypodermal cell is the initial sporogenous cell. The first cell of the ligule is distinguished by a black nucleolus. Figs. 3 and 4. Radial section of portion of strobilus showing two super- ficial archesporial cells (shaded). Fic. 5. Radial section of strobilus showing three-celled sporophyll (near apex), a two-celled sporangium (shaded) near the base of the subtending sporophyll, and the early stage of the vascular system. Fic. 6. Radial section of strobilus showing four-celled sporangium pos- sibly formed by two archesporial cells which divided by transverse walls. A stage subsequent to figs. 3? and ¢ Fic. 7. Radial section of older sporangium. Sporogenous cells are shaded, and are represented with black nucleoli. Fic. 8. Vertical section of sporangium showing definite radial arrange- ment of cells and plane of cleavage dividing it into two groups of cells, each the progeny of one of the two superficial archesporial cells shown in /g5. 3 and 4. Fic. 9. Vertical median section of sporangium more advanced. Sporan- gium wall at this stage consists of one layer. Fic. 10. Median transverse section of sporangium showing the sporan- gium cells, the tapetum, the inner and the outer layers of the sporangium sie . Section of sporangium after the spore mother cells are estab- ites ue this stage the sporogenous cells are distinguished by two con- spicuous masses of granular matter which lie against the nucleus iG. 12. Group of sporogenous cells in synapsis stage. Probably micro- spore mother cells. . 13. Detail of small portion of megasporangium showing megaspore ie cell (distinguished by the lumps of matter on either side of the nucleus) lying near the glandular tapetal cells. Two sterile sporogenous colt are at the left and below the megaspore mother cell. Fics. 14-24. Stages in the development of the megaspores. Fig. 16 shows the characteristic sextuple spindle stage previous to the separation into spores, as seen in fig. 78. The four nuclei pass from the bases to the apices of the newly formed spores, as shown in figs. 18-22 Figs. 25-28. Details of cell division in the final ‘ages of development of the female prothallium. Fig. 25 represents a portion of a section of the stage represented in fg. 49 to show thickness of protoplasmic vesicle and the invasion of fibrillae blocking off the separate nuclei. FIG. 29. Section of archegonium. The egg and ventral canal cell are lying in a plane perpendicular to the long axis of the archegonium. The egg is the larger cell with black nucleus. 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA 189 FiG. 30. Section of archegonium in which the egg lies below the ven- tral canal cell, and the neck canal cell above it. Two tiers of neck cells are FIG. 31. Stage younger than that shown above, before neck cells are Fig. 32. Archegonium with egg and ventral canal cell laterally placed. No trace of neck canal cell. Neck consists of four tiers of cells, each tier comprising an upper (superficial) cover cell, and one neck cell. The cells abutting on the venter belong to the prothallium. Figs. 33 and 34. Mature archegonia. In fig. 37 a see is lying on the receptive spot of the egg. Fig. 74 shows a one-celled em PLATE VI, Selaginella apus. Oc. 2, otl immersion y. FIG. 35. Section of young megasporangium at earliest stage when mega- spore mother cell is distinguishable from sterile cells. For detail see fig. 77. Fig. 36. Section of older stage of megasporangium. Megaspore mother cell has moved near center of sporangium. - 37. Oblique section of megasporangium showing young tetrad, and sterile mother cells, floating in slime composed in part of disintegrated sterile cells and in part of a secretion poured out by tapetum. Fig. 38. Section of somewhat older stage of megasporangium containing group of four megaspores (only three are represented). Each spore consists limpid fluid and possessing a very small nucleus. e spaces intervening between vesicle and median layer and between the latter and the exospore are filled with fluid, FIGS. 39-41. Successive serial sections of a tetrad, to illustrate the fact that the exospore is a continuous envelope common to the group of spores and splits as they move apart. Fig. 42. Section of group of megaspores but little older than those in fig. 38.- Filamentous processes shown between spores. The mother cell membrane, beyond whose boundary the radiations do not pass, envelops the tetrad. 43. Section of two megaspores more advanced. The spore at the left is cut nearly ir half from apex to base, that at the right is a slice across the apex. Protoplasmic vesicle grown but little larger than in fig. 2. Spine-like processes upon exospore in connection with radiations. Mother cell membrane still evident. FiG. 44. Section of the female gametophyte showing the protoplasmic vesicle with two nuclei. The radiations extend across the spaces between vesicle and median layer, between median layer and exospore, and between the latter and the original spore mother cell membrane. 1go BOTANICAL GAZETTE [SEPTEMBER Fic. 45. Section of female gametophyte whose vesicle has five nuclei and no cell walls. The exospore has grown to the size of the megasporangium as shown in fig. 78. The median layer has divided into the endospore (rep- resented in section as a broad black band), and the mesospore which can be distinguished only as a delicate layer (represented by a single line) just with- out the endospore. The exuspore is of spongy appearance and is still in contact with the persistent spore mother cell membrane. Fluid, presumably protoplasmic in nature, fills intervening spaces. 1G. 46. A transverse median section of a female gametophyte in “film” stage. The megaspore, endospore and protoplasmic vesicle (studded in its apical region with large free nuclei) have stretched to the dimensions of the exospore, against whose inner surface they form a layer consisting of three delicate lamellae. The central vacuole is filled with clear fluid. The same stage is shown in figs. g7 and g8 the former representing a surface view, the latter a median section. PLATE VII, Selaginella apus. ‘ All figures except 56 and 58 drawn with oc. 2, obj. }. Fig. 56 is drawn with same combination as plate V; fig. 58 with oc. 2, obj. }. ) 1G. 47. Surface view of the protoplasmic vesicle as if seen through spore walls. 1G. 48. Vertical median section of the same stage. The nuclei are massed in the apical region and the interior of the vesicle is occupied by 4 vacuole. 1G. 49. Surface view of the gametophyte at the moment when proto- plasmic fibrillae appear in the apical region and radiate over the surface of the gametophyte. The nuclei in the lower part of the figure are in process of division. The vesicle has increased in thickness (cf. fig.50), and is beset with proteid granules. Contents of vacuole a homogeneous liquid. 1G. 50. Median section of same. Fic. 51. Median vertical section of female gametophyte. -Protoplasmic vesicle much thicker in apical region where the nuclei are disposed in several layers diminishing to a single one in the equatorial region. Fibrillae per- meating the nucleated portion have outlined indefinite areas but there are no cell walls, Contents of vacuole an emulsion. Fie, 52. Section showing a portion of the megasporangium with female gametophyte iw situ to show the differentiation of the wall. In the lower- most part of the figure the fragile cells of the area of dehiscence are seen 10 cross section. The inner layer of the wall probably supplies the outer with nourishment. The vestiges of the tapetum are ‘upon its inner face, as a pavement layer, and a few sterile mother cells lie at the left next the wall. 1G. 53. Vertical median section of female gametophyte showing the differentiation of cells into an apical superficial layer from which develop Cetera 1901] SPORANGIA AND GAMETOPHYTES OF SELAGINELLA IgI archegonia, and a lower vegetative region. The delimitation of the arche- gonial region is shown by the trefoil shaped cleft seen in the lowermost spore of fig. 58. Figs. 54 and 55. Section of female gametophyte showing archegonia, Details of the process of division in the unshaded cells of fg. 54 are rep- resented in figs. 26-28. FIG. 56. Two embryos, one of two cells, the other of three, lying side by side in one venter. All walls shown are parallel to the axis of the arche- gonium. Compare relative position of these embryos with that of the egg and ventral canal cell in figs. 29, 32. FiG. 57. Embryo zz situ. The root lies toward the apex of the gamet- ophyte. At the right the club-shaped foot of large cells is embedded in the prothallial tissue. The embryo is bent with respect to its long axis so that the stem apex and cotyledons are brought nearer the root than appears in the drawing, which is a reconstruction from serial sections. The two leaves with their relatively large ligules are developed successively and envelop the tip of the stem (shown by dotted line). The embryo is still within the spore wall and sporangium. Fig. 58. Vertical section of megasporangium containing three female gametophytes. The lowermost shows the trefoil-shaped cleft in the apical region of the spore wall, exposing the archegonia. The two other oblique sections are not so advanced in development. The pad of tissue at the base of the sporangium lies above a region of storage cells in contact with the vascular bundles of axis and sporophyte. PLATE VIII. Selaginella apus. All drawings made with oc. ?, oil immersion, obj. yy, excepting 60-62. Figs. 60, 61 were drawn with oc. 2, oil immersion, obj. yy. Fig. 62 with oc. 3, obj. 2. FIG. 59. Section of microsporangium, showing microspore mother cells, goneoes inner and outer layers of sporangium wall. G. 60. Median vertical section of microsporangium with mother cells in rene stage. Sub-archesporial pad developing at base of sporangi Fic. 61. Slightly oblique section of microsporangium showing sic hecoees in groups of four (tetrads). F1G. 62. Median vertical section of microsporangium to show sporangium wall at maturity. Dehiscence region cut across at apex. Sporangium Opens in two valves, the subarchesporial pad at top of pedicel serving asa fulcrum. Figs. 63, 64. Microspore mother cells during first division of nucleus. Figs. 65, 66. Sextuple spindle stage with nuclear plates. Fig. 67. Exterior view of young microspore. 192 BOTANICAL GAZETTE [SEPTEMBER Fic. 68. Section of tetrad, showing continuity of exospore around the four microspores, and spore mother cell membrane enveloping the tetrad. The protoplasm is a thin vesicle with a single nucleus, parietally placed, and surrounding a large central vacuole. Fic. 69. Exterior view of mature microspore showing pebbled exospore. Fic. 70. Vertical section of male gametophyte after first division of spore into generative and vegetative cells. . Male gametophyte as seen by focusing through spore coats. The ee generative nucleus lies near the wall, the vegetative cell not in foc Bice 72. Male gametophyte showing generative cell in section. G. 74. Male gametophyte showing lenticular vegetative cell at left. The eas of the content constitutes the generative cell with a centrally placed nucleus. Masses of deeply staining proteid matter appearing. G. 75. Section of male gametophyte in which masses of proteid matter have increased in number. Vegetative cell appressed to wall at left 1G. 76. The protoplasm is aggregated around the centrally placed generative nucleus and has sent out radiating processes, connecting it with a peripheral layer. The vacuoles thus isolated are filled with a semi-fluid ame matter, , 78. A common appearance, where a large symmetrical mass of proteid matter (solid black) in the central vacuole may be mistaken for a nucleus, 1G. 79. First division of generative nucleus. Persistent vegetative cell appressed to wall in lower portion of figure IGS. 80-87. Phases of the early divisions leading to the spermatozoid mother cells. Figs. 88-95. Sections in various planes showing the two cell complexes and their relation to the dissolving proteid masses by which they are envel- oped. The masses of food matter are separated by strands of protoplasm in Jigs. 88,93, 95 FiG. 96. Exterior view of male gametophyte displaying trefoil shaped cleft in apical portion. The complexes of spermatozoid mother cells protrude through the gap. FIG. 97. Male gametophyte with spermatozoid mother cells as seen by focusing through spore coats. The unusually persistent vegetative cell is represented lying above the mother cells. F1Gs. 98, 99. Male gametophyte at maturity. The endospore protrudes like a short pollen tube through a gap on the ruptured exospore, and contains a slimy homogeneous fluid which later is discharged with the spermato zoids. FIG. 100. A transverse section of a female gametophyte displaying the 1901] SPORANG/A AND GAMETOPHYTES OF SELAGINELLA 193 eggs and spermatozoids. The necks of the archegonia have been sliced off somewhat obliquely PLATE 1X. Selaginella rupestris. G. - Apex of a strobilus in radial section. A single archesporial cell hee at the base of a sporophyll. The cell with black nucleus is the initial cell of the next younger sporophyll of the same ra 1G. 102. First division of archesporial cell into a ibid and a superficial cell. FIG. 103. The superficial cell of the first division has divided into three wall cells, and the hypodermal cell into two sporogenous cells, forming a wedge-shaped mass. ; FIG. 104. Radial section of portion of strobilus. The largest sporophyll has ceased to grow at the apex which is converted into a branched spine The initial cells of the ligule at the base of the sporophyll are represented with black nuclei. The tissue is rupturing to form air chambers in the basal region. The sporangium has been outstripped in growth by the sporophyll next above it. The two shaded cells in the upper part of the figure represent the same stage of sporangium as seen in fig. 702 FIG. 105. Section of sporangium. The wall is nearly completed and the tapetum is becoming differentiated from the sporogenous complex of cells. The limits of the latter in the basal region not so clearly defined as repre sented in the figure. Fig. 106. Detail showing two megaspores and several sterile mother FiGs. 107-117. Phases undergone by megaspore mother cells preceding divisions into spores. Fic. 118. First division of megaspore mother cells into two spores. FIG. 119. Section through sister megaspores. A portion of the sculptured €xospore is removed from the upper surface of each, showing the proto- plasmic vesicle with a small nucleus floating in liquid. The spore mother cell membrane envelops the two spores. Fig. 120. Section showing incomplete division of the protoplasm of two megaspores. The nucleus of the spore mother cell has not divided. Fig. 121. Section through a single megaspore whose contour and large nucleus suggest that it is the direct product of a spore mother cell which has failed to divide. Figs. 122, 123. Older stages of megaspor Fig. 124. Median vertical section of tip s pails showing relation of Sporangia to sporophylls and vascular system. The meristematic regions at the bases of the sporangium pedicels grow upward slightly by causing the Megasporangia to appear sunken in the hollows thus formed. There were only megasporangia in this strobilus. 194 BOTANICAL GAZETTE [SEPTEMBER Fig. 125. Transverse section of strobilus through microsporangia. In the center of the figure is the axial strand surrounded by air chambers inter- spersed with meristematic tissue. At the right and left are two micro- sporangia, each showing the septum which incompletely separates it into two loculi. The rest of the diagram represents the cross sections of six sporo- phylls with their ligules. Fic. 126. Apical portion of strobilus showing young sporophytes pro- truding from undetached megasporangia. HuLL BoTANIcaAL LABORATORY, THE UNIVERSITY OF CHICAGO, THE ECOLOGICAL RELATIONS OF THE VEGETA- TION OF WESTERN TEXAS. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORA- TORY... AX: WILLIAM L. BRay,. (WITH TWENTY-FOUR TEXT FIGURES) [Continued from p. 123.] THE GREAT PLAINS. THIS great region is naturally divided into two sub-regions, (1) the prairie plains, from which, except for scattered rem- nants, the original Great plains deposits have been removed, and embraced in the central and east central provinces of Hill (fig.1); and (2) the remaining areas of Great plains proper (Llano Estacado, Edwards plateau, and Stockton plateau). The prairie plains.—This sub-region embraces the grass coun- try of central and north Texas west of meridian 97.5°, an area Over 50,000 square miles in extent. As previously stated, it is a vast region of denudation, presumably from a former plain which must have stood at a much higher level. Whatever may have been the overlying formations, the present stage of erosion pre- sents exposures of various formations, each of which owes its individuality chiefly to the character of the strata now undergo- ing reduction. Taken as a whole the dominant vegetation is a grass formation, but upon the areas of sand and gravel exposure, and the hills, bluffs, and streamways, timber formations prevail. The provinces are the Grand prairie and those of the central denuded region; embracing (1) the granite country, (2) the Car- boniferous area (Brownwood and Palo Pinto countries), (3) the upper cross timbers, (4) the red beds country, and (5) the Cal- lahan divide. THE GRAND PRAIRIE— Geologically this is a Lower Creta- ceous area extending from the Colorado river (its eastern Igor] 195 196 BOTANICAL GAZETTE [SEPTEMBER margin near Austin) northward beyond the Red river, its eastern © boundary being the Black prairie or rich black land area of Texas. The western boundary of this province is a prominent and irregular escarpment overlooking the central denuded region, and the upper cross timbers at the north. The prairies, which are the dominant physiographic feature, are wide undula- tions interrupted by broad erosion valleys and occasional butte remnants of the former plain. The elevation of the province is from 800 to 1200 feet. The soil covering is but a thin veneer- ing overlying alternating beds of chalky limestones and marls, which are very poor receivers or retainers of moisture. This condition suffices to offset the comparatively high annual rain- fall (28 to 32 inches), so that this is a province of pronounced xerophytic grass formations, but still a transition from the meso- phytic Austro-riparian to the xerophytic Lower Sonoran zone. In its temperature relations the province stands almost on the transition to the Upper Sonoran and Carolinian zones respect- ively, thus differing notably from the Rio Grande plain province which passes into semitropical. Pure grass formations.— Ecologically the grass formations are strongly xerophytic and partake of the characters of the grass vegetation of the more westerly plains, though in a moderate degree. As already seen, the mean annual rainfall is sufficient, other things being equal, to give existence to semi-mesophytic formations. But the periods of brilliant sunshine and dry air which prevail, especially during the growing season, very quickly dry out the superficial covering of soil and drive vegetation into the resting stage. Under present conditions of pasturage the grass covering has been sadly reduced from its former occurrence. Formerly it was very compact and luxuriant. With the break- ing up of this, moisture relations have changed, and the forma- tion is now more xerophytic. Floristically it is impossible to say, with the meager data available, how the association of spe- cies compares with that of prairies and plateau plains westward, or to what extent the formation has changed under pasturage- It is certain from the general aspect of the formation, however, ae Igor] VEGETATION OF WESTERN TEXAS 197 that the dominant species here are not dominant ones on the more western plains. In brief, the ecological type known as prairie grasses exists here, as distinguished from the high plains type. The grass vegetation of the Grand prairie is accompanied by avery abundant vegetation of prairie annuals and herbaceous Fic, 7.—Rio Grande chaparral at Laredo: sage (Leucophyllum), “all thorn” (Rocneicies Mexican persimmon (Diospyros Texanum), and eight or ten other species perennials, with lignescent, tuberous, or bulbous underground parts. These assist in giving the province individuality as a grass formation. formations of prairie annuals—Since the most conspicuous formations of this kind occur in other provinces, this type will be discussed later under a separate heading. Chalk soil formation of herbaceous perennials.—The species asso- ciated in this formation number perhaps thirty-five or forty. It occurs throughout the province on the thinnest soils, but comes 198 BOTANICAL GAZETTE [SEPTEMBER out especially on bare chalk slopes. Sometimes a single species prevails upon such places, but commonly there occurs a mixture embracing nearly all the species. Thus a careful collection upon a typical chalk area will yield more than three fourths of the species of the typical formation (fig. 74). In all species the underground parts are reservoirs of food (and not infrequently of water) protected against the extremes of heat and drouth to which the arid soil is subjected. When the soil is wet by rains these subterranean organs quickly throw up vegetative organs, and perhaps flower and mature fruit before the brief period ends and they are again driven into the resting stage. The underground parts are of several types, as succulent fibrous roots with a thick zone of mucilaginous or saponaceous tissue (Krameria secundiflora, Yucca rupicola); woody subterra- nean tuberous caudex (Liatris pycnostachya); mucilaginous bulbs with impervious coats (Allium, Cooperia), and deep irregular or fusiform roots with hard flinty (sclerenchymatous) coat (Asclepias decumbens). THE CENTRAL DENUDED REGION.—The granite and Carbonifer- ous areas and the upper cross timbers together constitute a belt lying below the western escarpment of the Grand prairie, where the prairies are more or less interrupted by a more broken relief, which in part of the granite area becomes positively mountainous. This belt is marked on the more level areas by a surface cover- ing or by deep beds of sand and gravel, where conditions favor the occurrence of an open timber formation and even a close forest covering in the upper cross timbers. Discussed from the standpoint of grass formation, the granite and Carboniferous areas (Burnet, Brownwood, and Palo Pinto countries of Hill, see fig. 1) are open-timber grass prairies, which with the spread of the mesquite (Prosopis juliflora) are losing their last remnants of pure grass formation. The grass formation of this belt (the upper cross timbers not included) appears to have about the same character as that of the sand plains of the Rio Grande plain. The ecological condi- tions are very similar except as to temperature, and the results EE — —— oe Igor] VEGETATION OF WESTERN TEXAS 199 of this difference are apparent even superficially in the absence of certain semi-tropical species common at the south, and in the presence of Upper Sonoran or Carolinian species whose range extends into the middle northern states, THE RED BEDS PROVINCE.—The vast level plains of reddish, chocolate soil make of this province one of the most unique fea- tures of the entire Texas region. This geological formation underlies the Staked plains, and has been exposed by the ero- sion of that area in the Pecos and Canadian river valleys. But the red beds plains lie between the Staked plains and the Grand prairie (more accurately the upper cross timbers), a width of 150 miles, and stretch away through Oklahoma and southern Kansas, a distance of 400 miles. Erosion and weathering have left wide, flat stretches of reddish landscape, either covered with fine, chocolate, silty sand, or exposing the bare rock. Violent bursts of rainfall wash away acres of prairie, often to the depth of several feet, carrying the chocolate flood silt to the coast country. Within the Texas region there are no large relief features in the red beds province, although the broad erosion valleys have to a certain extent the ‘‘ Bad lands” aspect. The great extent of this province and its geographical position give it climatic relations which are apparent in the character of the vegetation. As to atmospheric moisture, the stretch of 150 miles (the breadth of the province) from east to west means a variation in rainfall from 27 to 20 inches, and of evaporation capacity from 50 to 65 inches. The vegetation of the western part of the Province has a more extreme xerophytic character than that of the eastern part. The northeastern parts sustain extensive mesophytic or semi-mesophytic culture formations with reason- able certainty. It is particularly a wheat belt. Toward the western border these formations become impossible except with seasons of higher rainfall. As to temperature, while the entire province falls in the Lower Sonoran zone, its position as a whole far to the north of the Rio Grande plain gives it floristic relations more in common 200 BOTANICAL GAZETTE [SEPTEMBER with the Upper Sonoran areas to the north. But within the province itself there exists this diversity, that the southwestern border receives much from the arid Sonoran flora of the south- west, at the northeast the incoming flora is a semi-mesophytic one from the northern prairies. This relation to the flora north- Fic. 8.— Edge of chaparral thicket, Port Lavaca. ward is promoted by the continuation of the red beds formation as far as southern Kansas. The ecological conditions are most favorable for a pure grass plains formation. While it lies beyond the zone of chaparral formations, the mesquite already covers most of the western half of the province, undoubtedly modifying the structure and content of the grass formation. Asto the general type of grass formation, the semi-arid climate makes of this a characteristic dry grass plains flora. Without a more specific study of this flora it will be impossible to show satisfactorily wherein lies its Igor | VEGETATION OF WESTERN TEXAS 201 individuality as compared with the Grand prairie on the east and the Edwards plateau and the Staked plains on the south and west. It is probable that there is a real distinction, and that it lies chiefly in the peculiarities of the red beds formation. Two specific formations are constantly to be distinguished, (1) the Hilaria Jamesit formation, which occupies the lower stretches where sandy loam lies deeper, and appears like a vast field of heavy cultivated hay; and (2) the Aristida fasciculata formation, which occurs on ridges alternating with the lower areas of FAitlaria Jamesti, and in its turn presents large tracts of waving grass fields. Formations of prairie annuals—It is not possible at present to say to what extent or wherein the prairie annual formations are distinct from those of other provinces. The soil conditions are such as to favor many species devoted to sandy soils. In the southern part of the province there are species from the south and southwest. At the north the prairie annual flora of Okla- homa and Kansas, in the same longitude, appears to be largely predominant. These formations prevail in such abundance in the case of some species as to form an important factor in the forage supply, especially in the early spring, when there is a solid mass formation. This disappears before the climax of the grass vegetation season (fig. 17). The Great plains proper.— There are included in this region three provinces of the higher plains, namely Edwards plateau, the Staked plains, and the Stockton plateau (fg. 7). THE Epwarps PLATEAU.—This province forms the continua- tion of the Great plains east of the Pecos, from the southern limit of the Staked plains to the downfall at the Balcones escarp- ment overlooking the Rio Grande plain. The entire southern half between the Colorado and Devils rivers is marked by such deep erosion as to be mountainous. While a grazing country, this is not characteristically a grass plains, but a timbered area, and hence to be discussed especially under timber formations. But the grass vegetation even here contests for predominance, except on the most rugged hills. , 202 BOTANICAL GAZETTE [SEPTEMBER The plains prairies —To the northwest there is an area of the Edwards plateau which has not yet come to the stage of active and deep erosion, but is a typical open level plains country, with low relief lines of erosion remnants, and draws which lead into the cafions of the southern erosion area. This plains prairie is embraced approximately in the counties of Upton, Glasscock, Stirling, Concho, Crockett, Sutton, Schleicher, Tom Green, and Irion. This Cretaceous area, both by its altitude and its westerly position, shares climatic conditions to a marked degree with the Stockton plateau and the Staked plains. Asa result, the grass vegetation is of a more xerophytic character than in any of the provinces previously discussed. It is, in fact, almost fairly within the ‘short grass” country. Like the Grand prairie, this Cretaceous formation increases the xerophytic tendencies, so that here we find again a strong element in the perennial vege- tation, which is more suffruticose than herbaceous, and made up of species with more xerophytic aspect and with affinities to the flora of the trans-Pecos provinces, But this element is in every way the counterpart of the chalk soil lignescent vegetation of the Grand prairie. In addition, the chaparral formations of the Stockton plateau and southwestward are encroaching upon this area. Tue STakeED pLains.—The Staked plains of the Texas region are simply a continuation southward of the Great plains area of western Nebraska, Kansas, and eastern Colorado, but cut off from the main area by the deeply eroded channels of the Cana- dian river on the north, and the Pecos on the west. To the north of the cafion of the Canadian river, Texas contains also 4 portion (the Pan Handle) of the main body of the plains. “This province is a vast constructional plain made up of flood débris from the mountains at whose base it lies. This mass of loose unconformable stuff is a monotonously flat plateau plain, whose only surface relief is the billowy swells and’ the shallow saucer-like depressions” (Hill). The open, porous, often sandy texture of the soil furnishes a favorable receiving area for | 1901 ] VEGETATION OF WESTERN TEXAS 203 rainfall. The soil water level is too far beneath the surface to be drawn upon even: by deep-rooted trees. The vegetation is dependent, therefore, upon the immediate surface for its mois- ture, and alternates quickly between active growth and the dor- mant conditions according as the surface is moist or dry. But the porosity of the soil enables it to retain a larger percentage of precipitated moisture, so that the xerophytic conditions are not so extreme as the low rainfall (15 to 18 inches), the high FIG. 9.— Characteristic occurrence of Acacia amentacea or black chaparral; on silty clay aauke plain near Eagle altitude (3500 to 4000 feet), the high evaporation capacity of the air (60 to 70 inches), and its high velocity would lead us to expect. The Staked plains do not terminate at the south in an escarpment, but pass into the plains portion of the Edwards plateau by a broken series of outlying buttes and sand hills and plains. These sand plains not only form the transition from the Staked plains, but extend northward néarer their western side beyond the center of this province. In its temperature relations the Staked plains is mainly Upper Sonoran, The transition from Lower Sonoran is made about the 3500 foot contour. Beginning at the southern limit of the plains, the Lower Sonoran vegetation, notably the chaparral 204 BOTANICAL GAZETTE [SEPTEMBER formations, is represented in great strength. In going north- ward, ascending the plains, this vegetation becomes gradually more dilute, finally leaving a pure grass plain. The most apparent index of transition is furnished by the mesquite, which persists long after every other Lower Sonoran shrub has disap- peared. Leaving a jungle of large mesquite below the escarp- ment, one comes upon a reduced growth upon the plains above, which becomes more and more dwarfed until at 3500 feet it is a mere clump of switches a foot tall, disappearing then absolutely. The same transition occurs in ascending the plains from the red beds province north of Red river, and where the Canadian cuts its cafion across the plains a Lower Sonoran flora has followed. The grass formation of the summit plains is a solid grass floor of the short buffalo grass type. These grasses, in harmony with the sudden transition from moist to arid conditions, are adjusted to rapid changes from the dormant condition to luxu- riant growth, and with the well-known characteristic that the growth already made when dry periods come on becomes pet- fectly cured as it stands, retaining all of the nutritious qualities which make these grasses economically valuable. All that remains alive during the dry periods are segments of the stem, which may be regarded as equivalent to buds or tubers oF rhizomes. These are especially rich in carbohydrates. They are not underground nor connected with the soil by living parts, but merely anchored by the dead fibrous roots of the old growth. The cured grass serves naturally as a protective covering for the dormant living parts, although these are not often killed when the dry covering is burned off. Indeed, it is this quality of being able to endure fires that gives to these grasses the great advantage they possess in their constant struggle against woody vegetation (fig. 23). The herbaceous species which accompany the grass vegeta- tion are chiefly annuals. The lignescent rooted perennials so characteristic of the Cretaceous plains and prairies are mostly wanting, which is in harmony with the open porous soil texture. The very abundant Yucca angustifolia is an exception to this 1901] VEGETATION OF WESTERN TEXAS 205 fairly constant condition. As to their affinity floristically, these annual species, while containing a considerable element common to Texas prairies south and southwest, are those which range over the whole stretch of plains to Nebraska and the northwest, thus indicating more positively the Upper Sonoran character of the province (fig. 78). The true value of the zonal areas of the Staked plains is not strongly brought out, because the dominant grass vegetation is not so sensitive to temperature differences. Yet it seems pretty certain, though not yet established, that some of the very important species of grasses found in southern and trans-Pecos Texas and north Mexico do not extend upon the Staked plains, but that rather the floristic content of the formation agrees with that of the plains northward. The grass formation is broken on the sand plains of this province because of the shifting character of the sands, This sand zone according to Dr. Havard, is entered upon going about twenty miles west of Odessa. ‘It extends thence south and east nearly to the Pecos and north to the very center of the plains. In this zone are the sand hills, a dreary chaotic belt of reddish sand, tossed by the wind into hillocks, cones, and ridges of various sizes and shapes.’’7 The lower belt of the sand zone marks the transition from Staked plains to the Cretaceous Edwards plateau. This is the only portion of the sand plains of which we have data as to the plant formation. Dr. Havard says of it: ‘‘The only grasses seen were a stout Andropogon (near A. furcatus) three to five feet high, with running roots holding the loose soil in their meshes; a Sporobolus (probably a form of S. eryptandrus), like- wise erect and tall; and a large form of Cenchrus myosuroides.”’ This indicates that the characteristic plains formation is entirely wanting. The herbaceous annuals are also those which prefer sandy soils. It appears from Dr. Havard’s report that woody vegetation is the more dominant formation. HE STOCKTON PLATEAU.—This is a subdivision of the 7 HAVARD, Contrib. U.S. Nat. Mus. 8: (no. 29) 1885. 206 BOTANICAL GAZETTE [SEPTEMBER Edwards plateau, cut off by the cafion of the Pecos river. On the west the province adjoins the eastern front ranges of the mountains. Most of this area is a typical grass plain formation, of the general type found on the Staked plains and on the plains portion of the Edwards plateau. It will be seen that t so G. 10.—Chaparral formation at Corpus Christi; the growth shown is ae tall as dias which covers the country generally in this region; the genera are C Opuntia, Condalia, Zizyphus, and Celtis, nearer the Rio Grande the surface is more broken, and that here woody vegetation predominates, also that the chaparral forma- tion is encroaching even upon the level grass plains. Compared with the Edwards plateau, this province as a whole is more elevated, and is in a zone of less rainfall and of greater evaporation capacity. It is more intimately a part of the arid sei a iam 1901] VEGETATION OF WESTERN TEXAS 207 plateau area of the Lower Sonoran zone, wich embraces north Mexico, southern New Mexico, and Arizona, and while it is only surmised that the grass formation would show this floristic rela- tion, it is certain that other formations show it in a marked degree. MOUNTAINS AND SOUTH PLATEAU SLOPE. The Rocky mountains proper terminate before reaching the Texas region, The mountain axis continues southward, how- ever, as an elevated plateau whose surface is much cut up by isolated mountain masses, some of which exceed gooo feet in altitude. These masses are composed of Carboniferous lime- stone and sandstone, of Cretaceous limestone, and of igneous material. The high plateaus among them are mostly bolson basins filled with the débris of their bordering mountains. The soil conditions here and the moisture conditions growing out of these furnish the direct causes which operate to determine the distribution of plant formation in a region which possesses extreme xerophytic conditions independently of edaphic fac- tors. Viewed from the standpoint ot the grass formation, this is a region possessing a grass floor climate in a surprising degree, for although the mountain masses are in some instances high and rugged, and covered with woody vegetation, the slopes of most of these up to 6000 and 7000 feet, the foothills, the wider plateau plains (for example, about Marfa and northwestward), and even portions of some of the desert bolsons, have a con- tinuous grass formation. But in the mountains of the Great bend and their intervening bolsons (except in the Rosillos mountains, where good and available range grasses are found, according to Havard), on bolsons with loose wind-swept débris, and on arid gravelly mesas, there is only the most scattering bunch-grass formation, or even no permanent vegetation at all. The bolson basins of this region, owing to the distribution of sedimentary filling, are lowest at the center and undrained. Consequently, the drainage water gathers at this place, in many cases accumulating in ponds or rather large lakes, and in all 2038 BOTANICAL GAZETTE | SEPTEMBER cases making alkaline flats of varying degrees of saltiness. The higher, less alkaline slopes of these bolsons are often covered with grama grass formation characteristic of the region. The alkaline flats, of course, possess a halophytic vegetation, of which salt grasses form a prominent part. On the whole, the grass vegetation of the mountain and south plateau provinces falls under three types of formation: (1) the close grass floor formation; (2) the open bunch grass forma- tion; and (3) the salt grass formation. Of these the third will be discussed under the special head of ‘halophytic formations.” The open bunch grass formation is floristically equivalent to the close grass floor formation. The open and bunched charac- ter of the formation is due to extreme dryness, and especially to the instability of the loose finer igneous débris, or the coarser gravelly or stony nature of the soils, The close grass floor formation may be designated “‘grama grass formation,” from the popular name of the one to several prevailing species. In this formation, the great majority of species making up the formations of the eastern provinces have been sifted out by increasingly arid conditions, and only those remain which mark the extreme limit of xerophytic adaptation in grasses. A summary of the adaptation features of this type of formation shows: (1) rapid transitions from active to dor- mant conditions ; (2) great resistance to extreme dryness and heat (including fires) while in the dormant stage ; (3) equally great recuperative power after extreme treatment, including apparent extermination from a given area; (4) large food stor- age of fats and sugars in portions which retain vitality during dormant periods, thus rendering quick growth possible ; (5) the quality of perfect drying zm situ, thus not only covering the soil and holding it in place, but also protecting the vital parts. PURE FORMATIONS OF PRAIRIE ANNUALS IN THE PRAIRIE PLAINS. In the preceding discussion of grass formations, the herba- ceous annuals of the prairies have been treated simply as acces- sory elements in the formations, but of value in characterizing Igor | VEGETATION OF WESTERN TEXAS 209 the formations differing somewhat in temperature and moisture conditions. But the annuals of the prairies in the west Texas region, as well as those in other parts of the Lower Sonoran zone, constitute a vegetation phenomenon which periodically over- shadows everything else. This phenomenon is the annual wave Fic. 11.— Hills covered with xerophytic timber, central mountainous region of Edwards plateau, of vegetative growth and floral display which sweeps over the prairies, temporarily submerging the grass vegetation itself. In the mutual adjustment of grasses and herbaceous annuals in this climate, the latter, mostly pure mesophytes, must take advan- tage not only of the returning rainy season but of the optimum period preceding the intense summer heat, and of the absence of competing grasses at that time of the year. It is a matter of record among cattle men that in the early spring, when grass forage is scarce, their herds often feed largely upon the 210 BOTANICAL GAZETTE [ SEPTEMBER abundant young vegetation of these prairie annuals. Thus the “tallow weeds” (Actinella linearifolia and Amblyolepis setigera), which cover many miles of range, are much prized. Not only does this annual vegetation sweep as a wave over the prairie in the early spring, but the individual species mass themselves in pure formations of incredible compactness and extent (fig.7g). Thus there may be successive waves of species. In harmony with the adaptation which leads to the massing of the species is the fact of the brilliance of floral display. This results in vast sheets of color. During the spring of 1900, which was unusually favorable for vegetation, the virgin prairie plains of Texas were literally a sea of color. In his survey of the Rio Grande plain during May, Vernon Bailey noted over twenty species which, as he expressed it, either occupied exclusively ‘‘acres of prairie” of “made miles of solid color.” This was the case in the lower Rio Grande country (Corpus Christi to Brownsville) previous to May 10. The season of display ended about four weeks later on the northern boundary. At Austin by the first of July (often even two weeks earlier) a solid grass formation occupies the ground which had just previously been exclusively occupied by the “blue bonnet” (Lupinus subcarnosus). The continuity of grass formation seems in no wise disturbed by the periodical dominance of the annuals. The species of these formations are of course dependent upon seed for their recurrence, and consequently are subject wholly to the caprice of rainfall during March and early April. When, as frequently happens, there is no rainfall of consequence during this period, the display of prairie annuals is very much reduced. Their adaptations are in the direction of seasonal adjustment rather than toward meeting xerophytic conditions. Il. WOODY VEGETATION. Since the region under discussion includes only so much of Texas as is marked by a distinctly xerophytic vegetation, it follows that, except for river bottom and sheltered cafion timber, the woody vegetation is as a whole one of xerophytic aspect. 1901} VEGETATION OF WESTERN TEXAS 311 But it is of varying degrees of xerophytism according to its relation to the zones of decreasing moisture from east to west, that in the eastern provinces possessing some of the elements of the vast mesophytic forest of east Texas, while the extreme of xerophytism is reached in the desert bolsons of extreme west Texas. Recalling the common conception of the west Texas region as a grass country, it becomes a matter of surprise to learn how great is the proportion covered by woody formations, that is arborescent or shrubby vegetation in sufficient amount to form the chief covering. This includes the following areas: the erosion area of the Edwards plateau; the hills, bluffs, and escarpment of the Grand prairie; the granite, Carboniferous, and upper cross timber areas; the higher mountain slopes and summits of the trans- Pecos mountains; the escarpment of the Staked plains and the Callahan divide; the chaparral of the Rio Grande plain, of trans-Pecos Texas, and of the lower Staked plains and adjacent Edwards plateau; the open streamways and the cafions; and finally many thousands of miles of mesquite-covered prairie. The energy and rate of encroachment of woody vegetation during the past half century lead one to believe that there is scarcely an area of consequence in the state that woody vegetation of some type will not occupy and cover more or less completely, granting of course that no artificial means are employed to check it. The woody vegetation is here discussed under several types of formation, based largely upon moisture relations. ARBORESCENT VEGETATION. XEROPHYTIC FOREST FORMATIONS OF THE EROSION MOUNTAIN- OUS AREA OF THE EpwarbDs PLATEAU (including the hill, bluff, and escarpment timber of the Grand prairie). —That portion of the Edwards plateau embraced between the Colorado and Devils rivers, and extending back 100 to 125 miles from the Balcones escarpment (southern downthrow of the Great plains) is very deeply cut by erosion channels, until the appearance is that of 212 BOTANICAL GAZETTE [SEPTEMBER a mountainous country. Here are found in full operation those agencies which have worn so much of the surface elsewhere from a higher to an approximately level lower plane. The Cretaceous limestones, which on the grass plains portion of the Edwards plateau lie still unbroken, have here been cut through 2.— Juniperus sabinioides formation; a prominent feature in the xerophytic Fic, timber region of the Edwards plateau province. and exposed on all sides, thus not only offering lines of penetra- tion for tree roots, but moisture facilities not available from an unbroken level surface. By this erosion, also have come about unequal conditions as between the summits and slopes of the hills, and the gorges and cafions between them. Certain Creta- ceous formations, friable on exposure to weathering, outcrop on the hill slopes or summits, thus exposing an unstable founda- tion for vegetation. On these outcrops, often entirely bare, certain very hardy species can secure a foothold, and so anchor 1901] VEGETATION OF WESTERN TEXAS 213 this crumbling rock, even permitting a soil accumulation. The existence of similar conditions in the exposed formations in the Grand prairie results in carrying the timber vegetation over the hills and bluffs and bordering escarpments of this province far to the northward of the main body (fig. zz). This timber forma- tion is an open stunted forest covering, which suffers extensive interruptions, but continues in spite of this to be the dominant vegetation feature of the areas just described. On some high arid divides it becomes reduced almost to shrubby dimensions. Floristically, the chief elements of this vegetation are Lower Sonoran, of which many species range southwest into the Mexi- can mountains, or westward across the continent. Inthe western part of this province the presence of western mountain species is very marked. At the east there remains a strong element com- mon to the Austro-riparian and Carolinian zones; but, aside from these indications of relationship, the province has a strong individuality. This is well shown by the fact that Austro-riparian and Carolinian species (when not present in the cafion timber) are vicariously represented by species stamped with the climatic characteristics of the region. Perhaps the most noteworthy case is that in which Juniperus Virginiana is succeeded by Juniperus sabinoides, the latter being a most characteristic xerophyte. As to the content of the forest formation, this may vary from a pure formation of one species to a more or less uniform mix- ture of 50 or 60 per cent. of all the species. This variation is generally due to variations in soil condition. For example, a pure formation of Juniperus sabinoides may be due to an extensive exposure of very friable limestone like the Glenrose beds. In some cases the prevalence of a single species is due to the accident of its having gained the ground first. The following species formations are the most notable in this province: Juniperus sabinoides formation —This is the most important formation, partly because of its ecological relations, partly from its economic value. The formation is popularly called ‘cedar brake.” Such brakes may be almost impenetrable over many Square miles. The tree branches diffusely from the ground up, 214 BOTANICAL GAZETTE [SEPTEMBER | making a compact oval outline when standing free, and a dense interlacing of branches in close formation. Frequently at the ground the trunk itself divides into several approximately equal members. A pure formation of this juniper occurs character- istically where a friable limestone exposure lies almost bare of soil except a loose débris collected by the forest itself. Where such an outcrop occupies a slope whose angle is sufficient to make the friable rock surface unstable, an open cedar forma- tion is the only woody vegetation present (fig. 72). Aside from the very general use made of cedar in commercial ways, it occupies an important relation to moisture and soil conditions. Not only is the formation found to occupy areas where soil cannot accumulate except for the presence Qf this growth, but when cut or burned off it tends ‘to reoccupy the ground most vigorously and to encroach upon the other species. Quercus breviloba formation.— This is the so-called ‘‘ mountain shin oak” of central Texas. On the summits of the high divides, where a thin soil is underlaid by unbroken limestone, the timber vegetation becomes very much dwarfed. In such cases the shin oak often becomes the dominant species and sometimes the only one, torming scrub thickets known as ‘‘shinneries.” The mature plant may not exceed three or four feet in height. A familiar illustration of shin oak country occurs on the high divide between the San Gabriel and the Colorado rivers in Burnet county. Quercus Virginiana formation.—This is in reality a dwarfed live oak formation which covers lower hill slopes or flats where the thin surface covering of black soil is strewn with coarse fragments of the broken underlying limestone. Such country, from its stony character is known as “hard scrabble,” and the timber covering, often almost wholly of dwarfed live oak, is dwarfed and gnarled into a fitting counterpart of the ground structure (fig. 73). ENCROACHMENT OF THE CRETACEOUS TIMBER FORMATION.— The forest covering of the erosion area tends not only to renew 1901] VEGETATION OF WESTERN TEXAS 215 itself when removed, but it is a vigorous agressor upon previ- ously open untimbered areas. This persistence may well be turned to account in any plans looking toward the maintenance of the forest covering on the more arid hills for prevention of soil erosion and destructive floods. Fic. 13.— Quercus Virginiana (mountain form) formation; Cretaceous hills fifteen a west of Au XEROPHYTIC FOREST FORMATION OF THE MOUNTAIN SLOPES OF TRANS-PEcos TExas.—This formation is the counterpart of that just described for the erosion area of the Edwards plateau. It is a part of the forest covering of the arid mountain slopes of the southwest, including south Colorado, New Mexico, Arizona, and northern Mexico. Its area in Texas is practically limited to the upper slopes of the mountains in the great bend of the Rio Grande (above 6000 feet); to the dry upper cafions in these mountains; to the middle slopes of the Guadalupe and Davis mountains and their dry cafions. The ecological conditions of these mountain slopes are more 216 BOTANICAL GAZETTE [SEPTEMBER extreme than in the erosion area of central Texas, as shown in the following data. The annual rainfall is from 14 to 26 inches (approximate averages for each area respectively ); the evapora- tion capacity is from 80 to 60 inches; the extremes of daily temperature in the trans-Pecos mountains is much greater than in the central region. Consequently the arborescent vegetation is one of more absolute adjustment to xerophytic conditions, and the semi-mesophytic species found in the central erosion area are either wholly excluded from the trans-Pecos area or rele- gated to the watered cafions, Of the arborescent species, Pinus edulis, Juniperus pachyphloea, J. occidentalis, J. monosperma, Quercus grisea, and Q. Emoryi are the principal ones. Not all of these are confined to the mountain slopes, for in watered cafions some of them form heavy timber of large size and good quality; but on the slopes their growth is dwarfed, the wood hard, the stems gnarled and misshapen, and they stand in a scant open forma- tion. The timber formation just described extends to certain physiographic features east of the Pecos river, These are the Staked plains escarpment, the remnant buttes of the Callahan divide (at least west of the 1ooth meridian), the sand hills of the southern Staked plains, and the cafion bluffs of the Devils and Nueces rivers. In the last instance, it is very dilute, being lost in the local formation, but in the other cases it is practically the only timber. On the escarpment of the Staked plains, even as far as the northeastern border, there occur Pinus edulis, Juniperus occidentalis (in many straggling groves of small growth, according to Havard), and Quercus grisea. On the remnant buttes of the western half of central Texas there are close formations of a much dwarfed Q. grisea, in which this species attains a growth of only two to five feet, covering the ground in true chaparral fashion. Here also is found a very scattered formation of Juniperus occidentalis, which occupies also the lower mesas, becom- ing one of the chaparral species covering them. 1901] VEGETATION OF WESTERN TEXAS 217 On the sand hills of the lower Staked plains region one of the western mountain oaks (Q. undulata or Q. Gambelit) occurs in low thickets. To this region and the buttes and mesas east- ward the trans-Pecos chaparral formation extends, and such reduced growths of Quercus and Juniperus as occur here become identified with the chaparral. [ Zo be concluded. | UNIVERSITY OF TEXAS. BRIEFER ARTICLES. NOTES OF TRAVEL. VII. A TROPICAL FOREST IN CERAM. THE steamer Japara of the Netherlands India Packet Boat Com- pany had a cargo of American pipe for a new oil well which was being sunk at Boela on the northeast coast of Ceram. To unload this pipe, the steamer anchored about noon on January 18 in the open roadstead, a half mile off shore. With our glasses Mr. Lathrop and I studied for some time one of those fascinating pictures of green and brown foliage which are the charm of steamboat travel in this tropical archipelago. We made up our minds that the low flat country and the hills behind it were covered with as rich a tropical forest as we had ever seen, but neither of us was prepared to find so many novel sights and interesting things as our short stay on land revealed. Boela is a tiny clearing on the ocean edge of the forest, and not only has no harbor but its roadstead shoals so gradually that at low tide even a ship’s boat grounds an eighth of a mile from the beach. So our party, the owners of the oil concession, the two travelers, and - several native men and women, were forced to wade that distance before they reached the shore. The water was warm and our slight discomfort was quickly forgotten in the beauty or the oddity of count- less things that attracted our attention. _ Asa first surprise, the beach was covered with thousands of moving shells of all sizes and shapes. These shells moved about in the most bewildering fashion. The rotting logs and even the mangrove bushes on the shore were literally alive with them. On examination each one was found to be the home of a hermit crab. Shells even smaller thap the head of a shirt-stud were inbabited, and it was with difficulty that we found any uninhabited specimens. In our stay ashore we did not see a single hermit crab which had left its shell. To add to this bewildering impression of moving strand shells, scores of small ash-colored fish were jumping about in the liveliest manner on the sand, often many yards from the water. ‘These fish wert 218 [SEPTEMBER 1901 | BRIEFER ARTICLES 219 so quick in their movements that it was many minutes before our boy succceded in capturing one with his hands. Although I did not observe any of these animals actually perched in the branches of the mangrove, I do not doubt the statement that they often climb low bushes near the strand, using their short ventral fins for feet. Boela, the possible future oil city of the Orient, was decidedly in embryo. A few huts, made of the midribs of the sago palm and thatched with the matted leaflets of the same plant, were clustered about the landing shed and surrounded by young plantings of manihot, bananas, and a few papaya trees. At the end of the little village was a more or less comfortable bungalow in which lived an American mechanic from Pittsburg who had charge of the oil drilling machinery. Three of the most curious dogs I have ever seen greeted us as we walked up the path to the bungalow. They were crosses between the hairless Mexican dog and the “kampong” or native cur. These dogs were quite hairless except the tip of the tail, which was furnished with a brush of bright reddish-yellow hair, and the top of the head, which bristled with a stiff mop of the same color. The only cat of the place was one of the short-tailed breed which is common in this region. As we strolled into the forest along a newly-made corduroy road, gorgeous butterflies flitted quickly or soared lazily across our path in such numbers that we were reminded of some of Alfred Russell Wallace’s descriptions of his best collecting experiences in Malacca. Only twice before had we seen such quantities of these tropical insects, once on a small island in the gulf of Siam, and the last time on one of the pathways near Petropolis, Brazil. The newly fallen trees were swarm- ing with insects and covered with fungi, and the log road was bright with red and yellow mushrooms of many different species. Every time we halted to examine a rotten stump or scratch away a few fallen leaves, swarms of black mosquitoes attacked us, and worried us with the suspicion that they were injecting malarial plasmodia into our lood. The new oil well, which was soon reached, was surrounded by little pools of the dark brown liquid, and many tall forest trees about were killed by the crude oil which had flowed from the well before it was closed. The bubbling oil and gas was sputtering and hissing under the cap which had been screwed down to keepit in. Although inhab- itants of Amboina who were interested in these wells expressed great hope of their success, as yet no large quantities of oil had been 220 BOTANICAL GAZETTE [SEPTEMBER discovered. The roads which were being cut into the interior were expected to open up and determine the size of this new oil field. Near one of the oil wells stood giant forest trees, with bases buttressed by thin irregular plates which reached to a height of ten or twelve feet from the ground. On one of these trees, twisting about its gray trunk, was an epiphytic fern with dimorphic leaves, the basal ones like sau- cers closely appressed to the thick fleshy rootstock which they pro- tected from the bright sunlight and its drying action. On the same tree, while examining this fern, we disturbed a num- ber of large gray ants which had built a paper-like nest on one of the broad flat buttresses. Like a company of veterans they rushed out of the nest and ranged themselves at regular distances from each other about their home. Every ant stood upright, curled its abdomen upward between its hind legs as if ready to sting, and with waving antennae awaited the enemy. On the parchment-like walls of the nest these warriors, as they hurried out, beat a sharp tattoo which was much louder but similar to that sounded by the termite warriors on the thin wooden walls of their galleries. The newly made roadways through the forest were strewn with fruits which had fallen from the tops of the tall trees. Some of these were of the brightest orange and others a clear lemon-yellow, others still were purple and brilliant green. None, however, were edible, though many were most attractive looking. In the tree-tops overhead, bright green parrots chattered, and a small cicada with a shrill but not unpleasant note kept up an incessant racket. Once a black parrot with red head flew low through the trees, looking like a flash of crimson light. Most of the birds, however, were in this season in the tops of the highest trees. As in all tropical forests I have ever visited, the flowers in Boela were scarce. One bright crimson representative of the lily family, with a spike as large as a pineapple, was the only showy species observed. The botanical fancy is attracted in these forests by the curious creepers, which fling their supple stems about the tall forest trees and spread out their dark green tops above those of their support: ing neighbors, until the latter are robbed of sunlight and slowly die. There are curious masses of stilt-like roots from which rise tall slender trunks; the pendant barbed tips of the rattan palm catch and hold you; delicate masses of green filmy ferns form soft pads on the fallen trunks ; dusky chocolate-brown masses of the myxomycete (Stemonitis) 1901 | BRIEFER ARTICLES 227 color your fingers as you touch them; ugly looking incipient Polyporei exude oily looking drops of brown fluid; and the rotting bark con- tains a host of microscopic fungi. Epiphyllous algae and lichens are always abundant in these dark forests and are often mimicked by insects. However little one may be interested in the insects of temperate regions, the constant buzzing of flies and wasps, and the fluttering of gor- geous butterflies in this virgin forest soon attract one, and leave a lasting impression of the activity of the animal world as compared with the world of plants. ‘The ants alone, in their host of forms and curious habits, aré sufficient to distract the most obstinate collector, and rob him of many hours he would have spent in adding new numbers to his herbarium. ‘They are the most intelligent beings of the forest, and the solitary traveler finds a sort of intellectual companionship in watching theirmovements. The termites of the tropics are second only to the tree ants in interest. At Boela only the tree-inhabiting forms were found, and none of those curious kinds were seen which cultivate fungi in their nests. The dead stumps and trees were every- where alive with them, and the decaying trees were rapidly removed by immense colonies of these wood-eaters. The virgin forests of the tropics would be well nigh impassible to collectors if the fallen branches and logs were not quickly riddled with the galleries of the white ants. They are above ground what the earthworm is below it. They reduce all dead wood to half-digested fragments. This portion of Ceram is very sparsely settled, so sparsely, in fact, that during the whole time we spent at Boela we saw only two or three native Alfurians, who came off to the steamer in their small dug-out canoes to sell their large but coarse-grained bananas. They are a darker, more uncleanly race than the Javanese or Amboinese, and show traces of Papuan origin. Imported coolies from Java and Amboina are depended upon to work the drills at the oil wells. But all idea of time was lost as we wandered through this forest, until the rapidly fading light (to say nothing of the rapidly devouring mosquitoes) gave warning that a hurried retreat must be made from this malaria-haunted shore. In the morning we saw Boela again, and it looked even more beautiful gilded by the rising sun than in the evening light.— Davin G. Faircuitp, Department of Agriculture, Washington, D. C. CURRGEN CoLIPERATURE. BOOK REVIEWS. The sea beach. WE FANCY that there is a demand for a popular account of the seaweeds. It is not so important that the names and classification should be given, for such may be found in several well-written manuals and texts. What is prob- ably most desired is a simple account of the life habits, and in part the life histories of such forms as are prominent in the tide pools, on rocky shores, in marshes, and in flats, and those that are washed up from the deeper waters after storms. A book by Augusta Foote Arnold’ attempts to satisfy the thirst for seaside lore, and this review will discuss some points in her treat- ment of the marine algae. They are given only 50 pages in a book some 500 pages long, the remainder of the work dealing with the marine inverte- brates, so it would be unjust to judge The sea-beach at ebb tide as a whole by the criticisms presented here. It is hard to understand precisely the attitude of this author as regards the relation of popular science to technical science, or to put matters some- what differently, the balance between the charm of natural history and the detail of a fully developed science. There is a little of both in this book, the science badly mutilated and sometimes incorrect, the natural history quite lacking the delicate fanciful touches that demand an imagination and yet must be tempered by many years of intimate contact with nature and by knowledge of scientific exactness and respect for it There is a brief account of the life habits, aeitcitvueiaii: and uses of algae, and some statements about naming and classification which would lead the reader to suppose that especial emphasis would be laid upon these depart- ments. And this is the case, for most of the space is devoted to the descrip- tion and illustration of genera and species. These are apparently arranged after the system of a standard text, much abridged of course, the outline being presented at the beginning of each group. These synopses can be nothing but a mere dictionary-like table of names, for the account does not pretend to give the morphological characters of the groups. There are no keys, no microscopical examination is required, and for identificaticn the reader (presumably beginner) must depend chiefly upon the figures. What * ARNOLD, AUGUSTA Foors: The sea-beach at ebb-tide. A guide to the study of the seaweeds and the lower animal life found between tide-marks. Small 8vo. pp. xiit+490. figs. 600+. New York: The Century Co., 1901 222 [SEPTEMBER Baath AD 1901] CURRENT LITERATURE 223 then can be the value of this classification, out of sympathy with the purposes of morphology on the one hand and the spirit of natura] history on the other? The illustrations are photographs of mounted algae on cards, and in some cases of preparations slightly magnified. While many are clear others seem to the writer quite valueless as a means of identification and unworthy of the book. Although the color of the specimen would help in this mechanical matching of mounts with figures, nevertheless there is sure to be much con- fusion. For example, how is Enteromorpha clathrata to be distinguished from certain Cladophoras? The figure of Ectocarpus viridis might do for several other species, the Callithamnia are quite impossible, and Polysiphonia fastigiata is certain to be confused with Sphacelaria. One occasionally finds statements that lead to the belief that the author is quite untrained in natural history. Thus, on page 29 is the — “Ectocarpus Hooperi, a species of Ectocarpus ie described by Mr. Hooper.” This does not seem to be a fact, and the impression conveyed that naturalists name species after themselves is an implication of conceit far from being warranted by the conduct of these modest members of society. The book has yet to be written that will tell the natural history of sea- weeds with the charm of manner shown inthe style of Miss Margaret Morley. And until such a treatment appears, it is much better that the amateur col- lector and observer of marine algae read Murray's /ntroduction to the study of sea-weeds, a simple and very interesting account, and one thoroughly grounded in science.— B, M. Davis NOTES FOR STUDENTS. PROFESSOR ARNOLDI? has taken up the somewhat incomplete work of Shaw, and has made a careful study of the development of the endosperm of Sequoia sempervirens. Free nuclear division takes place in the usual manner in an evenly distributed peripheral layer of protoplasm, but very soon there is a denser accumulation of protoplasm at the lower end of the sac. When the formation of walls begins, three regions of the endosperm may be dis- tinguished, the upper, the lower, and the middle. The upper, and particu- larly the lower, develop faster than the middle, so that the ends of the sac become filled with a solid tissue while the nuclei are still almost free in the middle portion. Each nucleus of the middle portion now becomes sur- rounded by a wall which is open on the inner side: the walls grow inward and when the center is reached, walls are formed at the inner ends of the cells. The nucleus now begins to divide, and each of these cells (“alveoli”) becomes divided into several cells. Archegonia are formed only from these *ARNOLDI, W.: Beitrage zur Morphologie einiger Gymnospermen. I. Die Ent- wickelung des Endosperms bei Seguota sempervirens. Bull. des Natur. de Moscow. Pp. 1-13. Pls. 7-8. 1890. 224 BOTANICAL GAZETTE [SEPTEMBER alveolar cells of the middle region. At the time of fertilization, the upper and lower portions of the endosperm consist of small-celled tissue, while the middle portion is alveolar. Sequoia is regarded as a connecting link between Gnetum and the angiosperms on the one hand and between gymno- sperms and the archegoniates on the other. In a later paper 3 he has described the archegonia and pollen tubes of the same species. The archegonia are very large, and some sections show as many as sixty. They sometimes occur singly, but are often grouped. In development they resemble the archegonia of the Cupressineae, since they are often in direct contact with each other and do not form any ventral canal cell. There are no proteid vacuoles. The neck consists of two cells, in this respect resembling the older gymnosperms. e pollen tube grows through the nucellus, not between the nucellus and integument, as described by Shaw. At the time of fertilization the pollen tube contains the two male cells of equal size, and two small nuclei, one of which is the tube nucleus and the other “the nucleus of the cell which united the generative cell with the microspore wall.” The general structure of the pollen tube and its contents agrees with the Cupressineae. ‘The morphological considerations, together with the geographical distribution, lead to the conclusion that Sequoia is nearly related to the ancient type from which the modern Araucarias and Cupressineae have descended. CHARLES J. CHAMBERLAIN. CONNECTING THREADS which establish protoplasmic continuity between adjacent cells have been studied by Mr. Hill‘ in the embryo and seedlings of Pinus pinea and in the mature tissues of P. si/vestris, Some attention was also paid to the endosperm of P. pizea. The endosperm consists chiefly of rather large rounded cells, but a close examination shows that in many cases an internal division has occurred. The threads are evenly distributed in the young walls, but are grouped in the older walls. Near the cotyledons the cells are smaller, the threads thicker, and there are traces of ferment action. Ferments from the cotyledon pass into the endosperm through the threads, and by the same route food materials pass from the endosperm to the embryo. In the seedling the absorptive side of the cotyledon is more abundantly supplied with threads than the side not exposed to the endosperm. There ee W.: Beitrage zur Morphologie und Entwickelungsgeschichte einiger spermen, II. Ueber die Corpuscula und Pollenschliauche bei Seguota semper P 9 ae Bull. des Natur. de Moscow. ls. 10-11. LL, A. W.: The distribution and character of connecting threads in the rag a Pinus silvestris and other allied species. Phil. Trans. Roy. Soc . London B. 194 : 83-125. pls. 37- | 2 Igor | CURRENT LITERATURE 225 are no threads in the external walls of the epidermis, and but very few con- necting the guard cells with their neighbors. All parenchyma cells show a general resemblance in the character of their threads, the threads on the end walls being irregularly scattered, while _ on the side walls they are grouped. In the phloem all the sieve tube threads show a characteristic median dot. The albuminous cells at the edge of the phloem of the leaf have their threads grouped in localized thickenings on the walls, and serve to pass materials from the mesophyll to the phloem. The very numerous threads of the root cap form a connection with the free surface of the root and with the periblem. In the mature tissue of /. s¢/vestris the threads in the cortical tissue are similar to those of the seedling. In the phloem there is no connection between the sieve tubes and the bast parenchyma or the starch medullary ray cells. The sieve tube threads on the radial walls have a median dot. The torus of the bordered pit is probably traversed by threads which soon disappear. In the leaf, the distribution is about the same as in the cotyledon. The endodermis, with very numerous threads, is in close connection with the cortex and the stele. In the pericycle, living cells are connected by threads, but there is no connection between the pericycle and the lignified transfusion tissue. In general, the main direction of threads in the cortex and phloem is tangential. The transitory nature of certain threads explains the absence of threads between the sieve tubes and medullary ray cells. Except in the medullary rays and in the cork cambium, the threads are chiefly on the radial walls. This suggests that in conifers food supplies and stimuli are conducted mostly in a tangential and vertical direction.—CHartes J. CHAMBERLAIN, Teme. Leo, A CORRECTION. IN the June number of the BoTANICAL GAZETTE (31: 441) there was published by Ernst A. Bessey, of the U.S. Department of Agriculture, a review of Bulletin 49 of the Oklahoma Agricultural Experiment Station entitled 4 rhizomorphic root-rot of fruit trees. The reviewer made the state- ment that ‘‘ This has been shown by von Schrenk and others to be caused, - all probability, by a hitherto unidentified rhizomorph-producing fungos. The reviewer having failed, on request, to cite references to these publications of “von Schrenk and others” in support of his statements, and having thought it ‘inadvisable’ to correct the same when asked to do 80, the fol- lowing brief statements are submitted for the information of those apie November 6, 1889, von Schrenk identified the fungus in question as ‘one of the most dangerous tree fungi known, Agaricus melleus.”’ (Letter bas Horticulturist of the Okl. Agr. Exp. Sta.) Some more of the same material was submitted to Dr. B. T. Galloway, and under date of November 10, 1899, Dr. Galloway stated that “the fungus is the edible mushroom, Armillaria mellea.” (Letter to Horticulturist Okla. Agr. Exp. Sta.) In the Abate. Judd Farmer for January 12, 1901, von Schrenk says, “I suspect that possibly common in both Europe and America, the Agaricus melleus, - honey mushroom.” This last statement is quoted in the Budletin of this Station above referred to. These statements will show clearly the state of the investigations of “von Schrenk and others” at the time stated. Perhaps in the future we shall have from these sources some valuable publications regarding this subject, pe present none such exists to my knowledge. It is hoped the above sere tions will clear away some misconceptions and prevent the inference gc this disease had already been investigated by members of the staff of the Department of Agriculture. E. MEAD WILCOX. OKLAHOMA AGr. Exp. STa., Stillwater, 226 [ SEPTEMBER No Wo. AT THE recent Denver meeting of the A. A. A. S. Dr. D. T. MacDougal was elected general secretary for the ensuing year. HE University of Glasgow, at its ninth jubilee, celebrated in June last, aes the degree LL.D. upon Professor W. G. Farlow, of Harvard University. PROFESSOR F. O. Bower of the University of Glasgow is one of three representatives appointed to attend the bicentennial celebrations of Yale University next October. THE University of Chicago at its last Convocation conferred upon Mr. A. A. Lawson and Miss Florence May Lyon the degree Ph.D. Miss Lyon has been appointed associate in botany and Head of Beecher Hall in the University of Chicago M. EUGENE AUTRAN has severed his connection with the Boissier herba- rium at Geneva, Switzerland, and has been appointed botanist to the botan- ical garden of Buenos Ayres. He is also a member of the staff of the botanical section of the een: Department of Agriculture. THE FOLLOWING RESOLUTION was adopted by section G of the A. A. A. S. at the recent Denver meetin ng: Resolved: That it is the sense of this section that it would be advisable to estab- offer most valuable opportunities for the prosecution of investigations in nearly all branches of botanical science, and would do much to supplement the facilities already offered by American institutions. Extended economic experiments in the tropics must rest more or less directly upon purely botanical research, and the establishment of such a laboratory would do much to strengthen the efficiency of the sveagaege Station. This resolution is not to be taken to mean that the research station should be placed in the same building or buildings with the oe station, but should be located at the point most favorable for the work in questio Dr. CHARLES Monr, the venerable botanist, for many years a resident of Mobile, Alabama, died at his home at Asheville, N. C.,on July 17. Dr. Mohr was for some years a special agent of the Forestry Division of the U.S. Department of Agriculture, for which he prepa ared a monograph on The timber pines of the southern United States, published in 1896. His most recent work is Plant life of Alabama, the result of exploration and study of T9o1] 227 228 BOTANICAL GAZETTE [ SEPTEMBER the flora for forty years. This volume, which has been nearly three years in the government press, was issued on July 31, unhappily too late for the author to see the full fruition of his labors. This work we shall review later. We learn from Science that he had in preparation a volume on the eco- nomic botany of Alabama, which is probably too incomplete for publication. He published some years ago a pamphlet on the Botanical resources of Ala- bana. . THE FOLLOWING PAPERS were read before the Botanical Society of America at the recent Denver meeting: J. C. ARTHUR, Clues to relationship among heteroecious plant rusts; W. J. BEAL, Some of the changes now tak- ing place in a forest of oak openings; C. E. Bessey, Early winter colors of plant formations upon the great plains; E.G. Britron and A. TAYLOR, The life history of Vittaria lineata; F. E. CLEMENTS, The fundamental phe- nomena of vegetation, The plant formations of the Rocky mountains (with lantern slides), and A system of nomenclature for phytogeography; W. A. MurriLt (by invitation), The anatomy of the embryo and seedling of 7suga Canadensis ; B. L. ROBINSON (address of retiring president), Problems and possibilities of systematic botany; Ww. TRELEASE, A suggested hybrid origin of Yucca gloriosa (with lantern slides). : The officers elected for the ensuing year are J. C. ARTHUR, president, B. T. GALLOWAY, vice President, D. T. MACDOUGAL, secretary ;, A. HOLLICK, treasurer, C. E, BESsEY and WM. TRELEASE, councilors. THE FOLLOWING PAPERS were read before section G of the A. A. A. 5. at the recent Denver meeting: W. J. BEAL, Lantern views of the Botanical garden at the Agricultural college of Michigan; C. E. Bessey, The morphol- ogy of the pine cone; ALICE Eastwoop, General botanical features of the Coast mountains of California; B. D. HaLstEep, Notes upon colors of salsify hybrids; A. C. LEwis, Contribution to the knowledge of the physiology of karyokinesis ; D. T. MacDouGAL, Thermal relations of plants, and Compara- tive climate of a meadow and a hemlock forest; AVEN NELSON, Some ape of the Wyoming desert flora; L. H. PAMMEL, The xerophytic vegetalon of the Uintah mountains; F. RAMALEy, Observations on £gregia Mensa and the plants of the eastern foothills (with lantern slides); A. D. SELEY Experiments with lime and solutions of formaldehyde in the prevention of onion smut, and germination of seeds of some common cultivated plants after prolonged immersion in liquid air; E. E. SLOSSON, Effect of salt . . ic solutions on seeds and plants; Wm. 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Ver, $0.50 1 FOR SALE BY BOOKDEALERS OR BY THE PUBLISHERS THE UNIVERSITY OF CHICAGO PRESS, Chicago, Illinois a neeepsnsteepainasghsiemtenuneanioe YOUNG MEN you are advised to Investigate Endowment Life Insurance It enables you to save money regularly You are absolutely certain of full returns for the money invested. Those dependent upon you are certain of protection in event of your death FILL OUT THIS SLIP AND SEND TO US Without committing myself to any action, I shall be glad to receive, free, particulars and rates of Endowment policies. EEE pe ee, Hale ee ee ee Name sesh aout oer peter cil apie, —_* ma ak pen ese Sts Addre. ey er Me ae ee Octeipation—— ai cA Sar Nica he RE ame a oe SUS gee sea Dept. 25 THE PRUDENTIAL INSURANCE COMPANY [29 OF pay AMERICA JOHN F. DRYDEN, President HOME OFFICE: Newark, N., J. a Se co eee NS = heme eh SSB a VPA AS eRe VOLUME XXXII eile NUMBER 4 BOTANICAL .GAZear OCTOBER, roor THE APPLICATION OF NORMAL SOLUTIONS TO BIOLOGICAL PROBLEMS. JAMES B. DANDENO. Jupcinc from recent investigations into the effects of ions upon plant and animal life, one is led to conclude that there is much confusion in regard to the interpretation of certain stand- ard chemical solutions. It is not to be doubted that solutions have a very important part in biological functions, and because of their importance it is deemed advisable to attempt to place this branch of an important subject upon a firmer basis. Since solutions of molecular concentration have been adopted for comparison, in place of the older percentage concentration solutions, there seems to be a great tendency to misinterpretation and confusion. Solutions prepared on the percentage basis are not now considered scientifically accurate when used for pur- poses of comparison, while those molecularly equivalent may be compared with scientific accuracy. The molecular and the equivalent solutions are the only ones here discussed. Much of the physiological work done with molecular solu- tions proves upon examination to be faulty because of a miscon- ception of the meaning of xormal solutions, gram-equivalent per liter solutions, and gram-molecule per liter solutions. There are no less than three interpretations of what a normal solution is. One takes it to be a gram-molecule per liter of solution. Another takes it to be a gram-equivalent per liter of solution. Still another regards it as a gram-molecule or a gram-equivalent per liter of 229 239 BOTANICAL GAZETTE [OCTOBER water. Some use the symbol x (meaning in chemistry a nor- mal solution) and say nothing about the method of prepara- tion, | There are two probable causes for this confusion. The ana- lytic chemists have differed in their use of solutions, (Allen, Chem. News 40: 239; also Analyst 13: 181, cited by Sutton 18," p. 28.) Some (a very few) German chemists use a gram- molecule per liter and call it a normal solution. By far the greater number of modern analytical chemists, however, use a gram-equivalent per liter of solution as a normal solution, The physical chemist uses both the above mentioned methods of preparation, but is always clear as to the meaning, and calls only the gram-equivalent per liter the normal solution. Those who use the chemical symbol without explanation fall under some shadow of doubt arising from the above mentioned con- fusion. The work, however, may be faultless and the interpre- tation accurate. The theory of the separation, in aqueous solu- tions, of the molecules of compounds into ions is now a generally accepted one, and it is partly because of the actions of these ions, and of the dissociation of the molecule into ions, that there is great need of uniformity and of chemical accuracy in the preparation of solutions. Here are some definitions of normal solutions. 1. Fresenius (2, p. 687): ‘Solutions of such strength that 1000“ contain an amount of acid or base equivalent to one gram of hydrogen are normal solutions, e. ¢., HCl, mol. wt. 36.46, wt. in 1000° of solution 36.46 eg. Oey « “ “ Na,CO, © Toast, oe: “4 #75304" Fresenius’ chemical table (p. 846) is based on the above men- tioned method of preparation. 2. SUTTON (18, p. 28): ‘Normal solutions are prepared so that one liter of solution at 16° C. shall contain the hydrogen equivalent of the active reagent in grams (H =1),” ¢.g-° * See bibliography at end. 1901 | NORMAL SOLUTIONS 231 HCl I mol. wt. in grams per liter of solution H,SO, 4 e . * : Na,PO, + sd a “ KMnO, 4+ " - v = Kz Ce, 0, + ‘ , " “(Talbot 19, p.65) 3. TALBor (19, p. 64): “A normal solution as defined by Mohr (Talbot 19, pp. 64, 65) contains in one liter ‘one equiva- lent of the active reagent in grams.’ The equivalent in grams may be defined as ‘that quantity of the active reagent which contains, replaces, unites with, or in any way, directly or indi- rectly, brings into reaction one gram of hydrogen.’” Mitter and Kivikani (11, p. 22), state regarding normality of solutions that ‘like volumes are equivalent to one another.” On the other side of the question, however, there are a few who use gram molecule per liter solutions and call them normal solutions. Of these, two perhaps are worth mentioning, Muter and Menschutkins, both cited by Sutton (18, p. 28). In Ostwald’s own work (13) he uses both gram-equivalent per liter and gram-molecule per liter of solution, but never con- fuses them. He uses the gram-molecule solutions for demon- strating molecular conductivity, and some of these tables are to be found in his Lehrbuch (2, pp. 722-772). Ostwald (15, p. 281) gives Kohlrausch’s (6) table of equivalent quantities (not molecular quantities) for several solutions and enunciates the principle that the equivalent conductivity is a direct measure of the velocity of the migration of the ions. Ostwald (14, p. 284) uses normal solutions (with table) in regard to surface-tension of solutions, and on pp. 170-172 he uses molecular solutions with table of reference for molecular lowering of the vapor-pressure. There are only two instances of different tables used by Ostwald in his work. It will thus be seen that he makes use of both kinds cf solutions according as they best suit his particular pur- pose. Van’t Hoff (23, p. 117), referring to Ostwald, uses the term ‘‘so-called molecular conductivity,” and he uses freely the term normality as applied to solutions. Hittorf’s table ( Le Blanc 7, 232 BOTANICAL GAZETTE [OCTOBER p. 78), is based on the gvam-equivalent per liter solution, and Le Blanc (7, p. 79), uses the symbol A to designate eguzvalent con- ductivity as distinguished from molecular conductivity, which is expressed by Ostwald as #. Moreover, Valson’s (24) law of the moduli (tables) is based upon gram-equivalent (not molecular) solutions, as well as almost all the analytical chemical tables now available. It is seen that the physical chemist recognizes two kinds of standard solutions, molecular solutions and equivalent solu- tions, calling the latter zormal solutions. The analytical chem-~ ist uses one kind of solution, the normal solution, z. ¢., the gram-equivalent per liter solution. The mistakes resulting from misinterpretation of normal solutions, and from a confusion of the terms gram-equivalent and gram-molecule per liter will be better understood if a few partic- ular cases be cited. Kahlenberg and True (5, p. 85), speak thus: ‘Chemically equivalent quantities (z. ¢., molecular quan- tities) of the different substances were not compared ;” also (p. 91), ‘expressed in gram-molecules or gram-equivalents per liter ;” showing that they regard a gram-molecule per liter as exactly the same as a gram-equivalent per liter, which, in the case of many of the substances referred to (¢.g., H,5O,), is not correct. What brings out more prominently the importance of the error is the fact that the solution of H,SO, (p. 92) was 4 normal solution (purchased from the chemist), that is a gram- equivalent solution, and was thought to be a gram-molecule solution. Their results show that 4255 gram-equivalent per liter of H,SO, is as toxic as y,!5,; gram-equivalent per liter of HCl. In other words, the experiments seem to prove that the solution of H,SO, was what they shought it to be, though what zt was not. Heald (4, p. 125), referring to Kahlenberg and True, states that ‘‘in these experiments the solutions were prepared according to gram-equivalents.”” On pages 119-123 of Kahlenberg and True we find written gram-mol. per liter. Similar mistakes were made by True (20, p. 184), where he says, referring to H,SO,, “and since it splits off two hydrogen | 1901 | NORMAL SOLUTIONS 233 ions from every molecule, it would have, in chemically equiva- lent quantities, twice the number of H ions found in HCl, and would have its death-limit at one half the concentration of the monobasic acid.” This shows clearly that this author regarded chemical equivalent and molecular solutions as the same. Kahlenberg and True (5, p. 6), say: ‘In the second column’ is the concentration just allowing growth, expressed in fractions of a gram-equivalent per liter of water.”” In the column referred to, the limit for H,SO, is 2,5, and for KHSO, is goa in equivalent solutions. Since the toxicity of potassium and of SO, at this concentration may be neglected (p. 6), the hydro- gen in the KHSO,, is as toxic as ¢wice as much hydrogen in the case of H,SO,, since there is in chemically equivalent quantities of H,SO, and KHSO, twice as much hydrogen in H,SO, as there is in the KHSO,. Another instance of the error arising from confusing the gram- equivalent per liter with the gram-molecule per liter is shown in the work of Kahlenberg and True (5, p. 109). They say, referring to Ostwald (16), ‘the most dilute solutions with which he worked contained one gram-equivalent in 1024 liters.’’ Ostwald’s tables in this reference are zof made on the gram-equivalent per liter plan, but on the gram-molecule per liter plan, and he so states (p. 174). ‘Die folgende Spaite enthalt unter ‘7’ den Wert der relativen molekularen Leitfahigkeit; der Wert von yu,,, auf welchen derselbe bezogen ist, findet sich in der Ueberschrift der Tabelle angegeben.” Also p. 171, ‘Wo p, die molekulare Leit- fahigkeit bei der Verdiinnung v (in Litern auf in Grammo- lekulargewicht), M,, den Grenzwert derselben bei unendlicher Verdiinnung und ¢ eine Konstante bedeutet. Driickt man die molekulare Leitfahigkeit in Bruchtheilen ihres Grenzwertes aus, setzt also m=/* .” i.e) Kahlenberg and True (5, p. 115) give in their tables 56, 57, 58, 59, “concentration gm.-equival. per liter,” and refer to Ost- wald (16, p. 380). Now Ostwald’s tables (pp. 369-422) are made zof on the plan gram-equivalent per liter but gram-molecule 234 BOTANICAL GAZETTE [OCTOBER per liter. No comment is necessary. Again Kahlenberg and True (5, p. 116), state, ‘‘as maleic acid at the dilution 1024 is dissociated 98.2 per cent. and fumaric acid 78.5 per cent., we should expect the latter to be less poisonous than the former if the toxic action be due to the H ions alone.” In the table to which they refer we find maleic acid 92.8 per cent. at 1024 and fumaric acid 78.5 at 20g8. They state further, referring to the two acids mentioned, ‘‘we do not place much reliance on the results obtained from these two acids as it is questionable whether the substances were perfectly pure.” The discrepancy above referred to suggests another reason for unreliability of results. True and Hunkel (22, p. 326), while using gram-molecule per liter in their paper, made reference to Kahlenberg and True (5, p. 92), where the tables are written gram-equivalent per liter. The one kind of table, of course, cannot apply directly to the other. In regard to the method of preparation of solutions by dis- solving a gram-equivalent of the salt, or a gram-molecule of the salt, as the case may be, in a liter of water, there is something to be said. Those who have made errors in this way are Pfeffer (17, p. 146), Detmer and Moor (1, p. 326), Kahlenberg and True (5, pp. 85, 87), True (20, pp. 184, 185), True (21, pp. 419, 411), Heald (4, p. 133), Garry (3, p. 298), True and Hunkel (22, p. 326), and Ostwald (14, p. 190). Pfeffer, Detmer and Moor, and Garry called them zormal solutions. Kahlenberg and True called them gram-molecule solutions and gram-equivalent solutions, apparently interchangeably, but meaning generally molecular solutions. Ostwald is using Tammann’s table which States that the solutions are made by dissolving ~ molecular weights, in grams, of the salts in 1000 grams of water.’ The objection to the above solutions is not so much to the *MacDovaat (10, p. §1) says: “A normal solution of ethyl alcohol is made by adding 46 grams of absolute alcohol to a liter of distilled water.” This, to be in any sense accurate, should read (after the word alcohol) fo sufficient water to make a liter of solution. MacDougal’s statement produces an error of about 58° in 1000%. a A i | b| / ; Igor] NORMAL SOLUTIONS 235 method of preparing them as to the application to them of chem- ical or physical tables prepared on a different basis; and to the error arising from their being called by names which mean some- thing different. A solution prepared by dissolving one gram- equivalent in a liter of water is quite different in concentration (in the case of strong solutions sapeckany) from that prepared by dissolving one gram lent in a liter of solution. The dif- ference is so apparent that explanation is unnecessary. The use of the chemical symbol z to represent solutions with- out some explanation as to the method of preparation of the solution, is likely to cause some doubt to be cast upon the work. One will naturally infer that those who use the symbol with- out explanation are as likely to err as those who use it with explanation, but in making explanation show an error of inter- pretation; though this inference might be unfair. Garry (3, p. 298) states: “To designate chemically equiva- lent solutions, the chemist uses fractional parts of the so-called normal solutions, z. e., the solution made by dissolving the equiv- alent gram-molecule in one liter of water.’ He uses the symbol N to designate his solutions, but if he prepared them after the method indicted above he is not warranted in using the symbol for normal solutions. Chemical tables showing degree of disso- ciation will, of course, not apply to solutions so made. True (20, p. 186) uses the term m-1024 in reference to Ost- wald’s tables (16), but he is not warranted in using this term because Ostwald’s tables just referred to are not based on normal solutions but are molecular solutions. Heald (4, p. 138) uses the term Vand refers to the same page of Zeitschrift for degree of dissociation. These tables do not apply. In the works of Loeb (9), D. J. Lingle (8), and A. Moore (12) the symbol z is used to designate the solutions, but no explanation is given, though in the context there are indications that the symbol z is properly applied. The results of the errors and misconceptions just referred to may be much or little according as the points brought out are of little or of much consequence. At all events, there seems need 226. BOTANICAL GAZETTE | OCTOBER of establishing this part of physiology upon a more secure and accurate basis. The analytical chemists and the physical chem- ists have done an enormous amount of investigation upon solu- tions, and they have established tables for various purposes. Their tables and results are convenient, clear, and ready for use ; and the only way to make proper use of them is to make the solutions according to the chemist’s standard. Sutton (18, p. 28, footnote) gives a piece of excellent advice in the following statement: ‘Anyhow it is to be hoped that those who communicate processes to the chemical journals, or abstracts of foreign articles for publication, will take care to distinguish between the conflicting systems.” If this advice is useful to the chemist, how much more useful should it be to those who are not chemists, but who in their work make use of tables of standard solutions made by the chemists. BoTANICAL MUSEUM OF HARVARD UNIVERSITY. BIBLIOGRAPHY CITED. 1. DETMER and Moor: Plant Physiology. 18098. 2. FRESENIUS: Volumetric Chem. Anal. 18 3. GAkRY: The effects of ions upon the Ritewation of flagellates. Amer. Jour. Physiol. 3: 298. 1900. HEALD: On the toxic effect of dilute solutions of acids and salts upon plants. Bot Gaz, 22: 125. 18096. : KAHLENBERG and TRUE: On the toxic action of dissolved salts and their electrolytic dissociation. Bot. Gaz. 22:81. 1896. . KoHLRauscH and HoLporn: Das Leitvermdgen der Elektrolyte. 1898. 7- LEBLANC: Electro-chemistry. 1896. 8. LINGLE: The action of certain ions on ventricular muscle. Amer. Jour. Physiol. 4: 265. 1900. : 9. LoEB: On ion-proteid compounds and their réle in the mechanics of life > on . a rr. MILLER and KILIKANI: Anal. Chem 12. Moore: Further evidence of the rater ‘effects of a pure NaCl solu- tion. Amer. Jour. Physiol. 4: 386. 13. OSTWALD: Lehrb. d. Allg. Chem. II. ee 2. “1893. : Solutions (trans. by Muir), 1891. : Outlines of Chemistry (trans. by Walker). 1895. 1901} NORMAL SOLUTIONS 237 16. OSTWALD: Zeitschrift f. Phys. Chem. 3: 170, 241, 369. 1899. 17. PFEFFER: Plant Physiol. (trans. by Ewart). 1900. 18. SUTTON: Volumetric Analysis, ed. 7. 1 19. TALBOT: Introduction to Quantitat. Chem. Anal. 1899. 20. TRUE: The toxic action of a series of acids and is their sodium salts on Lupinus albus. Amer. Jour. Science. 9: 183. : Physiological action of certain Seeculele pen. Bot. Gaz. 20-5 407.01 22. TRUE and Riacar: The eues effect exerted on living plants by phenols. Bot. Cent. 76: I 23. VAN’T HoFF: Lectures on ane Thtee I. 1900. 24. VALSON: Compt. Rend. 73: 441. 1874. 2I. GAMETOGENESIS AND FERTILIZATION IN ALBUGO. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. XXIX FRANK LINCOLN STEVENS. (WITH PLATES I-IV) [Concluded from p. 169.) Ill. GENERAL CONSIDERATIONS, OBSERVATIONAL evidence that kinoplasm and trophoplasm (Strasburger 1892) are true morphological elements of the cell has steadily increased, and striking experimental evidence has recently been adduced to confirm this conception (Hottes, ined.). In 1892 Strasburger proposed a theory of fertilization founded upon the assumption that sexual cells are incapable of develop- ment, owing toa kinoplasmic starvation. This theory was based partly upon observations on Ulothrix, and the relation of the cilia in sexual and asexual cells. Much confirmatory evidence has since been received, and in Strasburger’s latest book (1900) the theory is developed more completely. As applied to sex- ually differentiated cells, the theory postulates kinoplasmic hun- ger in the female and trophoplasmic hunger in the male. The phenomena attending zonation in Albugo and Peronospora are capable of explanation in the light of Strasburger’s theory, and in turn materially strengthen the theory itself. In Albugo and Peronospora the marshaling of the nuclei into a hollow sphere, a most conspicuous phenomenon, is quite inexplicable on the ground of atavism or phylogeny, nor can it have to do with wall- building. Why then do the nuclei habitually leave the ooplasm, apparently to perform no useful function in the periplasm, only to return and function as female pronuclei ? A study of the accompanying plates shows plainly that the periplasm is of a distinctly different character from the ooplasm. 238 [OCTOBER Igor | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 239 The latter is typically very dense and alveolar, does not stain with gentian-violet but takes the orange G. The former stains darkly with the gentian-violet, and is filar, not alveolar. The processes leading to zonation may indeed be characterized pro- visionally as a differentiating of the oogonium into two regions, a periplasm rich in kinoplasm, and an oosphere rich in tropho- plasm. This statement is borne out by all the positive charac- ters of the trophoplasm and by the stain reaction and structure of the kinoplasm. The nuclei are in mitosis and do not lose their membrane until zonation is sharp, nor do they reenter the ooplasm until their membrane is lost. The kinoplasmic nuclear membrane is apparently left in the periplasm, and its absence is evident during the second mitosis, thus resulting in that marked differ- ence in character between the first and the second mitosis, which is illustrated in the figures, a difference which was noted in A. Buti in my earlier paper (1899, p. 231). The nuclear membrane in the second mitosis is very thin or perhaps absent; the achro- matic figure is weak and consequently often distorted and irreg- ular. Every indication is that of an absence of kinoplasm. Thus the behavior of both oogonial nuclei and cytoplasm confirms in a striking way the view that kinoplasm is important in sexual differentiation, and suggests that the nuclei pass to the periphery to rid themselves of superfluous kinoplasm, possibly to prevent parthenogenetic development in the oosphere. If this be the true reason for the migration of the nuclei, it logically follows that kinoplasm is not readily convertible into tro- Phoplasm, at least not in the conditions that prevail in these oospheres. In the antheridium a behavior complementary to that exhib- ited in the oogonium is seen. The antheridial protoplasm stains intensely with the gentian-violet before fertilization, but after fertilization the cytoplasm left in the antheridium fails to give this reaction. The antheridial nuclei as they lie in the tube pos- sess a heavy membrane and stain darkly, giving every indication that they are rich in kinoplasm. In A. candida zonation is not 240 BOTANICAL GAZETTE [OCTOBER so marked, and the nuclei do not pass to the periplasm during division. In this species, however, there is a great preponder- ance of trophoplasm, owing to the highly developed coenocen- trum, as well as to the fact that the trophoplasm of the whole oosphere is surrendered to one nucleus. In general, the phenomena of oogenesis and spermatogenesis in Albugo afford remarkable confirmatory evidence for Stras- burger’s theory of fertilization. If the definite establishment of this theory should occur, botanists will come into more accord with those zoologists who accept the theory of Boveri that the sperm contributes the centrosome (kinoplasm), which is the one element needed by the egg to restore its capacity for division. Possibly the results of Klebs (1896), Loeb (1899), Nathansohn (1900), and others who have artificially induced parthenogenesis, may be explicable in the light of this theory, since it is at least conceivable that the environmental conditions which are sup- plied in these experiments may be identical with those which favor or retard the development of kinoplasm in the cell. Indeed, the results already attained by Hottes (1900) point in this direc- tion. No definite separating membrane can be detected in any of the species at the time of zonation, although the delimitation of periplasm and ooplasm is very sharp. Analysis of the condition shows that the differentiation is solely dependent upon the dif- ference in character between the ooplasm and periplasm that has been described in previous paragraphs. It is outside of the ooplasm that the nuclei accumulate, and here in A. Bit, A. Por- tulacae, and A. Tragopogonis they divide, some of the daughter nuclei returning to the oosphere. The plasmoderma is formed at about the time that the pri- mary oospheric nuclei renter, and at a period slightly later evi- dence of plasmolysis may be found. It appeared possible from some conditions seen in A. Bin (Stevens 1899, jigs. 65-67) that the nuclei might take some part in the formation of the new plasmoderma, but critical study shows that no constant relation is maintained between mitosis and plasmoderma formation. The ‘ j ! j \ 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 241 plasmoderma appears to arise directly from the cytoplasm ina manner closely resembling that which Mottier (1900) describes as a rearrangement of alveolar planes. The alveolae are here so small that it is impossible to assert with certainty that the proc- esses are identical. Simultaneous division of the nuclei in oogenesis and spermat- ogenesis is a phenomenon of wide distribution among the algae and fungi. The simultaneity itself is not remarkable, since it is frequently characteristic of multinucleate masses of cytoplasm, as endosperm nuclei of angiosperms, latex vessels (Pirotta and Buscalioni 1898), and plasmodia. The simultaneity in oogenesis is, however, of quite a different nature. Numerous vegetative nuclei, probably of very different ages, accumulate in the rudi- mentary sex organs and are there cut off from the parent cell. These nuclei pass simultaneously into mitosis, while the nuclei in the vegetative mycelium do not do so.5 This simultaneous mitosis, while it may be regarded with Hartog® (1891, p. 23) as a “ phylogenetic reminiscence,” is still often something more, and in the case of Albugo it is appar- ently a step necessary to the sexual differentiation of the gametes. Whether a reduction in chromosomes occurs in connection with this gamete production is uncertain. To be sure Berlese (1898) claims to count the chromosomes during mitosis and fusion, and to establish definitely that reduction occurs in germi- nation. The nuclear phenomena which he describes are so dif- ferent from the conditions seen by Wager (1896), Davis (1900), and myself (1899), that the evidence must be accepted with reserve. The distinct difference in character between first and second mitosis in Albugo is, as I have said in another part of this paper, probably due to change in kinoplasmic content. 5 Frequently nuclei in the immediate vicinity of the oogonium show a slight ten- dency to divide, and may even attain to the spirem stage (Stevens 1899, fg: 45). e can only regard the nuclear divininas in oogonium and antheridium as Phylogenetic reminiscences of the formation of gametes by cell division ; the peri- plasm is thus equivalent to a number of ee es gametes which have taken on the function of epispore formation; the multitude of gametes are sacrificed to the Ww. 242 BOTANICAL GAZETTE [OCTOBER In three species of Albugo there are two mitoses, in another species probably two, in the gametangia; while in Peronospora (Wager 1900) there is only one. In Sphaeroplea (Klebahn 1899) the antheridial nuclei divide repeatedly, while the egg nucleus does not suffer visible change. In Vaucheria, even if there is a division in the oogonium, as seems possible (Olt- manns 1895, p. 392), it is probably not a differentiating divi- sion, since all of the nuclei but one wander back to the parent filament, and are presumably capable of vegetative function. From these examples it appears that the number of divisions in both oogenesis and spermatogenesis varies in different species, and differs in the two sexes of the same species; while in some forms there seems to be no mitosis directly concerned in the genesis of the female pronucleus. These conditions render improbable the existence of a reduction in the number of chromo- somes during gametogenesis in these algae and fungi. Oogene- sis and spermatogenesis begin almost simultaneously fora given pair or group of sex organs, yet all efforts to correlate their inception with any development external to the organs them- selves were vain. A comparative study of four species shows no constant relation between the male and female organs in the sequence of their development, which seems to proceed inde- pendently in each organ. It is not probable, as might at first seem, that the inception of the antheridial tube is caused by the presence of the oosphere, since fig. 50 presents a case where the antheridial tube grew and functioned normally, yet without a parallel development in the oogonium, or indeed the existence of an oosphere. The factor which determines how many primary oospheric nuclei shall enter the ooplasm is uncertain. Clearly the position of the nucleus during its first mitosis determines whether or not one of the daughter nuclei shall enter the ooplasm. Yet this position seems to be governed by no law, the greatest irregu- larity existing, as is general in cases of simultaneous division in multinucleate masses of cytoplasm. This irregularity is equally prevalent at the time of zonation (figs. 2, 28). It is quite ' { : : 1901 | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 243 possible, therefore, that mere accident determines how many of the nuclei are to be so oriented as to contribute daughter nuclei to the oosphere. It is needless to attribute to the cytoplasm any special selective power which causes definite nuclei or a definite number of them to return, although such selection is exhibited in some plants. For example, in the oogonium of Vaucheria - (Oltmanns 1895) one nucleus maintains its position near the beak, amid violent activities of the surrounding cytoplasm, while the chloroplasts and numerous other nuclei are withdrawn. In the oogonium of Peronospora, according to Wager (1900), one nucleus of the many is selected to reenter the ooplasm and func- tion as the female pronucleus. The elimination of supernumerary nuclei by digestion in the surrounding cytoplasm finds analogy in vegetative cells in the Sieve tubes of Pinus (Strumpf 1898); and still more striking in the sexual cells of Achlya (Trow 1899, p. 156), where there is an average of ten times as many nuclei as there are eggs to be produced. The phenomenon is also analogous to that described by Riickert (1892) and Oppel (1892), where several male nuclei enter the egg, the superfluous ones degenerating and functioning as yolk nuclei. In Actinosphaeria Hertwig (1898) also describes a reduction from multinucleate to uninucleate condition before fertilization, which is essentially similar to that seen in Albugo, the supernumerary nuclei being dissolved in the surrounding cytoplasm. In this species, as in Albugo, the nuclei are all alike, showing no such differentiation as is common among the infusoria. The phenomena seen in the Fucaceae (Oltmanns 1889), particularly in Himanthalia lorea, may be classed in this Same category. Here the cytoplasm in early stages contains eight nuclei, the number being later reduced to one by casting off the seven superfluous ones. In Vaucheria (Oltmanns 1895) the reduction from a multi- nucleate to a uninucleate condition is effected in quite a different manner, by protoplasmic streaming, the supernumerary nuclei being carried back into the parent hypha. Thus the reduction occurs before the oosphere is differentiated, and not in the 244 BOTANICAL GAZETTE | OCTOBER ooplasm proper. Klebahn (1899) describes in Sphaeroplea a case where the reduction from the multinucleate to the uninu- cleate condition does not occur until after fertilization, even if then, there being several nuclei in the oosphere only one of. which is fertilized. The nuclei which receive no sperms are recognized in later stages by their smaller chromatin content, . but their fate on germination was not followed. Golenkin (1900) ° agrees with Klebahn in finding a multinucleate egg, one nucleus of which functions; but Golenkin says that the nuclei then all fuse into one. Such a condition offers serious difficulties of interpretation in the light of the present theories regarding the cell. Since uninucleate oospheres were present among the mul- tinucleate ones, it is possible that the condition observed by Golenkin was pathologic, a view which is strengthened by the fact that he was unable to germinate the spores after two years’ trial. In Saprolegniaceae Trow (1895, p. 630, and 1808, p. 166) notes a clumping or possibly pairing of the nuclei as they degen- erate. I have also noted this phenomenon in the foregoing pages. Yet this is in no way to be confounded with the process of gen- eral fusion as described by Hartog (1891, p. 25) and Golenkin (1900), since these writers derive a functional nucleus from the ultimate result of successive fusions, while in the case observed by Trow, and in that seen by the writer, the nuclei thus appear- ing to fuse are really in the process of degeneration. Inasmuch as it has been possible in all cases to follow the parallel development of the oospheres, it can hardly be doubted that in A. Tragopogonis and A. candida, as in Achlya, the Fuca- ceae, etc., the supernumerary nuclei represent potential pro- nuclei, and that each nucleus in the oosphere or A. Tragopogonts and A. candida is homologous with one of the nuclei in the oosphere of A. Blt or A. Portulacae. The coenocentrum has to some extent been sucess in connection with the description of A. Tragopogonts and A. can- dida. It yet remains to compare the structure in the different species. In A. Portulacae it is least developed, consisting simply Igor] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 245 of a large zone of darkly staining cytoplasm which contains at its center the alveolar (trophoplasmic) region. This region sel- dom contains a globule such as characterizes the other three species. In A. Aft the structure is much more prominent and endures for a longer period. It is of more complicated struc- ture than in A. Portulacae, owing to the presence of a distinct central globule. In A. Tragopogonis the coenocentrum is still more highly developed. The central globule seems to be formed by the trophoplasm of the central region, or rather by the coa- lescence of the contents of its vacuoles. This globule in a later period becomes granular, the granules staining like nucleoli. A. eandida possesses the most highly developed coenocentrum which, while closely resembling that of A. Zragopogonis, differs in that from its earliest formation till near the end of its func- tional activity it is thickly beset with coarse granules (jigs. 73, 77) that in size and stain reaction agree with the nucleoli of this species. In its function as well as structure this organ advances in complexity in the series here presented. In A. Portulacae there is no extensive accumulation of nutrient material in the vacuoles of the trophoplasm. In A. Bét this accumulation is marked. In A. Tragopogonis the central globule shows strong chemotactic attraction for the nuclei and serves as nourishment for one or more of them. In A. candida this function of nutrition reaches greater perfection, as is shown by the attachment of the nuclei to the coenocentrum rather than their mere approximation to it. The coenocentrum develops earlier in the more highly differen- tiated species, and thus by exerting its attractive influence upon the nuclei before zonation strongly influences ontogeny. It has likewise probably been an important factor in changing the gen- eral character of oogenesis in phylogeny. The presence of the receptive papilla in the four species of Albugo, as well as in Peronospora (Wager 1900, p. 270), attests to its importance either in the present or in ancestral species. Young stages in the development of this structure show that the plasmoderma adheres to the wall immediately under the 246 BOTANICAL GAZETTE [OCTOBER developing papilla, and exhibits a granular cytoplasm at this point. The granulation probably indicates the presence of a cellulose enzyme, formed here to soften the wall which alters chemically, as is evidenced by its response to stains and by its swollen condition (Stevens 1899, figs. 47, 50). The antheridial tube at this stage of development is probably unable to pene- trate the cellulose wall of the oogonium, and that duty rests with the female cell. Perhaps this curious structure can better be interpreted by a glance at possible ancestral forms. In algal aquatic forms, in which the gametangia open into the water, each sex organ opens independently, the female usually first. If the Peronosporeae have been derived from some such ancestors and these habits have been retained, in species where the walls of the antheridium and oogonium are in contact the origin of the receptive papilla is clear. Gametangia are usually of greatest turgor at the time of opening; therefore, in the present case the bulging, conse- quent upon the softening of the partition wall, is from the oogonium toward the antheridium. The formation of the recep- tive papilla in Albugo occurs immediately before the maturity of the oosphere, precisely as does the analogous phenomenon in Vaucheria. ~The term ‘receptive papilla”’ is a misnomer, since this is not in any morphological sense a receptive structure, nor is it homol- ogous with the receptive spot of the egg. In one case the differentiated area is part of an egg and functions as a place of reception for the sperm (as the eggs of Sphaeroplea, Sapro- legnia, Oedogonium, etc.); in another case it is a zymogenic region of the protoplast adjacent to the point where an opening is to be made in an oogonial wall, either to furnish exit for the female gametes or for the entrance of the male elements, and is homologous with the opening spot of sporangia generally, as Cladophora, Bryopsis, Sphaeroplea, and Oedogonium. In the Saprolegniaceae the papilla does not furnish the place of entrance for the antheridium tube, although in Albugo it does (Zopf 1890, p. 293). The two regions occasion no danger of Igo! | GAMETOGENESIS AND FERTILIZATION IN ALBUGO 247 confusion in the case of the multiovulate (vieleiig) oogonia, nor in cases where the egg is clearly differentiated in a surrounding periplasm; neither should they in forms like Vaucheria and Oedogonium. The migration of antheridial nuclei into the tube seems to be independent of the developmental condition and metabolic activity of the oogonium, and is probably due to a negative rather than positive chemotropism, possibly of such a nature as that suggested by Hartog (1888) for the sporangia of the Sapro- legniaceae. The objections raised by Humphrey (1892) to Hartog’s view do not find application here, since he was con- sidering cases where the sporangium failed to expel any of its spores. In Pyronema, where a condition similar to that of Albugo often prevails, Harper (1900, p. 362) explains the migration of some antheridial nuclei and the passivity of others by assuming that “the chemotactic or other stimulus which leads the male nuclei to migrate through the tube to the oogonium would in this case be assumed to have exhausted itself when a number equal to the number of egg nuclei had reached the oogonium.” Such an hypothesis does not appear adequate for Albugo, for to assumé that a stimulus which can arouse a given number of nuclei to migrate cannot bring the Same activity to a greater number presents serious difficulties, and the supernumerary males are not found em rouse, as would be the case if they were stopped when the female nuclei have consorts and the hypothetical stimulating agent has been exhausted. The failure of some nuclei to leave the antheridium is more probably due to a lack of irritability on the part of some sperms than to a lack of the stimulating substance. The phenomenon of the passage of nuclei from the body of the antheridium into its tube is no more comparable to the seek- ing of the female by a sperm than is the emptying of an algal Sporangium, or an antheridium of the mosses or ferns, or the Passage of the nuclei and cytoplasm from a pollen grain make the tube, all of which are clearly independent of chemotactic influence originating in organs of opposite sex, inasmuch as 248 BOTANICAL GAZETTE [OCTOBER they can occur in the entire absence of the female. It is after expulsion from the paternal gametangium that the chemotactic influence of the female unit is exerted upon the males. In the ferns, mosses, and most algae the chemotactic influence of the female extends over a comparatively wide region. In the spermatophytes the area is much restricted, since the pollen tube opens near the oosphere. In the Albuginaceae, where the male gametangium opens directly into the female gametangium, the region over which this influence may be exerted is still more limited. Indeed it may be questioned whether the force which brings the nuclei of the coenogametes (Davis 1900) together in pairs is at all the same as that which brings gametes together in the open. The pairing of nuclei in multinucleate masses of cytoplasm is comparable to the pairing of the male and female nuclei after the sperm has entered the cytoplasm of the egg, as is readily apparent from a consideration of the con- ditions presented in the larger eggs, as Fucus. Pfeffer demon- strated that the gametes in the open are drawn together by chemotactic attraction, and Wilson (1900) assumes that a simi- lar attraction brings the nuclei together in the cytoplasm of the egg. Apparently two different forces operate, one to bring the sperm to the egg and induce penetration, the other to bring the pronuclei together inthe ooplasm. Conklin (1899) has already distinguished these as distinct factors, and attempts to prove that in some cases at least cytoplasmic currents are responsible for the movement of the nuclei in the cytoplasm. Such an explanation does not seem adequate for the multinucleate oosphere of Albugo, as it would involve complexity inconceiva- bly great. An explanation resting on chemotaxis is more tenable. Conditions where an antheridial tube has reached the oosphere after that organ has been fertilized by another tube Jig. 48), as well as cases where two tubes open into one oosphere (fig. 51), show there is no correlation between the number of female nuclei to be fertilized and the number of nuclei which pass from the antheridial tube. This is also emphasized in fig. 50, where a tube is opening into an oogonium ee Toor] GAMETOGENESIS AND FERTILIZATION 1N ALBUGO 249 having no oosphere, and into which an abundance of nuclei are poured. From these conditions, as well as direct observation, it is clear that the number of males and females is not exactly equal. In A. 7ragopogonis unpaired male pronuclei are some- times seen in the fertilized oosphere, but they are eventually digested by the ooplasm. Similarly, those male nuclei which do not pass into the antheridial tube, and those lying in an antheridium which forms no tube, fail to resume vegetative function and may be found im situ in stages of degeneration. The inability of such nuclei to function vegetatively attests their sexual differentiation and accounts for their subsequent rapid elimination. One observation may help to disclose the force which directs the antheridial tube in its penetration of the oosphere. Two abnormal oospheres were seen quite devoid of nuclei. In one of these only the plasmoderma was developed (fig. 52), in the other periplasm and ooplasm were separated by a wall. In both cases the only antheridial tube present had gone astray in the periplasm. The absence of oospheric nuclei and the misdirec- tion of the tube in the same oosphere may be only a peculiar coincidence. Immediately after the opening of the antheridial tube a wall is seen surrounding the oospore. This develops with great rapidity, often attaining considerable thickness before the pro- nuclei have begun to fuse. Later thick walls, composed princi- pally of cellulose for the nourishment of the germinating spore, are laid down from the inside, and on the outside heavy brown walls which are characteristically corrugated.’ In teratological forms, which abound in all species of Albugo, antheridial tubes where they lie in contact with the periplasm are frequently seen coated with the characteristic pectiniferous deposit. This occurrence was noted by DeBary (1863), and 7 For brevity I shall hereafter use the term _esemteoats to hein sueme this charac teristic brown coating, inasmuch as pectin seems to b t which is ee ere and absent from the other parts of ie fungus. From the literature at hand the deposit appears to be really a mixture of cellulose, callose, cutin, and pectin, although its actual composition is yet open to question (Magnin 1895, Zalewski 1883). 250 BOTANICAL GAZETTE [OCTOBER mention of it is frequently found in works on the morphology of the group. Occasionally I have found isolated balls of this pectiniferous deposit lying in the periplasm. These observa- tions in a measure confirm the generally accepted belief that the outer wall is laid down by the periplasm, a view originally pro- posed by DeBary (1863). Sometimes an antheridial tube penetrates an oogonium that has not yet differentiated periplasm from ooplasm ( fig. 43). In such case it often becomes coated with the characteristic pectin- iferous wall of the species. More frequently it is aborted, reaching a length not greater than one-fifth the diameter of the oogonium. In such cases the adjacent oogonial walls and the remains of the aborted tube receive a pectiniferous coat (/igs. 44-40). These phenomena tend to prove that undifferentiated cytoplasm of the oogonium has the ability to form the pectin- iferous layer, and that in the absence of the plasmoderma of the oosphere it is apparently a matter of indifference what plasmoderma is to receive the deposit, although the pectin is laid down in contact with or by means of some plasmoderma or tonoplast in all cases. Oogonia containing many small pectiniferous spheres (fi. 46) are quite frequently found, but the origin of the spheres cannot with certainty be determined. They are always accompanied by an aborted antheridial tube, and it may well be that these pectiniferous spheres represent deposits upon the lining mem- brane of vacuoles, thus emphasizing the similarity between the tonoplast and the plasmoderma in accord with the view of DeVries and Pfeffer. The fact that an aborted antheridial tube is present suggests that a stimulus may emanate from the antheridium which arouses the protoplasm to pectin production. This idea receives further support when it is recognized that the formation of the pectiniferous deposit begins and is most promi- nent near the antheridial tube (fig. 44). It thus often results in the tube becoming incrusted in a pectiniferous wall. Fig. 45 represents an oogonium prematurely penetrated by two anthe- ridial tubes from opposite ends. These tubes have aborted, but | : 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 251 each is accompanied by a pectiniferous formation from which the middle region of the oogonium is exempt. Occasionally, by the formation of the oospore wall, a super- numerary antheridial tube is pushed aside in the periplasm. It then swells until it assumes a club-shaped appearance as it presses against the wall. In such cases the pectiniferous layer is formed over the whole mass, consisting of the mass of sperm nuclei and the remains of the antheridial tube (fig. ¢8). Con- ditions like these result, in the ripe spore, in a structure which looks much as though an antheridium lying beside an oospore had been encased (fig. 49). Such malformations may be dis- tinguished from antheridia by the still persistent oogonial wall, which would of course not include the antheridium, but do include these masses. If emanations from the antheridial tube stimulate the cyto- plasm of the oogonium to form the pectiniferous layer, why does not the tube become coated as it penetrates the oosphere ? This question probably receives its answer in the fact that the ooplasm has not the power to form pectin. It produces cellu- lose deposits, but not even in teratological cases (jig. 47) does it give evidence of any ability to form the pectiniferous layer. It is also notable that the ability to deposit the characteristic thick cellulose layer and to accumulate the typical oily globules is limited to the ooplasm. It seems, therefore, that in the activities leading to zonation there is a differentiation of the cytoplasm, that a capability vested in the oogonial cytoplasm is lost to the ooplasm, and its manifestation in later stages becomes limited to the periplasmic regions.* Whether this differentiation consists merely in a shifting of partially elaborated products of metabolism or in a segregation of living cytoplasm into two regions, one possessing a different constitution from the other, must be left an open question. ‘ The stimulating effect from the antheridium may be mant- fested when the tube has penetrated only half way to the center ® The figures of Magnin (1895, fg. 71) might at first glance appear to contradic this, but Magnin’s figures represent callose which is really present both in inner and outer walls, 252 BOTANICAL GAZETTE [OCTOBER of the oosphere without emptying, the stimulating agent pre- sumably passing through the wall of the tube when this becomes very thin. The conditions justify the hypothesis that the proto- plasm of the oogonium, or the periplasm as the case may be, contains the food constituents needed for the rather extensive production of the pectiniferous deposit, and that the substance contributed by the antheridial tube is more properly comparable to an enzyme than a food. It is a stimulant to activity rather than a material to act or to be acted upon. The protoplasm of the antheridium of Albugo, moreover, seems never to produce pectin, thus favoring the hypothesis that the necessary food materials are limited to the protoplasm of the oogonium. The antheridial tube in normal conditions opens near the center of the oosphere, from whence the stimulating agent diffuses outward, awakening no response in the ooplasm because this is incapable of response. When this stimulating agent reaches the oospheric plasmoderma it meets the periplasm, which immediately lays down the rudiment of the outer wall. The fact that the tube opens in the center of the oosphere explains the absence of pectin formation in the periplasm, since the large area of the oospheric plasmoderma is ample to receive all the pectin that is to be laid down. Moreover, the thickening wall probably retards the passage of the stimulating agent into the periplasm, so that the peripheral portion of the periplasm is not incited to formation of the pectiniferous layer, and there- fore the oogonial wall receives no pectiniferous coat. An answer to the question originally propounded by Cornu (1872) “si ce plasma extérieur a la propriété de se déposer en couche membraneuse, sans étre élaboré au préabable, pourquoi ne se dépose-t-il pas aussi sur les parvis d l’oogone ?”’ is thus sug- gested. Normally it is the periplasm that responds to the stimulus, and the periplasm is bounded by its own plasmoderma and con- tains its own nuclei. It is an independent unit distinct from the oosphere, although it is destined to sacrifice itself for the pro- tection of the oospore in a manner analogous to the tapetal 1901] . GAMETOGENESIS AND FERTILIZATION IN ALBUGO 253 cells in many pteridophytes (Strasburger 1889). Normally, therefore, the production of pectin in Albugo is analogous to the so-called secondary effects of fertilization commonly seen in the higher plants, since it is an effect manifested by a cell other than those directly concerned in the act of fertilization. SUMMARY OF SECTIONS II AND III. The processes leading to zonation may be regarded as the differentiation of an ooplasm rich in trophoplasm. The nuclei pass outward, possibly to leave part of their kinoplasm outside of the ooplasm, in order to lessen the possibility of partheno- genetic development. The antheridial nuclei give evidence of heightened kinoplasmic content. The cell plate is formed, without the participation of the nuclei, by a rearrangement of alevolar planes. The simultaneous mitosis in gametogenesis is a phylogenetic reminiscence, and was of value in ancestral forms in increasing the number of gametes. No constant time relation is maintained between the phases of oogenesis and spermatogenesis, but each after its inception seems to proceed independent of the other. The orientation of the nuclear figure determines which, and consequently how many, primary oospheric nuclei shall enter the oosphere. This orientation seems to be merely accidental. The supernumerary nuclei are phylogenetically gametes, and their dissolution finds analogy in the Saprolegniaceae and Fuca- ceae, in Actinosphaeria, and in cases of physiological poly- spermy. The receptive papilla is the result of a softening of the oogonial wall by the oogonial contents, accompanied by high turgor in the oogonium. It is probably a vestigial character recalling an algal ancestry. It is a structure of the oogonium, and therefore is not homologous with the receptive spot, which is a differentiated region of the oosphere. The migration of the sperms from the antheridium is homol- ogous with the emptying of a sporangium, rather than with 254 BOTANICAL GAZETTE [OCTOBER the seeking of the female by a male. The number of anther- idial nuclei which migrate into the oosphere bears no constant relation to the number of waiting female nuclei. The failure of some nuclei to leave is probably due to a lack of irritability. Superfluous nuclei of either sex which cannot resume vege- tative function degenerate. The periplasm has the ability to form the pectiniferous deposit, but the differentiated ooplasm cannot.. Emanations from the antheridial tube seem to be needed to stimulate the ooplasm to this activity. The pectiniferous layer is deposited on or by a plasmoderma or tonoplast. The four species, A. Portulacae, A. Bliti, A. Tragopogonts, and A.candida constitute a series in which the coenocentrum increases in complexity, the receptive papilla decreases, and the number of functional nuclei decreases. Of these A. Portulacae is prob- ably the most primitive, and A. candida the most highly special- ized form. The coenocentrum was an important factor in evolution from the multinucleate to the uninucleate condition of oosphere. The division of the fusion nucleus before passing to the winter condition is a consequence of the uninucleate condition, and constitutes the initial step in germination. Delay in the division of the fusion nucleus in a uninucleate oospore is associated with retarded and slow fusion of the sexual nuclei, and is explicable as a consequence of slowness in com- pletion of the last steps of fusion. The relation between Albugo, Peronospora, and Saprolegnia is emphasized by their cytological character, and all are probably derived from a common ancestor having a multinucleate oosphere. The derivation of Peronospora and Saprolegnia from the Chy- tridineae is rendered improbable. Pythium is more closely related to the Albuginaceae than to the Saprolegniaceae. The peripheral gathering of the protoplasm in the oogonium of Saprolegnia may indicate closer relation to Peronospora than to Albugo. | | | | . | . Igor] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 255 If the Phycomycetes are related to Vaucheria it is from a period before the attainment of the uninucleate oosphere by Vaucheria. The coenogamete is homologous with some or all of the gametes of a plurigametic gametangium, not with the individual gametes of such a structure. There is a remarkable agreement between Albugo and Pyronema in many details. The coenogamete is a result of pushing the synplast habit from the vegetative body into the reproductive organs. The synplast of the Phycomycetes is a unit in both morpho- logical and physiological sense, although it is philogenetically the equivalent of many units. BOTANICAL INSTITUTE, BONN. BIBLIOGRAPHY. ARTARI, 1890: Zur Entwickelungsgeschichte des Wassernetzes. Bull. de la Soc. Impér. des Naturalistes de Moscou, No. 2. BERLESE, 1898: Ueber die Befruchtung und ee der Oosphire bei den Peronosporeen. Jahrb. f. wiss. Bot. 31 : —196. BERTHOLD, 1880: Die geschlechtliche Pigieses von Dasycladus clavae- Jormis Ag. Bot. Zeit. 38:648-651. BLACKMAN, Igoo: The primitive Algae and the Flagellata, an account of modern work bearing on the evolution of the Algae. Ann. Bot. 14: 647- CHIMIELEWsKY, 1889: Zur Frage iiber die Copulation der Kerne beim Geschlechtsprocess der Pilze. Arbeiten d. neurussichen Naturf.- Gesselschaft 13 :113. CONKLIN, 1898: Protoplasmic movement as a factor of differentiation. Biol. Lect. at Woods Hole 17-43 Cornu, 1872: Monographie des Saprolégniées. Ann. Sci. Nat. Bot. V. 16 :5—198. DANGEARD, 1890: Recherches histologiques sur les champignons. Le Botaniste 2 :125. rae 1899: The spore mother cell of Anthoceros. Bot. Gaz. 28 : 89-108. 1960: The fertilization of A/bugo candida. Bot. Gaz. 29 : 297-310. ei Bary, 1863: Recherches sur le dével t de quelq hampig parasites. Ann. Sci. Nat. Bot. IV. 20: and STRASBURGER, 1877: [Aastitelanie mediterranea. Bot. Zeit. 35 : 713-728. 256 BOTANICAL GAZETTE [OCTOBER —— 1881: Zur Kenntniss der Peronosporeen. Bot. Zeit. 39 :617—625. -—— 1884: Vergleichende Morphologie und Biologie der Pilze, Mycetozoen und Bacterien. DRUNER, 1895: Studien iiber den Mechanismus der Zelltheilung. Jenaische Zeitsch. 29: 2. FISCHER, 1892: Phycomycetes. Rabenhorst’s Kryptogamenflora IV. GOLENKIN, 1900: Algologische Mittheilungen. Ueber die Befruchtung bei Sphaeroplea annulina und iiber die Structur der Zellkerne bei einigen riinen Algen. Bull. Soc. Imp. Nat. Moscow 1899 : 343. HABERLANDT, 1896: Physiologische Pflanzenanatomie. [2d ed.] HARPER, Ig00: Sexual reproduction in Pyronema nutans and the mor- phology of the ascocarp. Ann. Bot. 14: 321-400 HARTOG, 1888: Recent researches on the heonolegiiacena, a critical abstract of Rothert’s results. Ann. Bot. 2:201-216. 1891 : Some problems of reproduction. Quart. Jour. Micr. Sci. 33 1-79. —— 1895: On the cytology of the vegetative and reproductive organs of Saprolegniaceae. Trans. Roy. Irish Acad. 30 : 649. HERTWIG, 1892: Die Zell und die Gewebe. —— 1898: Ueber Kerntheilung, Richtungs-Kérperbildung und Befruchtung von Actinosphaerium Eichorni. Abhand. bayer. Akad. Wiss. 19:—- [part 3.] HUMPHREY, 1892: The Saprolegniaceae of the United States, with notes on other species. Trans. Ann. Phil. Soc. 17:63-148. IsTVANFFI, 1895: Ueber die Rolle der Zellkerns bei der Entwickelung der Pilze. Ber. d. deut. bot. Gesells. 456. IWANOFF, 1898: Zur sated cele hietie von Botrydium granulatum Woron. et Rostaf. Soc. Imp. Nat. St. Petersburg. Compt. Rend. —:155- JORDAN, 1893: The habits and development of the newt. Jour. Morph. 366. KLEBAHN, 1899: Die Befruchtung von Sphaeroplea annulina Ag. Fest schrift fiir Schwendener 81-10 KLEBS, 1896: Die Bedingungen der Fortpflanzung bei -einigen Algen sund Pilzen. KOHL, 1897: Zur Physiologie des Zellkerns. Bot. Centralb. 72: KORSCHELT, 1889: Beitrage zur Morphologie und oe Pe Zellkernes Zool. Jahrb. Abt. fiir Anat. und Ontog. der Thiere 4 LAGERHEIM, 1899: Mykologische Studien. II. pacan diigo iiber: die Monoblepharideen. Meddelanden fran Stockholms Hégskola no. 199- Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar 25 : 42- u LOEB, 1899: On the nature of the process of “fertilization, etc. Am. Jour. Phyiol. 3:135. i 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 257 MAGNIN, 1895: Recherches anatomiques sur les Péronosporées. Bull. Soc. Hist. d’Autun 8: 3-56. MOTTIER, 1900: Nuclear and cell division in Dictyota dichotoma. Ann. Bot. 14:16 -192. Munson, 1898: The ovarian egg of Limulus. Jour. Morph. 15: 2. NATHANSOHN, 1900: Ueber Parthenogenesis bei Marsilia und ihre Abhangig- keit von der Temperatur. Ber. d. deut. bot. Gesell. 18: 99-109. NoLL, 1888: Die Wirkungsweise von Schwerkraft und Licht auf die Gestaltung der Pflanze. Naturwiss. Rundschau a°57 OLTMANNS, 1889: Beitrage zur Kenntniss der Fucaceen. Abhand. Gesam. 4. —— 1895: Ueber die Entwickelung der Sexualorgane bei Vaucheria. Flora 80 : 388-420. —— 1898: Die Entwickelung der Sexualorgane bei Coleochaete pulvinata. Flora 85 : 1-14. OPPEL, 1892: :Die Befruchtung des Reptiliensies. Archiv. f. mikr, Anat. 39:215-216. pls. g-72. PFEFFER, 1897: Pflanzenphysiologie. [2d ed.] PIROTTA and BUSCALIONI, 1898: Sulla presenza di elementi vascolari multi- nucleati nelle Dioscoreaceae. Ann. R. Inst. Bot. Roma 7: 237-254. RAUWENHOFF, 1888: Recherches sur le Spaeroplea annulina Ag. Arch. Néerland sc. exact. et nat. 22: gI- : ROSTAFINSKI und WorRoONIN, 1877: ait Bot. Zeit. 35 : 649-664 RUcKERrtT, 1892: Ueber phyisologische bie ase bei meroblastischen Wirbelthiereiern. Anat. Anzeiger 7: 320 SCHROETER, 1893: Mucorineae. Die natiirlichen Pflanzenfamilien 17: 119. STEVENS, 1899: The compound oosphere of Adbugo Bliti. Bot. Gaz. 28:14 9. STRASBURGER, 1880: Zellbildung und Zelltheilung. [3d ed 1889 : Ueber das Wachsthum vegetabilischer Zellhiute. Hist. Beitrage 2: 1-186, Sermmmit So) Schwarmsporen, Gameten, pflanzliche Spermatozoiden, und das. Wesen der Befruchtung. Hist. Beitrige 4: 48-158. —— 1893: Ueber 1 oe der Kerne und die Zellgrésse. Hist. Beitrage 5 : 97-1 —— 1897: Ueber aie Jahrb. f. wiss. Bot. 30 : 406-422. ——— 1900: Ueber Reductionstheilung, Spindelbildung, Centrosomen und Clemalaae | im Pflanzenreich. STRUMPF, 1808: Zhistologii sosny (Zur Histologie der Kiefer). Anzeiger Akad. Wiss. Krakau. TROw, 1895: The karyology of Saprolegnia. Ann, Bot. 9: 609-652. 258 BOTANICAL GAZETTE [OCTOBER —— 1899: Observations on the biology and cytology of a new variety of Achlya Americana. Ann. Bot. 13: 131-179. VERWORN, 1897: Allgemeine Pigaiskigic. WAGER, 1896: On the structure and eduction of Cystopus candidus. Ann. Bot. 10: 295-342 — Igoo: On the fextilitation of Peronospora parasitica. Ann, Bot. : O77 a ILSON, 1900: The cell in development and inheritance. ZALEWSKI, 1883: Zur Kenntniss der Gattung Cystopus. Bot. Centralb. 15: 1-10. ZIMMERMANN, 1896: Die Morphologie und Physiologie des pflanzlichen Zellkernes. ZopF, 1890: Die Pilze in morphologischer, physiologischer, biologischer und systematischer Beziehung. Schenk’s Handbuch der Botanik 4:271-756. EXPLANATION OF PLATES I-IV. All figures are from material killed in chrom-acetic acid and stained with Flemming’s triple stain. The figures were sketched with the aid of an Abbé camera, using the Leitz ,, objective, aperture 1.30, and oculars I and 4. PLATE I, Albugo Portulacae. Fig. 1. Early stage in massing of Seca nuclei in advanced sagt receptive papilla prominent. X . 2. Later stage in oogenesis; nuclei near metaphase ; ooplasm and perp of very different structure, but not sharply delimited. x 857- . Zonation ; nuclei in metaphase; ooplasm and periplasm sharply aie as. coenocentrum prominent, consisting of a loosely vacuolate center srecdonabid 2 a denser, slightly granular region which stains darker with the orange G. ve Fig. 4. ena later than fig. 7, primary oospheric nuclei entering oosphere ; ooplasm and periplasm sharply delimited; ooplasm typically alveolar, staining very lightly with orange G; periplasm staining densely with gentian violet, filar in structure; three reentering nuclei show each a very weak but distinct polar ray; coenocentrum has disappeared. X 857. Fic. 5. Primary oospheric nuclei before second division; receptive papilla very prominent; later than jig. 4. X 857 Fic. 6. Receptive papilla open; oospheric nuclei in second mitosis. x oe G. 7. Antheridial ‘tube in oblique section showing many nuclei and no ees oospheric nuclei in second mitosis. x 857. Fig. 8. Antheridial tube in section slightly st te sperm nuclei numer ous, elongated ; stage slightly older than in fig. 7. X Igor] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 259 Fig. 9. Transverse section of the antheridial tube, showing its multi- nucleate character. Oosphere of about the age shown in fig. 7. X 13 Fig. 10. Nuclei pairing after opening of antheridial tube, a distinct aa surrounding oospore; remains of antheridial tube visible in periplasm; degenerating nuclei in antheridium. x 8 Fig. 11. Fusion complete; stage before accumulation of oils and before the outer walls are complete. x 857. Fig. 12. Portion of mature wall. x 857. PLATE Il, Albugo Candida. Fig. 13. Very young coenocentrum showing that the granules pass in from the surrounding ooplasm; oosphere slightly younger than stage next a xX 1366. G. 14. Early oogenesis; oosphere roughly outlined by a withdrawal of the icin from the oogonium wall, leaving only a loose periplasm ehind; nuclei approximately in metaphase; coenocentrum well developed. 1366. Fig. 15. After first division; several ora clustered around the coeno- centrum ; some already gone to the periplasm. Fig. 16. Later than fig. 75; nearly all of we nuclei have retreated to the periplasm; those remaining in the ooplasm in mitosis. X 1366. Fig. 17. Later than fg. 76, mitosis complete. x 1366. Fic. 18. Very slightly later than fg. 77; nucleus in metaphase, aude Owing to attraction by coenocentrum; coenocentrum densely and coarsely granular, FIG. 19. Two nuclei attached to coenocentrum; one in late anaphase, the aie | in telophase ; coenocentrum more dense than in fig. 78 ; oosphere Similar to that shown in fig. 7g. X 13 Fic. 20. Similar to fiz: 79; one eee in late anaphase attached to coenocentrum. x 1366. Figs. 21, 22. One nucleus much enlarged attached to coenocentrum by a pseudopodium-like extension ; all other nuclei lie in the periplasm. X 1366. IG. 23. Before fusion; male and female nuclei and a supernumerary nucleus near the coenocentrum. X 1366 Fig, 24. Metaphase of first division of fusion nucleus. X 1366. Fic. 25. Anaphase of first division of fusion nucleus. X 1366. Fig. 26. An antheridial tube bearing two nuclei; the female nucleus lying in aha ooplasm near the coenocentrum. X 1366. PLATE Ill, Albugo Tragopogonis. Fig. 27. Early stage of oogenesis; protoplasm collected in one central mass; nuclei approaching metaphase; a slight indication of the coenocen- trum apparent in the center. x 200 BOTANICAL GAZETTE | OC1OBER Fic. 28. Condition slightly later than 7g. 27 ; oogonium just before zona- tion; nuclei approximately at metaphase in both oogonium and antheridium ; no sharp separation between ooplasm and periplasm. X 857. Fic. 29. Slightly later than fig. 28; anaphase of first division; daughter nuclei entering ooplasm; ooplasm and periplasm sharply differentiated. xX 1366 Fic. 30. Immediately after the first division; primary oospheric nuclei moving toward coenocentrum, two of them strongly elongated; differentia- tion between ooplasm and periplasm sharp; plasmoderma probably present and many periplasmic nuclei pressing against it; coenocentrum prominent. re Fig. 31. Slightly later than fg. 70; ooplasm and periplasm separated by definite plasmoderma ; oosphere multinucleate ; coenocentrum contracted to a spherical homogeneous globule surrounded by dense protoplasm; primary oospheric nuclei attracted by coenocentrum. X 857. FiG. 32. Metaphase of second mitosis; several nuclei attached to the coenocentrum ; periplasmic nuclei not dividing ; plasmoderma well defined ; coenocentrum staining darkly and not homogeneous. X 8 Fig. 33. Antheridial tube nearly ready to open, eas several nuclei; oosphere in same condjtion as shown in fig. 32, z. e., nuclei in second mitosis. xX 1366. Fic. 34. Similar to #g. 72, nuclei in second anaphase; ten dividing nuclei discernible in this one section. x 857. F1G. 35. One female nucleus lying beside the coenocentrum — somewhat larger than daughter nuclei of second division; general appearance of the oosphere much like that in fig. 76. X 1366 Fic. 36. After the opening of the antheridial tube; definite wall sur- rounding oosphere; remains of antheridial tube in oosphere ; female nucleus in resting condition much larger than in fig. 75 and lying beside the remains of the coenocentrum, which has lost its characteristic form ; sperm nucleus somewhat elongated. x 857. Fig. 37. Male and female nuclei lying in contact, both in resting condi- tion, enveloped by the remains of coenocentrum; male larger than in earlier stages ; compare figs. 75, 36. X 1366. Fic. 38. Similar to fig. 37, also showing degenerate nuclei from same oosphere. X 1366. 1G. 39. Fusion nucleus and remains of coenocentrum. x 1366. Fic. 40. Anaphase of first division of fusion nucleus. x 1366. Fic. 41. Winter condition of spore, showing thirteen nuclei in one sec tion. X 857. Fig. 42. Receptive papilla; oosphere like that shown in fig. 32. 1901] GAMETOGENESIS AND FERTILIZATION IN ALBUGO 261 PLATE IV, Teratological forms from various Shay (X 1366 or 857, and slightly reduced tn reproduction). 1G. 43. Antheridial tube penetrating oogonium in which no oosphere has been differentiated; tube branched and heavily coated with pectin; pectin also on oogonial wall in neighborhood of antheridium. A. 7ragofo- gonts, 1G. 44. Oogonium in which no oosphere was developed; antheridial tube small; deposit of pectin on oogonial wall in neighborhood of antherid- ium ; also isolated pectin deposits, presumably in vacuoles; large deposit on the remains of the antheridial tube. A. Portudacae. IG. 45. An oogonium similar to that shown in fig. gg, two antheridia in contact with an oogonium; deposits of pectin lining the walls and projecting into the oogonium in the neighborhood of the antheridium ; isolated balls of pectin lying free in the oogonium. the rounding up of cells as if going over into palmella. In the columns following those indicating the response is given the time of response in days; 2-12 denotes that the response was observed two days after the making of the culture, and continued for ten days. The second response is the result of evaporation, and comes after a somewhat longer time than the first. The time for it is given in days after the making of the culture. In general, solutions of low osmotic pressure produce zoo- Spores and filaments, while those of high pressure produce only the round-celled form. This is in accord with the results obtained with solutions of mineral salts. However there is one difference in the behavior of the palmella form in solutions of electrolytes and non-electrolytes. This is brought out by Table 294 BOTANICAL GAZETTE [ OCTOBER TABLE I. PALMELLA IN SUGAR SOLUTIONS. LacTosE CANE SUGAR Co t Re- : d Re- | +7; - - . d Re- < tration petal Time as ae eaten sponse Time its se —_ n is n sa Sp. F 2-12 P 30 a Sp. F 5-10 n 7 —~ an, Sp. Bs 27 Fae Sp F 2-9 — | Sp.F P Sp. F 10 p-?. 4-9 30 a pt. 3 a nu x “TO. Sp. F. 9 2% Sp. F 2-18 on 2n oe Re a a Sp. F 2-18 fla *x* P cm ede & Sp* Fj 2-9 Pp 9 a Sp. F 2-18 P 30 vw |Sp.** F, 2 = 14 mie Sp. F 8-14 Io B ce) hid No spores Sh to Pp 14 a Sp.* F 8 r 14 eae 70 10 P 10 Teor EP 10 TABLE, Wi. FILAMENTS IN SUGAR SOLUTIONS. LACTOSE CANE SUGAR Concentration Response Time Concentration Response Time peers es ei : 6-1 3 Sp. F 3 ne No spores 6 Io F es Sp.* F 4-11 10 P> ial No spores Io ia No spores 14 ro P> 10 r> ye edl on - : ‘4 at E pad at, P a 10-12 Io 1a 10 e Bb atc 7H — P 10 — P 10 thls P 9 100} og eo ee eee | Igor] PHYSIOLOGY OF POLYMORPHISM IN ALGAE 295 III, which shows the maximum limit for zoospore production in each case. It is to be compared with Table V of the former article,? from which the figures for mineral solutions have been taken. The figures have been altered to round numbers to avoid confusion. TABLE Ili. MAXIMUM LIMITS FOR ZOOSPORE PRODUCTION. Sotutions oF NoN-ELEC- 3 TROLYTES ae Mineral Original Form Soludions Lactose Cane Sugar 2n in 6 7" Panella onion 5008s or 4 a = . nt Filamentous ....... Eu mas ee. Io Io Io The filamentous form responds in exactly the same manner whether the solution is of electrolyte or non-electrolyte. But to bring about an inhibition of zoospore production in the pal- mella form requires a much higher osmotic pressure when this is produced by non-electrolytes than in the other case. Also the two sugars used differ in that the limit for cane sugar is much higher than for lactose. Why is this? The volume of the cells in the culture is so small, as compared with that of the surround- ing fluid, that the difference just spoken of cannot be explained by the supposition that the carbohydrate is absorbed and thus the concentration of the medium lowered. The amount of absorption possible would be entirely inadequate to alter the concentration to any appreciable degree. It seems more prob- able that by absorption of sugar the concentration of the cell sap is increased, thus decreasing the difference between the osmotic conditions within and without the plant, and so weakening the Stimulus. This would occur if the carbohydrate molecules were to penetrate the cells (7. ¢., be absorbed) more rapidly than the ions of an electrolyte. It is possible that cane sugar is absorbed * Loc cit., p. 313. 296 BOTANICAL GAZETTE [CCTOBER much more readily than lactose; hence might arise the differ- ence between the effect of these two substances upon the organism. Considering these results, together with those heretofore published, theresseems little room for doubt that the response of this plant is brought about by a change in its relation to water. An increase in the osmotic pressure of the surrounding fluid must invariably extract water from the cell (since plant membranes are readily permeable to water), and a decrease of such pressure must cause the cell to take up more. However, an increase in the amount of sugar in the medium might influence the plant otherwise than im the way just mentioned. There might be, for instance, a chemical effect produced by the car- bohydrate molecules. Also, it is readily conceivable that an increase in the number of electrically charged ions in the medium might exert some specific influence upon the protoplasm aside from mere change of water relation. But it is hardly conceivable that any chemical influence exerted by sugar molecules could be identical in its effect with an influence exerted by electrolyte ions. Thus we are almost driven to the following conclusions: (1) since solutions of electrolytes and non-electrolytes affect the organism in the same way, they must exert a common influ- ence upon the cells; and (2) since it is inconceivable that there is any chemical influence common to the two forms of solution, the response must be due to the one factor which is common to both, namely, change in the water relation. Whether this change in the water content of the cells acts merely through the mechan- ical effects of a change in the turgor pressure of the cell sap, oF whether the response is brought about by a more subtle adjust- ment within the protoplasm itself, we have at present no means of telling. It seems probable that both these factors are opera- tive. It is of interest to note here that while the organism often dies in a mineral solution whose pressure is * and invariably dies in stronger mineral solutions, yet it lives indefinitely and apparently without injury in a normal sugar solution. This is Igor] PHYSIOLOGY OF POLYMORPHISM IN ALGAE 297 probably due to the poisonous action of some of the electrolyte ions used. To determine, if possible, what this action may be, will be the object of further research. The same culture may be made to change its form several times in sugar solutions just as in mineral solutions, by adding water or allowing it to evaporate. Nearly all the cultures made in sugar solutions have been controlled by others made at the Same time in mineral solutions, and the control cultures have not deviated at all from those discussed in my previous paper. Further, healthy material taken from sugar solutions behaves in water and in mineral solutions precisely as though it had been grown in a mineral solution. 2. MIXED SOLUTIONS. These solutions contained both sugar and mineral salts. The response of the alga to the weaker ones is the same as though they were composed either of sugar alone or of salts alone. For the stronger solutions the same is true of the filamentous form. But the palmella form gives something of its characteristic response to sugar or electrolytes, according as one or the other of these substances predominates. However, this was not very well marked in the experiments. In general, the plant behaves in the same manner in a mixed solution as in a simple one. 2. POROUS PLATE CULTURES. These were made on unglazed porcelain, such as is used in chemical work. A piece of plate four or five centimeters square was laid on the bottom of the culture dish, and sufficient solu- tion was poured over to stand within a millimeter or two of the upper surface of the plate. Thus the plate was saturated, but no free liquid was upon its surface. The alga cells were placed upon the plate and the whole was covered as usual. The results of these cultures are perfectly uniform, and are exactly what would have been expected. The osmotic pressure of the solu- tion which saturates the plate determines the response of the plant. Typical results from the series of plate cultures are pre- sented in table IV. The abbreviations are those used in tables 298 BOTANICAL GAZETTE [OC1OBER I and II. Ev denotes an uncovered culture with rapid evapora- tion. TABEE FV- POROUS PLATE CULTURES. Solution Original hes onse Time Solution Original Response Time form. P ; " i. form. P . : biiacd 3% 100 L P Sp. F. 9 ay K ig Be 10 8 . 42a x ee S P Sp. F. 9 a. 5 | Sp. F; 9 pn | 4m cal ean Be eo a rarer | ae 0.5 4| B 155 1.8708 Se ey ae errors f ; C 165 1.9915 P $06. 2902. | sccenees D Me. 2.2329 FOG S725..| cestch. 3 J K 3 3.621 S&S een ees i, A .235 | .06 | 2.8364 | 3.42 | 235:3416 | 228.5578 t 4 43 1.0 B 26 3.1382 ee Be eo ae ee era ! $23. Cc 295 3.5606 905.4948 foecezees. Li? 2 322 3.8865 ae, + ee J fi K .4 09 | 4.828 5.13 | 400 368 369.0180 | | |; A 32 sv) 3:BG84 eet ee ORE Damn B emceaiaat x Berea tone! roadway, New Y: NEW WABASH At tate 5 wn oO 4 a ° t3'] ° 53 “t eo A 3 oq oe 2 + fon oO especialy feces ed for dhese cars. The dinin g cars ag nty-nine persons, and hav eample kitchen s e the ace e café cars wi ors seat eighteen ersons in she have a li an g m i ba observation = a car, which will seat four- persons. cars also contain a private ca : — it seating canes ity for eight persons. 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Paid-up Cash Capital, . . $1,000,000.00 BOSEIS, 4% 2. .., 30,561,030.00 RAavilities, . 4.) . . 4. 26,317,003-25 EXCESS, 3% per cent basis, 4,543,126.81 Life Insurance in force, . . . $109,019,851.00 Returned to Policy holders, . 42,643,384-92 J. G. BATTERSON, President. S. C. DUNHAM, Vice President. H. J. MESSENGER, Actuary. JOHN E. MORRIS, Secretary. E. V. PRESTON, Sup’t of Agencies ty mesg = the well-known Bs ha of the IN URIC ACID DIA: WATER GOUT, RHEU- BUFFALO LiTniA WATE = oa WATER oy nae URIC ACID AND PHOSPHATIC SEDIMENTS, ETC., ETC. John. key Shoemaker, M. D.. LL.D., Professor of Matena Meética” a Therapeutics huss Cotarsece College of Philadelphia, etc., in the New ‘York Medical pated june, 22, 1899: “ is mad efficient in Rheumatism and Gost. it the BUFFALO LITHIA WATE ER Pant Uric Acid and Posphatic sediments, well as other inane difficult of elimination, while tthe same time it — a moderately. stimu- fant geiiect witha enal cells, and thereby facilitates the swift removal of. insoluable materials from eb oi elieh action insoluable substances will precipitate in the ekideeus and Bladder. The see cutters produced by Stone, jevenee with consecutive pyelitis and cystitis, are avoided by prompt elimination. Inquestionably, although the speedy removal ot Acid an other products of faulty tissue change is of ace ae henefit yet toP thi their formation IS a service still more important. hen it corrects. those. This service is performed by the BUFFALO L 1A WATER « dtesstive failures. The <— Hunter McGuire > M.D., LL. D., Formerly President and Professor of Cte ui ery, eg 4 College big Medicine, ” Richmond, Va., and Ex-Presiaent of the American Medi Te Says ” as an alkaline diuretic “a invaluable. tn Uric Acid THIA WATER Gravel, and indeed in diseases generally dependent Upon a Uric Acid 0 LTH it is a remedy of extraordinary pabioec I have prescribed itin Cases of Rheumatic ig which had peer the =. remedies, with wonderfully good oe I have used it also my Own case, being a great sufferer fr rom this malady, derived more dicniefit from it than from any pea serail Dr. P. B. Barringer, Professor of Physiology and Surgery, University of Virginia: Pd more than twenty years of-practice | have used Lithia as an anti-uric acid agent any times, e tried itin a great variety of forms, both in the NATURAL WATERS a nT Ss As the result of this experience, | have no hesitation in ne. se for prompt idea have found ne enting uric acid deposits in the agers with : BUFFALO LITHIA WATER i. aay: cane rh ar hag with it as a solvent ne hah ins oe (calcull) has “aiieg ims limited, fea Il hesitate to compare ey ioe €r torms to their disadvantage: but 2 first class of conditions above ot forth | feel that + BUFFALO LITHIA WATER ALONE.’” Dr. Thomas H. Buckler, of Paris (Formerly of Baltimore), Suggestor oy Lithia as a Sol vent for Uric Acid, says: *% Nothing I could sa say would add I have frequently Ayes . ee st hae d GOUT, and wit this object | have esults in URIC ACID DIATHESIS, RHEUM an - ine carbonate, a ordered it to E r et hia is in no form so valuable as where it exists in ae Ee ues nature’s mode of.solution and division in orm i it is roti ste 3 Water which has passed through Lepidotite and ei lh Minerai formations.” Dr. J. ~. Mal — of Chennstry: University of Virginia. Extract from report of analysis llet t Calcul? fitéche rged by BUFFALO Spring No. 2. patients under the action of LITHIA WATER * R Sy and MAINLY that the action of the water is EXERTED agen PGRIC A Eip AND T HE URATES, but when these constituents Sa along with and as cementing matter to Phosphatic or Oxalic Calculus materials, the Me So detached and broken down as to disintegrate the Calculus as a whole in these ae “also us admitting of Urethral discharge.’ vames t. C bell, M.D., , LL. D., Formerly Professor of Prysiology and Surgery in the kgeumed q the ane of Vira. pit President of the National Board of Health, says: hn Uric Acid Diathesis is a well-known thera- peutic ¢ resource. it should be recognized by the Profession as an article of Materia Medica.”’ WATER is for sale by Grocers and Druggists generally. ii TESTIMONIALS WHICH DEFY ALL IMPUTATION OR QUESTIONS SENT TC ANY ADDRESS PROPRIETOR. BUFFALO LITHIA SPRINGS . VIRGINIA — a a LOOK AT THE LABELS! a THE GENUINE CHOCOLATE pa ° IN PENIS LUNE While Liss MOST AND BEST es WaLTER BAKER & Co..Ltp. ESTABLISHED 1780. DORCHESTER, MASS. Maeno eesrerensanconersunes EES NERDS RIBS Arcs Dette 9 WEBER PIANOS are used and enthusiastically endorsed by the WORLD’S GREATEST SINGERS because of that pure, rich and SYMPATHETIC TONE in the possession of which they stand alone WEBER WAREROOMS: 108 Fifth Avenue, New York. 266 Wabash Avenue, Chicago. 181 Tremont Street, Boston eR ae ig The “Could never havert what SAPOLIO. ‘* , yvaula Rs doing for women today.., ; dint of bard w ork littte Scour | % er kitc ae aterail s W Ta ss That would se pase for e polish teday But t the vost: of the scouril THE BOTANICAL GAZETTE EDITORS JOHN M. COULTER ann CHARLES R. BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS J. C. ARTHUR FRITZ NOL L | Purdue University University of Bonx : CASIMIR DECANDOLLE VOLNEY M. Sikora Geneva : iversify Eg Michigan J. B. DETONI ROLAND THAXTE Cniversity of Padua flarvard sea ADOLF ENGLER WILLIAM Pa seat aa University of Berlin issourt Botanical Garden LEON GUIGNARD H. MARSHALL WARD L’ Ecole de Pharmacie, Paris University of Cambridge ROBERT A. HARPER EUGEN. WARMING University of Wisconsin University of Copemhagen _JINZ6 pata: MURA VEIT WirrRogs vial University, Tokys oyal Academy = Sttences Stockhoi: CHICAGO, ILLINOIS Published by the Gnibdersity of Chicago Che Aniversity of Chicago press COPYRIGHT Ig0I BY THE UNIVERSITY OF CHICAGO Vol. XXXII NOVEMBER, 1901 No. 5 ho Z The soap made by PEARS. j in Great Britain, is incom- parably the purest and best for the toilet and bath. oe ors Of stores sell i, all sorts of people use i All rights secured. be rem 7, AS Sens: should be | ak payable to the order of The commend of Chica Hotanical Gazette A Monthly Journal Embracing all Departments of Botanical Science ee el year, $4.00. Foreign, $4.50. Single Numbers, 40 Cents tion price must be paid in advance. No numbers are sent after the expiration of the time paid for. FOREIGN AGENTS: Great Britain— Wm. WEsLEY & Son, 28 Essex Continental Europe—GrBrtipER BORNTRAEGER, St., Strand, London. 18 Shillings 6 pence. Berlin SW. 46, Schdnebergerstr. r7a. 19 Marks Vol. XXXII, No. 5 Issued November 25, 1905 CONTENTS NEW OR LITTLE KNOWN UNICELLULAR ALGAE. II Et cab ela VIRIDIS AND EXCENTRUSPHAERA. (WITH PLATES X-xI1) George Thomas Moor 309 DEVELOPMENT OF THE POLLEN IN SOME ASCLEPIADACEAE. ContTRIBUTIONS E Hutt BoranicaL LaporaTory. XXXII. (WITH PLATE xl) 7.C. Frye 325 ON THE ae ae OF RED COLOR IN a a a pant IN eee ies ENG F.. Grace Smith 332 SOME PLANT ABNORMALITIES. (WITH THIRTY-SIX FIGURES) George Harrison Shull - 343 BRIEFER ARTICLES. MEISSNER ON EVERGREEN NEEDLES. (WITH ONE a ay Edwin a een - 356 THE INSTABILITY OF THE ROCHESTER NOMENCLATUR M. L. Fern 359 FLOWER VIsITs OF OLiGoTROPIC BEES. III. Charles Pilertws - - - - ay CURRENT LITERATURE. Boo EVIEWS - - > - - . - - - + 365 LATEX AND MUCILAGE A MANUAL OF BACTERIOLOGY THE FLORA OF ALABAMA METHODS IN PLANT HIsTOLOGY MINOR NOTICES - - - - - - - - =! gpg NOTES FOR STUDENTS - " - . z = : : eae NEWS i ‘ < : ; . . - Ps - . - 379 Separates, if desired, must be ordered in advance of publication. Not less reg 50 separates of lead- ing articles will be printed, of which 25 (without pened oe be fur! —< gratis, the actual cost of the b temainder (and covers, if desired) to be paid for by th or. Separates of “ brite toe ” (with or without covers) will also be supplied at cost.. The ll A.cl. 7 matter. } BOOKSELLERS STATIONERS A. C. 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The a Tad of ps aah to Solutions, Considered from the Standpoint of the Chemical Theory of Diwociation, SUBJECTS FOR 1908: fA h f Thallophytes, Str +6 S : a én. h . _% * Cra gc 2. Life Hist y poroz GLOVER M. ALLEN, Secretary. Boston ea ang F NATURAL Pea ston, ae. eg § BaP if Ses [JUST PUBLISHED] Send 12 cents, U. S. stamps, for f Wm. Wesley & Son’s Botanical Catalogue, 1901 ‘ SSS SSS55555554 BOTANY 28 Essex Street, Strand, - - =. London, England. os a: So oe bas ec ae Ree ee Being CONTENTS: No. 137-138 of Transactions of Scientific Societies Periodicals Bibliography History Biographies and Portraits , scars Herbals Early Botanical Science Linnaeus NATURAL Handbooks HISTORY and SCIENTIFIC BOOK CIRCULAR Microscopy Morphology and Physiology Encyclopaedic works Classification Nomenclature Cryptogams Phanerogams Fossil Plants Natural distribution of Plants (Floras) Agriculture and Horticulture to the end of the 18th century Gardening Landscape Gardening The Flower and Ornamental Garden Husbandry Tropical Agriculture Commercial Plants F04 pages Medical Botany Forestry + -—=—Diseases of Plants William Wesley & Son, Booksellers, — More than 3300 works, classified under 42 headings SSSR SSS54 or e Every Botanist Should be familiar with the prominent works of GEBRUDER BORNTRAEGER Publishers. Die Glykoside. Chemische Monographie der Pflanzengly- koside nebst systematischer Darstellung der kinstlichen Glykoside von Dr. I. I. L. von Ryn, Director der Reichsver- suchsstation in Mastricht. 8vo. Cloth, $2.50. Das Werk giebt—wie es bisher noch nirgends geschehen — eine ein gehend chemische Behandlung der Glykoside—nicht nur eine kurzgefasste Zusammen- stellung der chemischen Eigenschaften dieser Korperklasse, sondern die Darstel- lungsmethode, die Griinde, welche zur Aufstellung der Constitutionsformein gefithrt haben etc., so dass das Buch in chemisch-pharmaceutischen wie pharma- kologischen Vise sowie unter den Studirenden und sonstigen Freunden der phytochemischen Forschung sicher mit grosser Freude begriisst werden wird. Die Harze und die Harzbehalter. Historisch- kritische und experimentelle, in Gemeinschaft mit zahlreichen Mitarbeitern ausgefiihrte Untersuchungen von Proressor Dr. A. Tscuircu, Director des pharmaceutischen Institutes der Uni- versitat Bern. Mit 6 Tafeln. 8vo. Half calf, $5.00. Das Werk stellt zum ersten Mal das gesammte Material dieser wichtigen Gruppe von Pflanzenproducten kritisch durch gearbettet dar. Die streng wissen- Schaftlichen Untersuchungen werden auch fir die Praktiker, besonders fiir die, die sich mit Harzen und Harzproducten beschiftigen, von Interesse sein, da fede rationelle Technik ja auf wissenschaftlicher Grundlage ruht. Write for free Catalogue; postpaid. Address: Gebriider Borntraeger, Publishers, BERLIN, S. W. 46 SCHONEBERGERSTRASSE 17a zx+1080 pages, 8vo Britton’s Manual of the Flora of the Northern States and Qanada By Proressor N. L. Brirron,: Director of the New York Botanical Garden Seve sce weet Sons. HIS manual is published in response to a demand for a handbook suit- able for ordinary school use, which shall meet modern requirements and outline modern conceptions of the science. trated Flora, prepared by Professor Britton in co-operation with Judge Addison Brown, and published in three volumes by Messrs. Chas. Scribner's The text has been revised and brought up to date, and much of novelty has been added. All illustrations are omitted, but specific reference has been made to all of the 4,162 figures in the /é/ustrated Flora. It is based on An J/lus- CONWAY sek ee er rs SClEaNCE The no work e xtant in the descriptions of in “ass series my ken he flowering ‘plants a can —- a moment be compared with it, eit ither for a skillful and ero geo t for versity of Minnesota, can botanica] publications which deals with h tion of the su sect matter work must consist. 29 West Twenty-third Street, New York CITY — Hol & ae 378 Wabash Avenue, CHICAGO When calling please ask for Mr. Grant Save on Books Whenever you need any book, or any information about books, write to me and you will receive prompt attention and low prices. My Stock of Books in all departments of literature is very complete. An Assortment of Gatalogues and special slips of books at re- duced prices sent for a 1o-cent stamp F. E. GRANT 23 West Srey Abvepeta Street,/New York The Botanical Gazette Edited by John M. Coulter, C. R. Barnes and J. C Arthur, with American and foreign associates. Monthly; at least 80 pages. Devoted to the science of Botany in all of its departments, con- taining results of research, book reviews, notes for $4.00 a year; foreign, All subscriptions and students, and news items. $4.50; single copies, 40 cts. requests for sample copies should be addressed to The University of Chicago University Press Division, Chicago, Ill. Second-Hand B CO O KS BOTANY, ENTOMOLOGY, AND MATHEMATICS For sale by CHARLES £: SMITH Bookseller etther in or out of ta lewa City, fowa Send for price lists Methods in Plant Histology BY CHARLES J. CHAMBERLAIN, A.M., Ph.D. Instructor tn Botany in the Untversity of "Chica 20 NOW READY ILLUSTRATED. PRICE $5.50, NET This book contains estes = collecting agen ore sacige ith ranges microscopic pis aay It based upon in cal technique and is the first co mplete suki to ie pu ublished on ‘this subject. It is the result a seve pee! Oger > work with went in Spahth ce at the Un versity: of roe and with University Extension classe i from the Uni. Mi dag i It aim gn ah to meet the ucassics oar nly of the student who has the assistance of an instructor in a fully equipped shri but also, thee student who aoa work by himself and with limited ee Free hand sectioning, the parafin m ethod, the collodion method, oom pei method are treated in ‘aanaghexsile detail. In — chapters spestiis c directions are given for making such preparations as are those who wish to study the plant kingdom from the Algae up t 6 the ae plants. Special attention is paid to the staining of Palekinetis c figures, because the student who masters this problem will find little difficulty in differentiating other structures. 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GEORGE THOMAS MOORE, (WITH PLATES X—XI!) Eremosphaera viridis. Amonc the many algae which have been supposed to rep- resent stages in the life history of other plants, perhaps none have had such a variety of positions ascribed to it as the beau- tiful spherical form known as Evemosphaera viridis. This species was described by De Bary in 1858 in his Untersuchungen iiber die Familie der Conjugaten as a desmid, although he had not observed anything resembling conjugation in the plant, and could only determine its affinities by the mode of division and its general Similarity to other well known desmid forms. About the same time, Henfrey (5) found this unicellular organism in bogs in Northumberland, and described it as Chlorosphaera Oliveri. A Sood account of the general structure of the plant is given, and because of certain appearances which he assumed to be anthe- ridia, Henfrey says ‘in the vicinity of Oedogonieae they (Chlo- rosphaera) will find their true place.” Hofmeister (6), who described Eremosphaera in his memoir on the Desmidiaceae and Diatomaceae, without giving it a name, regarded it as a link between the Desmidiaceae and Palmelleae. Twenty-five years later De Wildeman in a measure con- firmed the view of De Bary, but stated that Eremosphaera 309 310 BOTANICAL GAZETTE [ NOVEMBER more probably belongs to some developmental stage of a desmid, possibly a zygospore, than represents the vegetative condition of any unicellular alga. More recently De Toni (3) describes the genus as an exceedingly doubtful one, suggesting that the large cells resemble fern prothallia, and that we may expect to find it related to the higher cryptogams. Wille (7), in Engler and Prantl, places Eremosphaera among the Pleuro- coccaceae. The last investigator to have had this alga under consideration is Chodat (1), who believes that its affinities are such as to put it with the Volvocaceae ; furthermore, the results of his observation show a condition of polymorphism which has not hitherto been suggested. In addition to the regular vegetative condition and the ordinary methods of multiplication, which will be described later, Chodat found a number of transi- tion stages which he variously designated as ‘ Gloeocystis” forms, ‘‘Palmella” forms, ‘‘Schizochlamys” forms, ‘ Centro- sphaera” forms, etc. According to Chodat, these various stages of Eremosphaera are actually the same thing as the genera which they resemble, and we can no longer recognize them as distinct species, but must include them all under the single name Evemosphaera viridis. It was with the desire to clear up the question as to the polymorphism of this form, and to settlé, if possible, its life history and affinities, that the investi- gation concerning this alga was undertaken. MATERIAL. The specimens first studied were secured from a small pool formed by a sluggish brook running through a marsh near Ridge hill, Mass. Although the pool is quite small, there is usually plenty of water, and at no time during the year is the material absent from this locality. There was no sphagnum growing in the pool, as seems usually to be the case where Eremosphaera is found, but an abundance of Zygnema, Spirogyra, and other Con- jugatae, notably Micrasterias, was frequently present. The plants were first discovered in May 1897, when they numbered about twenty to the cubic centimeter. Since that time they Igor] EREMOSPHAERA AND EXCENTROSPHAERA 311 have been collected during almost every month of the year, always in considerable quantities. In August, 1899, while on Naushon, one of the Elizabeth islands, near the place on the east shore known as Tarpaulin cove, Eremosphaera was found in its typical habitat, namely a low sphagnum swamp. There was practically no difference between the forms from the two localities except that of size, and this was quite characteristic. Out of the two hundred and fifty measurements of the specimens from Ridge hill (fg. 7), the average diameter was 75.45 with a maximum of 105 and a minimum of 67.5; while of the same number of spheres from Naushon (fig. 3) the average diameter was but 35 with a min- imum of 31.54 and a maximum of 40.7. The figures above referred to will give a fair idea of the comparative sizes of the two lots of material. This marked difference in size has been noticed by Chodat (1), who in the spring of 1892 found Eremosphaera which was almost exclusively the large form, while in 1894 he discovered among Sphagnum and Carex a quantity of the small variety. The relative sizes of these two extremes is not given, except for the statement that by careful search one might find ‘‘giant indi- viduals” of 170". De Bary, in his original notice of the plant, gave the measurements as about 60, while Rabenhorst says 43-49". Kischner gives 100-145, and De Toni covers both extremes by citing 100~150m, as the usual size, 30-804 spocuucas being occasionally found. It is evident, then, that the plant is one varying within wide limits. The maximum of the Naushon form not coming within 26m of the minimum of the Ridge hill form, if size be a sufficient criterion there would be no diff- culty in recognizing a variety ™m1nor, measuring from 30-414, and a variety major, measuring from 67-100 or more microns. There can be no question that the smaller forms, found by me at Tarpaulin cove, were mature plants. The arrangement of the chromatophores and general cell contents was in every way iden- tical with the larger forms, and division took place as readily in cells 334 in diameter as in those measuring 1004, Furthermore, 312 BOTANICAL GAZETTE [ NOVEMBER cultures grown from material from the two localities have retained their characteristic size through a cultivation of more than eighteen months in one case, and for over three years in the other. GENERAL STRUCTURE. In appearance Eremosphaera resembles a perfect gelatinous sphere with numerous minute chromatophores usually lining the wall. The arrangement of the chromataphores varies greatly, hardly any two specimens showing exactly the same pattern. Usually they are scattered about in an irregular fashion, some- times singly, sometimes in groups (fig. 7); or they may radiate from the center in a quite definite manner (fig. zo). That light has much to do with the arrangement of the color-bodies may be seen from fig. 4g. This specimen while kept in subdued light showed a cell of almost solid green hue, so thickly were the chro- mataphores lining the wall. After five minutes in bright sunlight, however, the condition shown in fig. ¢ was obtained, and in watch- ing the effect under the microscope the chromatophores could be seen sliding along the protoplasmic strands which radiated from the centrally placed nucleus. When the chromatophores are at the periphery of the sphere and are not too great in number, the nucleus is easily recognized without the aid of stains. The method recommended by Cho- dat (1) of clearing with chloral solution and staining with car- mine brings it out well, and also reveals one or more nucleoli (fig. 2). At times the nucleus and the protoplasmic mass sur- rounding it become quite granular, giving the appearance of some foreign body within the cell. The strands which radiate from the nucleus connect with the protoplasm lining the cell wall and form quite a complete network (fig. 5). The chromatophores are irregular both as to outline and as to size, varying from circular, through broadly elliptical, to nar- row fusiform (jig. 16). Occasionally they are angular, of a rhomboidal outline, a type especially common in the variety minor. The honeycomb appearance described by Chodat (1) Igor] EREMOSPHAERA AND EXCENTROSPHAERA 313 was not visible in my material, neither was the platelike expan- sion which he says is drawn out from the middle of the chro- matophore and may consist of two or three bent wings, causing an irregularly shaped dark spot in its center. A good sized pyrenoid is always distinctly visible, there being sometimes as many as three or four in each chromatophore. If the chromatophore be treated after the manner of Mayer, first adding a dilute iodine solution and then chloral hydrate, the crystalline structure of the pyrenoid is easily seen (fg. z8), and the layer of starch sur- rounding it made visible. The wall of Eremosphaera is normally thin and of but a single layer in thickness, but it has the property of great gela- tinization, and is often laid down in successive layers. This condition gives rise to the forms characterized by Chodat as the Gloeocystis, Palmella, and Schizochlamys stages. The duplica- tion of the wall does not seem to be confined to particular periods of development, but may occur at any time. Each wall soon becomes separate and distinct from the adjoining ones, and with care the several coats may be broken one at a time and the con- tents allowed to slip out. That such forms are really what botanists recognize as Gloeocystis or Schizochlamys is doubtful. While it is possible that Emerosphaera, surrounded by a number of gelatinous walls, might be mistaken by a hasty observer for a similar condition in either Gloeocystis or Schizochlamys, it does not necessarily follow that these genera are one and the same thing. Schizochlamys, I have been able to cultivate for a con- siderable length of time, and it has never exhibited any evidence of being related to Eremosphaera. Since we can have the same duplication of the wall in Chlamydomonas and other genera widely separated from the ones under discussion, it hardly seems a sufficient basis upon which to unite a number of forms frequently showing marked differences in their life histories. In addition to the successive formation of coats around Eremosphaera, there sometimes occur peculiar growths on the inner surface of the wall, which are the result of a number of layers of cellulose being formed about a central point (jig. It). a iS c 314 BOTANICAL GAZETTE [NOVEMBER These concentric layers can occasionally be separated by crush- ing, but usually they are very compact. They project inward towards the center of the cell, and in case a second continuous wall is formed, it remains indented wherever these excrescences appear. As many as twelve secondary walls have been counted around a single sphere, and there seems to be no limit to their formation. ; CULTURE METHODS. A plant which has had as many developmental stages ascribed to it as Eremosphaera naturally necessitates the most careful application of pure culture methods. Consequently the cultivation of this plant was attempted as soon as it was pro- cured, and cultures have been kept running successfully for about three years. As it was not convenient to visit the original locality frequently, and to obtain fresh material in that way, it seemed best to maintain a number of gross cultures from which pure transfers might be made at any time. No trouble was experienced in this, and water from the original pool, containing diatoms, numerous filamentous algae, and Eremosphaera, has been kept continuously in the laboratory. Bacteria appeared for a short time, but the water soon cleared, and all the algae have maintained themselves in good condition. These gross cultures were kept either in crystallizing dishes or wide-mouthed bottles, over which was a sheet of glass to prevent too rapid evaporation. It was found that the crystallizing dish was most convenient for this purpose because of the ease with which material could be picked out from it. The large Eremosphaera cells were readily found with a hand lens, and then transferred by means of a pipette to a watch glass. Here they were washed several times in sterilized water, and then examined under the microscope, before being placed on the culture medium. Various methods of cultivation were tried. The well-known solution of Knop was used in strengths from 0.2 per cent. up to I per cent., both as a fluid culture and in connection with agar agar and gelatin. The gelatin was soon abandoned, however, on account of the low temperature at which it liquefied, since ae 1901] EREMOSPHAERA AND EXCENTROSPHAERA 315 agar agar (5 to 7 grams to the liter of nutrient solution) enabled one to keep perfect trace of the cells, and was more satisfactory for many reasons. Stender dishes were used for culture vessels because it was desirable to have as large an amount of the medium as possible, so that the cultures would last for some time, and in order that there might be a considerable surface for growth. The same precautions of sterilization were observed that would have been necessary for bacteriological work, and check cultures indicated that these methods were successful. Usually after sterilization there would be a considerable amount of moisture collected upon the surface of the agar, and it was in this fluid that the Eremosphaera cells were deposited. If placed directly upon the surface of the agar it seemed impos- sible for them to persist, but when transferred to the moisture and then gradually, through the evaporation of the superfluous liquid, brought in contact with the nutrient agar there was no difficulty in making them grow as well as in fluid media. Van Tieghem cell methods were not successful. The plants lived for a considerable length of time and the chlorophyll gran- ules moved under the influence of sunlight, but there was no development of any kind. Cultures kept in tightly sealed cells for several months would gradually present an appearance resembling the formation of spores, and at first this was thought to be the case. Upon investigation, however, it was found that this effect was due entirely to the rounding off of the chromato- phores, which were closely packed at the periphery of the sphere. Upon being restored to natural conditions such cells resumed the characteristic arrangement of the color bodies, and the nucleus at no time presented any other appearance than that found in the normal plant. Ward cells, which allowed a constant supply of air and yet prevented any contamination from the outside, were partially successful. The most satisfactory method for making direct observations of cultures, however, was to isolate a single individ- ual and place it on a slide either in sterilized water or a ines solution. A large cover-glass was then placed over it, being 316 BOTANICAL GAZETTE | NOVEMBER propped up by three or four minute drops of a mixture of bees- wax and vaseline. A bell jar lined with moist filter paper was sufficient to prevent the evaporation of the solution for twenty- four hours, and a drop of nutrient fluid added at one side of the cover-glass replaced any loss during the examination under the microscope. Such cultures require daily examination, of course, but when the life history of an organism is being followed out this is an advantage rather than an objection. The wax sup- ports for the cover-glass were preferable to bits of glass or filter paper because they permitted the crushing out of a specimen, if necessary, and held the cell in just the desired position. Cul- tures of this kind gave the best of results, the algae apparently developing in a perfectly normal manner. MULTIPLICATON AND DEVELOPMENT. De Bary, in his observations upon Eremosphaera, discovered that it multiplied by simple division into two or four cells, each of which soon attained the size of the original plant. Henfrey likewise recognized this condition. Such a division is accom- plished by the formation of a new wall between the two halves of the original cell. This wall extends completely around each half, and as the nucleus divides and the chromatophores, with the protoplasm, form into two irregular masses, the gelatinous wall pushes in from the periphery, forming the division (jig. 6). The two cells thus formed gradually increase in size, so that in a short time the original wall is ruptured and the daughter plants are liberated. These usually escape as perfect spheres, but may retain their flattened appearance for a short time, as is shown in jig. 7. More rarely a second division may take place at right angles to the first, thus producing four daughter cells instead of two (fig. 8). Wolle (8) and Chodat (1) both speak of even further successive divisions, but my material has never shown more than four cells formed in this way. This method of simple division has been believed to be the only means of propagation in Eremosphaera, and it was not until the result of the researches of Chodat became known, that 1901] EREMOSPHAERA AND EXCENTROSPHAERA 317 a more complicated life history, including the formation of zoospores, was suspected. Henfrey, in his discussion of the plant already referred to, suggested the possibility of anther- ozoids, but this condition was undoubtedly due to the presence of some parasite, which Henfrey himself recognized as a possible cause, While considerable doubt has been cast upon Chodat’s inter- pretation of what he saw, the fact that certain of the stages which he describes certainly exist in material found in this country, makes it necessary that a most careful search be made for all the forms supposed by him to be connected with Ere- mosphaera. In addition to the well-known division into two or four spheres, Chodat (1) describes a kind of sporangium-building condition where division goes on until the diameter of the ulti- mate cells is ten to twenty times smaller than at first. This stage I have been unable to find, and nothing of the sort seems to occur in the material I studied, either in gross or pure cultures. In addition to these non-motile spores, Chodat des- cribes ciliated spores. Just how and when these may be formed is not clear, but it is to be inferred that they arise from some ‘“‘palmella” condition and not directly from the adult plant. These zoospores, according to Chodat, are usually elliptical, and are always surrounded by a gelatinous coat, reminding one of certain forms of Chlamydomonas. Each zoospore has a red eye Spot, a radiating chromatophore, and an evident nucleus. In most cases there are two cilia, but exceptionally three. The question as to whether they actually had four remained unde- cided, as did also the possibility of their conjugation. They seem to have been irregular both as to size and shape, for Chodat (1) says, ‘da ihre Mutterzellen an Grosse sehr verschieden sein konnen, so sind infolgedessen die Zoosporen auch sehr verschie- den, sowohl was die Grosse als auch die Form anbetrifft.” Although a most careful search has been made among mate- rial growing under all conditions, not the slightest trace has ever been discovered of any motile spore in evident relation to Ere- mosphaera. Occasionally plants would be found of a lighter 318 BOTANICAL GAZETTE | NOVEMBER green color, filled with spherical bodies (fig. 73), which in gen- eral resembled a sporangium. But while these cells were observed daily for weeks they not only failed to show any signs of further development, but gradually disintegrated instead. After having observed algae which were crushed and noting the manner in which the chromatophores formed a colorless membrane about themselves, with the green collecting at one side, as in fig. 77, it would seem almost certain that the con- ditions I had been observing were due to injury, and the admis- sion of a small amount of water caused the abnormal appearance suggesting spores. On one occasion, a plant of this sort showed four or five biciliate organisms within it, which at first were sup- posed to be zoospores (fig. rg). Their subsequent development (fig. 15), however, proved them to have no connection with the alga, and after three days, when the culture was crushed out, it was found that the organisms possessed no chlorophyll, a fact which could not be made clear with certainty while they were embedded among the chromatophores. Failing to find zoospores under natural conditions, it was hoped that some change in the environment might be produced which would cause zoospore formation. Plants, closely encysted in numerous gelatinous coats and which had been growing in a nutrient solution, were removed to fresh water and to variously modified media. Some cultures were allowed to remain in the sunlight, others were kept in darkness. Specimens taken direct from nature were placed in numerous solutions and subjected to all possible changes of temperature and moisture. Aeration was tried for lengths of time varying from a few hours to several weeks, and cells were kept on ice over night, and then gradually brought into the temperature of the laboratory. But none of these methods yielded any thing more than the previously des- cribed method of division. Consequently, after having had cultures running for over three years, and making repeated examinations of material collected under natural conditions, it must be said that at no time was there any appearance which gave the slightest evidence of zoospore formation, and it 1901 } EREMOSPHAERA AND EXCENTROSPHAERA 319 is certain that in the forms found growing in this country zoospores must be very rarely, if ever, formed. The rejuvenescence described by Chodat (1), wherein a cell instead of dividing simply slips out from the old sheath, was frequently seen and is represented in fig. 5. Whether this can really be termed a rejuvenescence, according to the ordinary use of the term, is a question. It seems as though this condition can hardly be different from those which show a succession of walls, except that in the first case the outer wall is not strong enough to hold the cell, and it slips out as indicated. There never appears to be any change in the contents of the cell, not €ven in the arrangement of the chromatophores. In the mud at the bottom of the water containing Eremos- phaera, there will usually be found a considerable number of resting cells. These are of the typical brick-red color, and gen- erally show successive layers of sheaths. They contain large quantities of oil, and in no way appear to differ from the same type of spore in other algae. So far as observed, the develop- ment of the resting spore was not accompanied by the formation of any new cells. The red color was gradually lost, the chro- matophores became more and more prominent, and the normal vegetative appearance of Eremosphaera was the result. Usually the resulting cell is quite large and division takes place in a very short time. No division of the resting spores was observed, the condition indicated in fig. 79 being an example of where two cells passed into the resting condition almost immediately after division, and before they had time to enlarge. This is not an infrequent occurrence. TAXONOMIC POSITION. It is evident from the foregoing that there is no reason for supposing that Eremosphaera is related to the desmids or to any of the Conjugatae; neither does it seem possible to show that it forms any part of the life history of any of the higher crypto- gams. The theories of Henfrey (5) and Hofmeister (6) may also be dismissed, so that there only remain the two positions 320 BOTANICAL GAZETTE [NOVEMBER ascribed to it by Chodat (rt) and Wille (7). Chodat, as the result of his study of Eremosphaera, came to the conclusion that it had evident affinities with the Volvocaceae, and if all the stages described by him are really passed through by this plant, it would seem as though some such disposition would have to be made of it. On the other hand, the negative results of my cul- tures and observations would seem to throw some doubt upon its supposed polymorphism, particularly with regard to a motile stage. It seems, therefore, that for the present at least, Ere- mosphaera should be classed with the group of green algae pos- sessing no zoospores, namely the Protococcoideae. Excentrosphaera, nov. gen. There still remains to be discussed the form described by Chodat as the ‘“‘Centrosphaera” stage. This form was present in small quantities in the first material collected at Ridge hill, and was immediately separated from the Eremosphaera cells and cultivated. The growth was luxurious in fluid nutrient media, and satisfactory, if not so abundant, on agar. In addition to these pure cultures, a number of dishes containing water from the pool which supplied the original material were kept in the laboratory, and in the course of a few months became filled with a large number of these plants. The conditions for growth under these various circumstances seemed favorable and normal, but there was at no time any evidence of the slightest connection with the developmental history of Eremosphaera. After having had this form under cultivation for about three years, and having seen it pass through its life cycle over and over without assuming 4 stage which in any way resembled those of Eremosphaera, I am obliged to conclude that it is an independent genus. Since it does not seem to have been described elsewhere, the name Excentrosphaera viridis has been given to this alga. Excentrosphaera, in its mature condition, may assume an out- line varying from that of a perfect circle, through all gradations to an ellipse, as well as occasional excentric and indefinite shapes. The general resemblance to the forms assumed by germinating Igo] EREMOSPHAERA AND EXCENTROSPHAERA 321 Ulothrix spores is striking, but the much larger size, the arrange- ment of the chromatophores, and suosequent development make it an easy matter to differentiate this plant from that genus. The numerous large chromatophores are crowded near the surface of the cell, and are usually arranged radially (figs. 27, 23, 24). As the cells assume irregular shapes the chromato- phores may be distributed in various ways, and there are often lacunae between the chlorophyll masses. Sometimes there are several layers of chromatophores, so that except for the nucleus and a small amount of protoplasm, the cell is almost filled with these bodies. There are numerous minute pyrenoids in each chromatophore, but these are not readily made out without the use of stains. The densely packed chromatophores render it difficult to see the centrally placed nucleus, with its nucleolus, but sections (fig. 25) or crushing will usually reveal it. When the cells are about to form spores the nucleus divides repeatedly (fig. 26), then the chromatophores break up, and ina short time the-entire cell has a homogeneous appearance similar to that shown in fig. 27. The spores, which are formed from this con- dition simultaneously, are 2-3 in diameter, non-motile, and without a red spot. They escape through a hole formed by the dissolution of the wall (fig. 22), and in about a month increase to the size of the mature plants. The developing spore usually remains spherical until it has reached its maximum size, and not until then does it begin to take on the irregular shape previously referred to. The wartlike projections on the wall, reported by Chodat (1), frequently occur in plants after maturity has been reached, but in small spherical forms are quite rare. These formations are made up of a series of layers of cellulose, and often increase until they are of considerable size (figs. 23, 24). ; Resting conditions, with a very thick wall and of a reddish color, were found in pure cultures, but the mode of development has not been observed up to the present time. As previously stated, this plant has been cultivated in a pure state for several years, and all the methods resorted to in an effort to bring about 322 BOTANICAL GAZETTE | NOVEMBER zoospore formation in Eremosphaera were repeated with Excen- trosphaera, but without success. The Eremosphaera material found on Naushon has never shown Excentrosphaera, nor has the Ridge hill material devel- oped it after the original lot was collected. Excentrosphaera has been found in stagnant ponds near Norwich, Vt., which contained, in addition, large amounts of Nitella, Spirogyra, Oedogonium, and related forms. A shallow pool, almost filled with Hydrodictyon, not far from Boston, has also furnished Excentrosphaera in considerable quantities. Neither of these lat- ter localities have ever shown Eremosphaera, although repeated search has been made for it. Unless we are to adopt Borzi’s ‘‘stadii anamorphici”’ for all the algae, it does not seem possible that this plant has any genetic connection with any other form. The external resemblance to Centrosphaera, which led Chodat to give that name to it as a supposed stage of Eremosphaera, is certainly striking, but the decided difference in habitat, together with the absence of motile spores and the difference in develop- ment, would seem to be sufficient to separate it from that genus. The affinities of Excentrosphaera, so far as known, must be with the Protococcaceae of Wille. Excentrosphaera, nov. gen. — Plant consisting of a single cell, in mature condition varying in outline from spherical and elliptical to irregular and excentric forms. Chromatophores large, angular, usually radiately arranged, closely lining the wall. Pyrenoids minute, numerous in each-chromatophore. Multipli- cation by means of non-motile spores (aplanospores), which escape by the dissolution of a part of the cell wall. Reaction to all external stimuli negative. E. viridis, nov. sp.— Plate XII, figs. 21-27. Characters of the genus. Plants of bright green color; size of mature cells 22-55. Spores 2-34. Growing with Eremosphaera, Geneva (?); with Eremosphaera, Micrasterias, Zygnema, etc., Ridge hill, Mass., the year around; in swamps with Nitella and various algae, Norwich, Vt., September—November; with Hydrodictyon in shallow pool, in vicinity of Boston, June-August. BOTANICAL GAZETTE, XXXII MOORE on EREMOSPHAERA PLATE X oe oe BOTANICAL GAZETTE, XXXII PLATE XI ee ee ee eee | MOORE on EREMOSPHAERA BOTANICAL GAZETTE, XXXII MOORE on EXCENTROSPHAERA PLATE Xifl 1901] EREMOSPHAERA AND EXCENTROSPHAERA 323 This work was commenced in the Cryptogamic Laboratory of Harvard University, and my sincere thanks are due to Dr. Farlow and to Dr. Thaxter for their helpful criticism of the investigation carried on while there. DARTMOUTH ini Hanover, N. H. BIBLIOGRAPHY. 1. CHODAT, R., Ueber die Entwickelung der Evemosphaera viridis DeBy. Bot. Zeit. 53: 137-148. 1895. 2. DE Bary, A., Untersuchungen iiber die Familie der Conjugaten. Leipzig, 1858. 3. DE Tonl, J. B., Sylloge Algarum 1 :616. 1889. 4. De WILDEMAN, E., Comptes rendus de la Soc. Roy. de Bot. Belge. 1894. 5. HENFREY, A., On Chlorosphaera. Trans. Micr. Soc. Lond. 7: 25-29. 1859. 6. HormeisTER, W. F. B., Ueber die Fortpflanzung der Desmidieen und Diatomeen. Leipzig. 1857. 7. WILLE, N., Die natiirlichen Pflanzenfamilien, Algen 17:58. 1897. 8. WOLLE, F., Freshwater algae of the United States. 1: 201. 1887. EXPLANATION OF PLATES. All the figures are from ink drawings sketched in with an Abbé camera. In the reproduction = are reduced about one fourth. Figures 44-78 are drawn with a Leitz +5 z (oil), oc. 3; all the others with a Leitz } oc. 3. The magnifications given are ihe original ones before reduction and allow for projection. PLATE X, Eremosphaera viridis De Bary. Fie. 1. Surface view of large variety (Ridge hill material). x 250. Fig. 2. Section of same showing nucleus and protoplasmic strands. X 250. Fig. 3. Surface view of small variety (Naushon material). X 250. Fic. 4. Surface view showing the retreat of the chromatophores under the influence of strong sunlight. X 250. Fig. 5. So-called “rejuvenescence.” x 250. Fic. 6. Beginning of the division into two. FiG. 7. Division completed and liberation of auughter cells. X 250. Fic. 8. Division of mother cell into four. x 250 FIG. 9. Division completed and liberation of daughter cells. x 250. Fic. 10. Escape of cell from old wall after “rejuvenescence.”” X 250. 324 BOTANICAL GAZETTE [NOVEMBER PLATE XI, E bh trtdzs De Bary. J Fic. 11. Surface view and section of wartlike formations in wall. X 250. Fig. 12. Section showing successive formation of walls. X 250. Fic. 13. Abnormal condition resembling zoosporangium. X 250. Fic. 14. Foreign organisms found within cell similar to fig. 73. X 830. Fic. 15. Ultimate development of ciliated organisms. X 830. Fig. 16, Normal appearance of chromatophores, with pyrenoids. 830. ’ Fic. 17. Chromatophores after being crushed out in water. X 830. Fic. 18. Pyrénoids after treating with chloral hydrate. X 830. F1G. 1g. Resting spores formed immediately after division. x 250. Fic. 20. Resting spore formed from mature plant. X 250. PLATE XII. Excentrosphaera viridis Moore. Fic, 21. General appearance with successive stages in development from spore. X 250. FiG. 22. Escape of spores. X 250. FIGs. 23, 24. Cellulose projections of wall. x 250. Fig. 25. Section through cell, stained to show pyrenoids. X 250. Fig. 26, Section showing first divisions of nucleus previous to spore for- mation. X 250. Fic. 27. Homogeneous appearance of cell previous to spore formation. X 250. } DEVELOPMENT OF THE POLLEN IN SOME ASCLEPIADACEAE CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY, XXXII. T. C. FRYE. (WITH PLATE XIII) It 1s well known that in the Cynanchoideae’ the microspores of each sporangium adhere in a mass, forming what is known as apollintum. This is true also of some of the Orchidaceae and Leguminosae. Since the three families mentioned are all highly specialized for insect pollination, the adherence of pollen may perhaps be regarded as having no special morphological signifi- cance. The Asclepiadaceae are further exceptional in the production of only two sporangia in each stamen. In the Balanophoraceae, Orchidaceae, and Mimosoideae, the number of microsporangia is variable. The large number in some of the Mimosoideae is attributed to the interjection of plates of sterile tissue. In the same way Lemna minor is said to form four, the normal number, from a single mass of archesporial tissue. The question arises whether, in the Cynanchoideae, there is any indi- cation of suppression or fusion of sporangia in the earliest stages of their development. More interesting does this become when we recall that in the small group Secamoneae the pollinium in each half anther is paired, but the parts adhere closely. Little is known about the formation of pollen in the Ascle- Piadaceae. In the Periplocoideae tetrads are formed, the micro- Spores adhering in groups of four when mature. Of the pollen development in the Cynanchoideae almost nothing is known. *The author follows the classification of Engler and Prantl in Dye nattirlichen Phlanzenfamilien. = * CALDWELL, O. W., Onthe life history of Lemna minor, Bort. GAZ. 27 : 37-66. 99. 1901] : 325 326 BOTANICAL GAZETTE [ NOVEMBER Chauveaud3 claims that the pollen mother cells of Cynanchum divide, but mentions only one division, with reduced number of chromosomes, the pollen grains forming from the daughter cells. Here one is left to infer that a mother cell formed only two microspores, or that there was a second division which escaped his observation. Wille+ examined Asclepias speciosa, but was unable to find tetrads. Strasburger’ finds no tetrad division in Asclepias Syriaca. He finds in the anthers what he terms sporog- enous cells, radially elongated, large, and rich in contents. These divide by cross walls into two, and the longest, near the middle of the sporangium, into four. He says the resulting cells are homologous with the pollen mother cells of other plants, and develop into microspores without further division. So far as I have been able to learn Zostera® and the Cynanchoideae are the only Angiosperms which are reported to form their pollen without tetrad divisions. It was thought that there were no tetrad divisions in the Cyperaceae until Juel? found that the nuclear division occurred, but no walls were formed. The reported different pollen formation in Cynanchum and Ascle- pias, the presence of tetrads in the Periplocoideae, and the small number of cases of pollen formation without tetrad division, all seem to warrant an investigation of the formation of microspores in the Cynanchoideae. In the spring of 1901 I undertook the task of looking into the pollen formation of some members of the Asclepiadaceae growing abundantly in the vicinity of Chicago. Those examined were Asclepias Cornuti, A. tuberosa, A. phytolaccoides, A. incarnata, A. verticillata, Acerates viridiflora, and A. longifolia. In A. Cornuti 1 was able to trace the development from the 3 De la reproduction chez les dompte-venin 41. (Diss.) Paris, 1892. Paha gape die Entwickelungsgeschichte der Pollenkérner der Angiospermen 4I. 5 Ueber das Wachsthum vyegetabilischer Zellhaute. Histologische Beitrige 2 : 80. ° HOFMEISTER, Bot. Zeit. 10: 121. 1852. ? Beitrage zur Kenntniss der bintovantned a eg Die Entwickelung der Pol- lenkGrner bei Carex. Jahrb. f. wiss. Bot. 58:6 ; 1901 | DEVELOPMENT OF THE POLLEN 327. archesporium to the pollen mother cell. The first indication of the formation of sporangia is the increase in length of the hypo- dermal cells on the inner side of the stamen on each side of its median line, thus forming the archesporium (jig. r). While both sporangia are formed from the same continuous layer, they are separate from their beginning ; and there are no indications of four. The tapetum on the dorsal side of the sporangium is formed from the third layer of cells—the one next the arche- sporium. The archesporial cells elongate at right angles to the inner surface of the stamen and divide transversely, resulting in an inner primary sporogenous, and an outer primary wall layer. The beginning of this division is shown in fig. 7. 2 By transverse divisions of the primary wall layer there arise four wall layers ( fig. 3), the inner one or two of which form the tapetum. The tapetal nature of these cells is already evident in their large nuclei and nucleoli, and their deeply staining contents, before they have completed their division ( fig. ¢).- In some places the tapetum is composed of one layer, in others of two. Where there are two the outer is often composed of flattened, the inner of isodiametric cells (figs. 6 and rg). While the primary wall layer forms its four layers, the plate of cells dorsal to the Sporangium also divides and takes on its tapetal character. The primary sporogenous cells contain large vacuoles, while their nuclei are near the middle (fig. 4) ; but even at this stage of development they grade into ordinary vegetative cells at the outer edge of the sporangium, so that it is difficult to determine just where they cease to be sporogenous cells ( fig. 3). Enough stages were seen in the other species mentioned to warrant the conclusion that the development of the pollen in them is the Same in general as in A. Cornuti. From this point the history was most completely followed in A. tuberosa. The primary sporogenous cells become the Spore mother cells without further division. This is indicated by the nature of the nuclei and by the form and number of the chromosomes. The size of the nucleus increases, and the chro- Matin collects in granular tangled threads, which together with 328 BOTANICAL GAZETTE [NOVEMBER the large nucleolus form the staining contents of the large transparent nuclear vacuole—the synapsis stage of the nucleus (fig. 7). This stage of the nucleus was also observed in Asclepias Cornuti, A. phytolaccoides, A. verticillata, and Acerates longifolia. At the beginning of mitosis the nucleolus often separates into several smaller nucleoli. The chromatin in A. éuberosa breaks into five chromosomes which split longitudinally while scattered in the nuclear vacuole (fig. 6). They are very small when split, but become much larger as the nuclear wall and the nucleoli disappear (fig. 6). After the disappearance of the nucleoli they are short, thick, and much larger than those in the vegetative cells. The number of chromosomes in the spore mother cells is approximately half that in the vegetative cells (compare fig. 6 with figs. 8and g). The exact number in the vegetative cells I was unable to determine. This reduction of chromosomes occurs in the spore mother cells of other plants. The mitotic phenomena attending the division of the cells referred to as spore mother cells closely agree, then, with those in the spore mother cells of other plants, and do not agree with those in the vegetative cells. This is to me conclusive evidence of their spore mother cell nature. Closely following this division, often before the cross wall is distinguishable, comes the second, with smaller chromosomes, which results in a row of four daughter cells representing a tetrad ( fig. zo). Both the rapid succession of these divisions and the forma- tion of four and only four cells from a mother cell homologize well with tetrad formation. - Each mother cell divides; those at the pointed upper end of the sporangium slope outward and downward, as shown in fig. 5, thus permitting the division of all the cells without great difference in size. The lower end of the sporangium is rounded, and hence the cells there are not greatly modified. These divisions I have no doubt are the ones Stras- burger saw in A. Syriaca, and this creates a doubt in my mind whether the mother cells near the ends of the sporangia formed only two microspores, as the report would lead one to believe. 3 eee : | 1901] DEVELOPMENT OF THE POLLEN 329 This formation of microspores in a row, while not the usual tetrad form, is not peculiar to the Asclepiadaceae. The cells are reported in other arrangements in Typha latifolia,’ Orchis mascula,? Juncus,’*? and some members of the Periplocoideae.™ The cells now become isodiametric, irregular, and crowded. Adjusting themselves to each other they soon lose all indication of their origin as a row of four. The protoplasm becomes filled with short coarse threads which stain readily and obscure the other cytoplasmic structures (fig. zz). The walls then thicken very rapidly. This is by the deposition of a layer from within the cells, as is evident later when intercellular spaces appear. When a mature pollinium is cut and squeezed the walls separat- ing the microspores split, and the microspores are squeezed out whole. Where a microspore is in contact with a pollinium wall it separates from it an inner layer (fig. 76,4). This suggests that most of the pollinium wall is of a different origin from the walls of the microspores. The nucleus divides after the walls are thickened, forming the generative and tube nuclei (fig. 73). The tube nucleus is much the larger and denser. The generative nucleus moves to the side of the cell ( fig. 74), and is cut off by a wall of considerable thickness (fig. 75). The tapetum gradu- ally disintegrates, leaving the pollinium loose in the anther, except for its attachment to the caudicle by its upper end. After the walls begin to thicken the thready protoplasm collects in masses within the cells (jig. 12). The masses disappear about the time of the formation of the generative and tube nuclei, and dense fusiform bodies composed of one or more Strands, and of unknown nature, make their appearance. In 4, verticillata they were short, thick, and smaller than in the other species. Their origin may be connected with the thready and dense protoplasm just preceding their appearance. They 8 SCHAFFNER, J. H., Development of the stamens and carpels of 7ypha /attfolia. Bor. Gaz. 24:93. 1897. °WILLE, Ueber die Entwickelungsgeschichte der Pollenkérner der Angio- spermen 39. 1886 7 Ibid. 42. = SCHUMANN, Engler and Prantl, Nat. Pflanzenfam. q?: 196. 1895. 33° BOTANICAL GAZETTE [NOVEMBER remained and were seen even in the pollen tubes. The pollen of all the species studied contained them, but they were most abundant in A. Cornuti. The tests for calcium oxalate, car- bonate,. phosphate, and sulfate, and for silicon and starch failed to reveal their nature. They stain like chromatin. A stage like that in fig. 76 was frequently found, and seems to indicate the fragmentation of the tube nucleus, although the series about it was not close enough to justify a definite con- clusion. Germination always bursts the pollinium along its outer edge, at the point where the diameter is greatest (fig. 77). A cross section shows the pollinium wall to be thinnest there. Germi- nation of the microspores was observed in A. Cornuti, A. tuberosa, and A, incarnata. SUMMARY. The development of the sporangia in the Asclepiadaceae studied is the same in general as in other plants, while there are no indications of the phylogenetic history of the reduction in number. The primary sporogenous cells without further divi- sion become the pollen mother cells. The latter divide each into four with the usual phenomena accompanying tetrad divi- sion, but through mutual adjustment and the close adherence of the microspores the evidences of grouping are lost. I wish to acknowledge my indebtedness to Dr. John M. Coulter and Dr. Charles J. Chamberlain for their kindly direc- tion and valuable suggestions. HULL Botanica, LABORATORY, THE UNIVERSITY oF CHICAGO. Note.— Just as this paper goes to press, one by Professor Strasburger, Zinige Bemerkungen 2u Pollenbildung bei Asclepias (Ber. deutsch. bot. Gesells. 19 : 450-454- 1901), dealing with Asclepias Cornuti and Cynanchum Vincetoxicum, has come to hand, confirming in a general way the results reported above. EXPLANATION OF PLATE XIII. The figures are reduced to one half their original size. The lenses used were Leitz objectives 3 and 7, and Zeiss *s oil immersion; oculars, Reichert BOTANICAL GAZETTE£, XXX PLATE X1il iS 0 Fy, Q q FRYE on POLLEN OF ASCLEPIAS CT TTR 1 rs tae £3" sll. en OE 1901] DEVELOPMENT OF THE POLLEN 331 1 and 2, and Zeiss compensating 4, 8, 12, and 18. All drawings were sketched with a Bausch and Lomb camera lucida, Figs. 1-4. 62| 9 . 42 8 me ’ 8 renchyma ......| 29), 19 16 4 Sclerenchyma..... 05 79 ih ) = of 3 Vase. bundles..... I Stele de 3 ; 4 i iq 1901 | DISTRIBUTION OF RED COLOR 339 WET, SUNNY PLACES, | Stem. Petiole. Leaf-blade. | Veins, | ae 15 7 10 6 e131) eee 10 8 3 I Oates Collenchyma..... 8 8 5 Sys I ; 4 4 . | Parenchyma...... 13 33 | 9 ae Ie 9] 8 8 a { ee ac a ee 2415 3 ia of 5 fc) asc. Bundles . fe) prele ee RAN eeeronetre et aioe sie: - WET, SHADY PLACES. Stem. Petiole. Leaf-blade. Veins, | 6 4 7 oO PINS cio. scarad 2 ATs af : a) Cortex s Collenchyma ..... at.% 15 | .? | ° 9 te) P. chyma Se TO a4 ed ae a 7 4 ~ cl i)" (a) a te) 2 Vasc. bundles..... I I — { tee 3 DISAPPEARANCE OF COLOR IN 54 PLANTS. Stem. Petiole. Leaf-blade. Veins. gasses tO O REIT 3 7 22 6 Hypodermis...... 7 5 3 2 Cortex os eros 12 13 | 10 enchyma..i <5 5 8 18 8 Sel enchyma..... Stele { Vasc, bundles. PR oi aes eee 2 fe) e) ° In frequency of color epidermis exceeds hypodermis or par- enchyma in stem and leaf-blade, but in each case the cortex shows the largest number of cases. In dry sunny places the epidermis has most color, the color gradually disappearing in the stem from the outside in. In dry shady and wet shady places the parenchyma exceeds, in wet sunny places the hypo- dermis or epidermis. It seems then as if sunlight tends to increase the red color in the epidermis or hypodermis. In cases of disappearance of color, the loss is greatest in the epidermis 34° BOTANICAL GAZETTE [NOVEMBER of the leaf-blade, and in the cortex of stem and petiole. In 26 cases of loss, the conditions had changed since spring from sunny to shady. This would make it seem as if red might be a protection from intense sunlight, but in 28 cases of appearance of color 13 were found in conditions which had changed from sunny to shady, and 5 were found in conditions which had changed from wet to dry. One might conclude that change of conditions favors change in color, from green to red or red to green, as has been asserted by others. Most of the plants with red color were gathered in dry sunny places. This does not accord with observations of Stahl, who finds in moist or shady places the most noticeable red col- oring. His theory does not seem applicable to the facts just given, for the conditions of sunlight and warmth in this case favor photosynthesis and conduction, also transpiration. The percentage is so large it could not be due to collecting more specimens in the spring or fall, when the temperature is lower, and thus be due to increased sugar concentration according to Overton’s theory. Taking the whole number of cases, the red color gives the largest percentages for stems and petioles, really the conducting parts of the shoot. One might think with Stahl that some advantage is derived from red color in photosynthesis and pas- sage of food material, but again why this is needed in most of these cases is not so clear, since conditions were already suffi- ciently favorable for transpiration, etc. The large amount of color in the cortex seems to favor Stahl’s theory also. If we apply Pick’s theory, that the red color protects the conducting parts from the injurious effects of light upon the changing sub- stances in them, we seem to have a strong argument in its favor, for the majority of the cases noted were recorded in dry, sunny places. In wet shady places only is the percentage of cases of red _ color in the leaves greater than in the stems and veins. In Stahl’s paper most attention is paid to color in leaves, so here is an agreement, if only a slight one, with his theory of the need of warmth to increase transpiration. UR eR I AIRoCtreRe ne none. a > me” oe 1901] ‘DISTRIBUTION OF RED COLOR 341 In stems, petioles, veins, and leaf-blades the largest percent- ages are for an even distribution of red color upon the surfaces of these parts. This would seem to show either that the shoot was evenly lighted or that some factor other than sunlight was the determining one in causing red color. However, the upper surfaces of the leaves, veins, and petioles show more color than under surfaces, and this accords with observations of others. Kerner’s theory of protection from too great light, and Stahl’s of increasing transpiration through the aid of red, apply here, although it is hard to see, as said before, why there is need of greater photosynthesis than is brought about by naturally favor- able conditions. Overton’s view is not applicable, for cool tem- perature is certainly not a factor in the conditions. The disap- pearance of red color in leaves when greater shade is present, and the great number of cases where red is located in the epi- dermis, are favorable to Kerner’s theory of protection. Another fact agreeing with the latter theory is in the case of upright stems, where one side, the side always toward the sun, Shows often the more color. This was noticed continually in Ambrosia. The red color is intensified at the joints of stems and peti- oles. This deepening of color, as well as that upon the upper surface of parts, is greatest in proportion in plants growing in wet shady situations. The theory of Stahl applies here, for the passing of food materials may be checked somewhat at the joint, where tubes pass into the stem; but Overton’s theory applies just as well, since conduction is delayed here and sugar must be concentrated in the cells of this region. Cool temperature does not appear here however. Any theory of protection from light is not applicable, since the joints are as much shielded as the other parts of the shoot. A large number of cases were found where the stem was very red near the ground. None of the theories seem to fit this fact. Cases where color is absent just at the joints and present upon the rest of the stem, where petioles are red near the blade 342 BOTANICAL GAZETTE [NOVEMBER only, where old tendrils are very red, where color is present in the hairs of leaves and not in the leaf surfaces, are unexplained by any theory ; also the appearance of red under the same con- ditions which usually favor the disappearance of color. Stahl has given a plausible explanation of color in plants of the moist tropics, where color is located in the cells of tissues near the surface. Overton explains autumnal and spring color- ation, but finds, although he can produce colorin internal tissues of the plant, he is seldom able to do so in epidermal layers. None of the explanations given apply to all the cases here recorded. In conclusion it is seen that the statistical observations pre- sented fit no one theory in all particulars. Some facts agree with no theory at all so far as known. The suggestion may be made that with further experimental study red color may be found subserving a number of purposes or having a number of different meanings; or, on the other hand, it may be due to an entirely unknown cause which may fit all cases. BOTANICAL LABORATORY, SMITH COLLEGE, Northampton, Mass. —- —— SOME PLANT ABNORMALITIES.! GEORGE HARRISON SHULL, THE investigation of the abnormal, either in the structure or function of organisms, is often of great value in arriving at cor- rect interpretation of normal conditions. But any specific abnormal form is rare as compared with the frequency of the normal condition, and as no one observer is likely to discover a great number of cases, it is important that all should be recorded. Already botanists have described very many instances of abnormal plant forms, so that the bibliography of plant tera- tology is already extensive. Thus Penzig? (1894) gives several thousand references to described cases of plant abnormalities. I. FASCIATION,. This is a common phenomenon, and so widely distributed that it has been observed by all who have come into any con- siderable contact with plants. The present cases are described here because of certain interesting attendant characters which may throw some light upon the nature of fasciation. A remarkable case of fasciation was observed in Lopéilon (Erigeron) Canadense L., where a stem had a breadth, in the dried State, of 8.5°™, and the overgrowth at the crest was so great as to throw it into marked undulations. The margins of this stem were apparently normal, as were also the leaves borne on them ; and one margin gave rise to a series of normal axillary branches. On the broad sides of the stem, however, the leaves were reduced toa narrowly linear form 0.5—2™™ by 2-4. This reduction was probably correlated with two other conditions, (1) the great crowding of the leaves, and (2) the greatly increased surface of the stem compared with its volume. It is certain *Contributions from the Biological Laboratory of Antioch College. No. 3. * 0. PENZIG, Pflanzen-teratologie. 2 vols, Genoa. *901) 343 344 BOTANICAL GAZETTE [ NOVEMBER that photosynthesis was carried on chiefly in the tissues of the stem itself, which had an unusually bright green color and deli- cate texture. No buds were produced in the axils of these reduced leaves until within 6-8™ of the crest, above which point each axil produced a sessile flower bud. The whole side of the stem was covered with numer- ous fine ridges which, on being traced to their origin, were found to originate in the midribs of the leaves. According to Masters? “the striae which these stems almost invariably present exhibit the lines of junction” between the stems by whose union the fasciated stem is formed ; but this certainly cannot be true of the striae in this case. In a fasciated stem of Echium vulgare L. there was found a greater width near the ground than at a point just below the widening at the crest. Here too the FIGs. 1, 2.—7, Leaf of Pelargonium sp. with striae are simply the grooves re blades. 2, Cross section through blades between ridges, which are of leaf shown in fg. 7, showing the reflection: ; all plainly traceable to the midribs of the leaves and are undoubtedly the lines beneath which lie the fibrovascular bundles originating in the leaves. By offering a different explanation of these striae the evidence of the union of stems in fasciation is lessened by so much, though not destroyed, since a union may exist without the existence of an evident line of union. 3 Vegetable Teratology 16. London, 1869. agent 1901 | SOME PLANT ABNORMALITIES 345 II, ABNORMAL FOLIAGE LEAVES. One of the most common abnormalities in Pelargonium is the formation of peltate and funnel-shaped leaves by the growth of leaf tissue where there is normally a sinus. On a specimen which had been observed to produce a number of peltate leaves, there was also found the exceedingly interesting form shown in figs. rand 2. The petiole was about 1.5 times as broad as a normal petiole and bore two perfectly formed blades. These blades were united from base to margin along a single vein and were placed with the under sides opposed to each other. This leaf gives an excellent illus- tration of Bateson’s*t (1894) law of reflection, 7. ¢., in the duplication of an organ the arrangement of the parts is the reverse of the arrange- — ment of homologous parts in the normal organ. So perfect was the reflection in this case that the more minute details of outline were reproduced almost as perfectly as in a mirror, In a leaf of fTicoria sp. (fig. iar collected several years ago by the writer, the terminal leaflet was so regularly and deeply lobed as to be almost compound. This modification was decidedly ‘‘progres- Sive,” although it occurred in the terminal leaflet, where, as ‘ oe BATESON, Material for the study of variation, 474-575. London and New or 1G. 3.— Leaf of Hicoria sp. showing a deeply lobed terminal leaflet. 346 BOTANICAL GAZETTE [NOVEMBER has been so well pointed out by Jackson’ (1899) the most primitive form is to be expected. There was a small gall just at the base of the leaflet, and as it lay close to the midrib and partially deranged its tissues, the peculiar form of this leaflet may have been due to a pathological condition. Ill. ABNORMAL FLORAL ORGANS. In the summer of 1900 a small plot of Lathyrus odoratus L. was found to be producing a considerable number of abnormal flowers. A few of these were dissected and the parts carefully drawn at the time ( figs. g-30). The inflorescence of this species usually has only three flowers, but occasionally varies from two to four. /ig. 4 represents a double inflorescence in which a two- flowered inflorescence has apparently arisen, axillary to the lower flower of a four-flowered inflorescence. Abnormal floral organs were numerous and interesting. I have shown the parts and arrangement of the normal flower in jigs. 5-7. The combi- nation of abnormalities in each flower can best be exhibited by description of the cases examined. In each case the floral organs not mentioned were normal. Case z. An outgrowth from the margin of the sinus between the upper calyx teeth assumed the form and coloring of the vexillum. Vexillatwo. (Fig, 8.) Case 2, Calyx with a petaloid outgrowth from the upper sinus. This outgrowth had a lanceolate form instead of the nearly orbicular form of the vexillum. Vexilla two. Stamens eleven, all united. Case 3. The upper edge of an upper calyx lobe dilated slightly and colored. Vexilla two, the outer having a downward hook like that of an ala, and somewhat narrower than normal. Stamens eleven, two nearly free at the base but united to the, other nine above. (Fig. 9.) Case 4. Calyx the same as in 7. Vexilla two. Stamens twelve, all united to form a tube. 5R. T. Jackson, Localized stages of development in plants and animals. Mem. Boston Soc. Nat. Hist. 5: 89-153. pls. 16-25. 1901] SOME PLANT ABNORMALITIES 347 11 15 FIGs. 4-16 —Abnormal floral organs in Lathyrus odoratus. For details see text. Case 5. Calyx normal in form but with a touch of color between the upper teeth. Vexilla two, the inner one having one margin enclosed within the carina. 348 BOTANICAL GAZETTE | NOVEMBER Case 6. Calyx with six teeth, and with a large expansion between the upper ones. One margin of this expansion was free and inserted within the calyx like a vexillum. This structure was probably a union of a vexillum with a petaloid outgrowth of the calyx, although no line of junction was apparent. Free vexillum none. Stamens ten, united into a split tube. (fig. 70.) Case 7. Calyx asin z. Vexilla two, one with a downward hook. Stamens eleven, united into a tube. Case 8. Calyx asin z. Rest of flower normal. Case 9. The tissue of two of the upper sinuses of the calyx was expanded into irregular petaloid bodies, the upper one with a divergent lateral lobe. Vexilla two, each having the form of half a normal vexillum, one with a prominent downward hook. Alae abnormally narrow, curved upward and outward, and lack- ing the usual overlapping edge. Stamens twelve, of which eleven were completely united, the twelfth partially adherent. (Figs. 11-13.) Case ro. Calyx with a slight petaloid expansion which was united edge to edge with the outer vexillum. Vexilla two, each having one margin enclosed in the carina. One ala external. (fig. 14.) ” Case rz. Calyx the same as in z. Vexilla two. One ala narrow and much curved upward. Stamens nine united, one free, and one nearly free. Case r2. Calyx with six teeth. Vexilla two, each with a lateral lobe, one lobe unusually large. (Fig. 15. Case 13. Calyx teeth six, the lower two narrower and more approximate than the others. Vexilla two. Stamens nine united, three free. Case 14. Vexillatwo. Stamens nine united, and two free. Case 15. Vexilla two, each with a lateral basal lobe on the upper side. (Fig. 76.) Casest6 and 17. Vexilla two, the second having one margin enclosed in the carina. Stamens ten united, one free. Case 18. Vexilla two, which were slightly united edge to edge at the base. Stamens diadelphous, two and nine. (fig. Z 7-) SOME PLANT ABNORMALITIES 349. Fics. 17-30.— Abnormal floral organs in Lathyrus odoratus. For details see text. Vexilla three, all alike and nearly normal in Case 79. form. Case 20, three free. (Fig. 78.) Case 21. Vexilla three, the middle one with one side folded Vexilla three, all united. Stamens nine united, 350 BOTANICAL GAZETTE | NOVEMBER back upon itself. Stamens triadelphous, one, two, and seven. (fig. 79.) Case 22. Vexillum with margins turned backward and coalesced, forming a funnel which was adherent to the calyx tube at the base. Stamens two free, nine normally united, with a twelfth united for two-thirds of its length. This filament was slightly enlarged and colored, and had its anther converted into a body of the same color and texture as the petals. (figs. 20, 21.) Case 23. Vexilla two, the second shaped like half the first and balanced on the other side by a petaloid stamen having a yellow spot, corresponding to an anther, on the inner margin. Other stamens normal, nine united and one free, the free stamen arising between the two petaloid bodies within the normal vexil- lum. (fig. 22.) Case 24. Vexillum doubled upon itself on both sides. Alae two on each side, the supernumerary alae having the hooks on the lower side, z.¢., toward the normal alae, in this respect appearing as a reflection of them. One of these supernumerary alae occupied the position of a free stamen. Stamens nine united, two free. (Fig. 23.) Case 25. Vexilla two, the first notched to one side of the center, the second with a large lateral lobe, and having one margin enclosed in the carina. The enclosed margin of the second vexillum bore a small appendage which was nearly spiral below and ended above in an erect wing. This appendage was probably the homologue of an anther. (Fig. 24.) Case 26. Vexilla two, the outer joined below, margin to margin, with one ala. Case 27. One ala joined to the first vexillum, this double petal external. Second vexillum of normal form, but with one margin included in the carina. Upper stamen free (as normal), beside it a broad filament with a lateral overlapping hook and colored like a petal. This broad filament was topped with an anther and had a normal filament branching from it at a point two-thirds of its length from the base. The remaining stamens were united into a split tube bearing upon its outer margin four a Ti 1901} SOME PLANT ABNORMALITIES 351 normal filaments, and one broader filament formed by the union of three filaments. This bore at its summit one sessile anther, and at each. side of this sessile anther a short-stalked one. ( Figs. 25-27.) Case 28. One ala external and lacking the usual hook. One side of the vexillum folded back and included in the carina. (Figs. 28, 29.) Case 29. Nine stamens united, the tenth united to the other nine for half the length of the tube, then spreading back with the form and coloring of a second vexillum. ( Fig. 30.) These abnormalities may be summarized as follows: 1. Abnormalities of the calyx. (a) In two cases there were found six calyx teeth, and (4) in eleven cases a petaloid inflation grew from the sinus between the upper teeth. This petaloid outgrowth varied from a small colored area in a nor- mally formed sinus to an expanse of tissue having the size, form, and color of a perfect vexillum. 2. Abnormalities of the vexillum. (a) In twenty instances there were two vexilla, and in three cases three vexilla were found. In about half, the second vexillum was normal in form, the variations from this being in two directions, (i) toward the wing with its downward hook on the one hand, and (ii) toward the stamen on the other. (4) In two cases there was a union between an ala and the outer vexillum. (c) Union of the vexilla was found in two instances. 3. Abnormalities of the alae. (a) In two flowers the alae Were unusually narrow and curved upward. (4) In two cases, as mentioned above, there was a union between an ala anda vexillum. (c) In one case there were two alae on each side. 4. Abnormalities of the stamens. (a2) An approach to the form and coloring of the petals. Almost every degree of peta- lody was found from a petaloid appendage or a broadened fila- ment with a hook, to the form of half a vexillum. (8) Three flowers had two free stamens, and two had three free stamens. (c) Thirteen had an unusual number of united stamens, mostly ten or eleven, in two instances reaching twelve. (d@) In several 352 BOTANICAL GAZETTE [NOVEMBER the stamens were triadelphous, one, two, and nine, or one, two, and seven. 5. Abnormal aestivation. (a) In three cases one ala was external to the vexillum. (6) In six instances the supernu- merary vexillum had one margin enclosed within the carina. (c) In several cases there was an abnormal folding of the vexillum upon itself. There have been but few cases recorded of the union of sepals in Clematis, and belonging as it does to a family which typically has all the parts distinct, the union of parts is all the more remarkable. Perhaps the most noteworthy case of this kind is that described by Jaeger® (1828), in which the sepals of Clematis Viticella L. were united into a bilabiate tubular perianth. Ch. Fermond? (1884) has found many cases of three-parted perianths in Clematis sp., and Cockerell® (1897) remarks upon the com- mon occurrence of the union of two sepals in C. ligusticifolia. In a specimen of Clematis growing at Edgewood, North Hamp- ton, O., said to be a seedling of C. Jackmani of the gardeners, there are found every year many cases of similar union of sepals, and in varying degrees of union, from the slight union of two sepals to the complete union of all the sepals into a regular, tubular, gamosepalous flower (figs. 37-34). This complete union, shown in fig. 34, does not occur often, but I have found two like the one figured. If this is a seedling of C. Jackmant, its variations are rather remarkable because of the almost abso- lute invariability of that form. Inthe “Star of India’*® Clem- atis there is a marked tendency to depart from the typical four-parted perianth, and to produce flowers having six sepals. In this species are found many cases of division or of union of sepals. ®Nov. Act. Acad. Caes. Leop. 1828 :641. /. 37. 7Essai de phytomorphie, ou étude des causes qui déterminent les principales .formes végétales. 2 vols. Paris. 8 Bot. Gaz. 24: 293. : ° This is the garden name. I have been unable to determine its specific relation- ship. 1901] SOME PLANT ABNORMALITIES 353 Two of the most striking cases of abnormality in Clematis are shown in figs. 35 and 36, although similar cases have been frequently described, indicating that the displacement and coloring of the leaves which are nearest the flowers is a common abnormality in the genus. The two figured were both collected from the same plant of the “Star of India” Clematis, although several seasons intervened between them. In fig. 35 there Fics. 31-34.— Abnormal floral organs in a seedling of Clematis Jachmant (of Sardeners); 37, normal flower; 32, two sepals united; 73, three sepals united; 34, th all the sepals united into a gamosepalous perianth. appears what at first seemed to be a reversion of the greater part of a sepal to an ordinary foliage leaf; the remainder retained the form, delicate texture, and rich coloring characteristic of the sepals. On the examination of the normal inflorescence, how- €ver, it appears that this organ, instead of being a partly reverted ‘Sepal, in reality is the result of a coalescence of a sepal and/a leaf from the node next below the flower. The chief evidences of this are found in the absence of one of the leaves from that node, and the presence of a marked decurrent line extending from the leafy organ down the side of the peduncle to the node. 354 BOTANICAL GAZETTE [NOVEMBER Fics. 35, 36.—Abnormal floral organs in “Star of India” Clematis (of garden- ers); 35, union of leaf and sepal; 36, union of peduncle and petiole, one leaf with leaflets partly colored (g), and one leaf entirely changed to a sepaloid body (¢g')- smi mall ul MM aaa ce ES ae ae 1901] SOME PLANT ABNORMALITIES 355 The bud which is normally axillary to the removed leaf failed to develop. fig. 36 is a somewhat similar case, but more complicated. Here, too, there is an adnation of the petiole of the leaf to the peduncle, and two of the three leaflets are partly colored like sepals. From the groove at the side of the coalesced petiole and peduncle arises an axillary branch. This branch bears, at the next node above, a leaf which in every respect resembles a sepal, and which failed to produce a bud in its axil. All the cases here described and figured are in the herbarium of the writer, except those of Lathyrus which were dissected, and the parts sketched and described from the fresh material. YELLOw Sprincs, O. BRIEFER ARTICLES. MEISSNER ON EVERGREEN NEEDLES. (WITH ONE FIGURE) A RECENT double number of the Botanische Zeitung” is devoted to a third treatise by Meissner on evergreen needles. This time the theme is chiefly the relation of the length of stem and needle, in sup- port of his previous attack on the idea advanced by Kraus,* and sup- ported by myself,3 that the thrifty growth of stem and leaf may be expected to occur together. Part I of Meissner’s paper contains in tabulated form numerous measurements of stems and needles of Pinus, Abies, and Picea. In a large part of these the variations in needle length are without any apparent relation to that of the stem. Nega- tive testimony of this kind is not very strong, for a more judicious than judicial selection of material can make it show almost anything. For instance, Meissner (p. 30 ef seg.) would prove by it that I am unreasonable in expecting the different parts of one tree to vary alike in the growth of stems and needles.* While it must be said that the frequency of exceptions has surprised me, it has still seemed to me in the observation and measurement of conifers, wild and cultivated, in various parts of the United States, that in a great majority of instances the stems and needles vary in length together. But perhaps Meissner would have drawn the opposite conclusion in the same forests, with an equal desire to see things as they are. Results obtained by the statis- tical method can hardly ever have the weight of those yielded by judicious experimentation. Part II deals with mutilation experiments. As would be expected, if the mutilation compels the tree to devote its plastic material to the *Ueber das Verhiltniss von Stamm- und Nadellange bei einigen Coniferen. Bot. Zeit. 59:25. 1901. * Abhandl. d. naturf. Gesells. zu Halle 16 : 363. 1886. 3A biological note on the size of evergreen needles. Bot. Gaz. 25 : 427: 1898 ‘The proportionate length of stem and needle on the main axis and its branches in one season is an altogether different subject. 356 [NOVEMBER aaa ENS Igor] BRIEFER ARTICLES 357 development of a few members, their growth will be stimulated: while robbing it of plastic material or the chance to form, it results in a stunted growth. Part III, on the annual period of growth of stems and leaves, is the most valuable part of the paper. Meissner shows by measurements that the chief growths in length of stems and needles fall at different times. In Pinus the stem almost finishes its season’s elongation, while the needles remain relatively short, the relations in Abies and Picea being just the reverse of this. The dry weights of stems and needles at various times during the months of growth vary in general with the lengths. A moment’s reflection will satisfy anybody who has seen evergreens grow that Meissner is correct. Transitory unfavorable conditions early in the season will therefore tend to hinder the growth of the stem of Pinus more than that of the leaves. The maximum growths of stems and needles at different times is a physiological explanation of the cases in which short year’s-stems bear long nee- dles, or vice versa. But while this makes exceptions intelligible, I am by no means ready to agree that there is no connection between the Sizes of stems and needles, or that Kraus and I were wrong in expect- ing both to vary, in nature, with the general condition of the plant. Neither of us could have thought of maintaining that leaves and stems could never vary independently ; that external factors, such as light, must directly influence both alike. Trees do not respond at once to changes in the weather to such an extent that they will often be Stunted in May and thrive in June, or grow luxuriantly in May and then abruptly halt. At least my observations on the whole resultant growth indicate that they do not. The influence of bad conditions on the stages of growth previous to active enlargement has been left largely out of account, and may appropriately receive a word here in the light of Meissner’s results. Nn my paper the area of cross sections of the needles was given, and was always least the season following transplanting. The difference was due to the size of the cells, the number not being materially different from that of other years. Thus in leaves of Pinus Austriaca (cf. doc. cit. p. 433) the number of cells in cross section was as follows: 1895 1896 1897 PRIN Wo ue cekaas ot 176 | 176 | 246 Outer parenchyma layer..| 77 7 96 Bndodermisc.4 3...2250% 39 38 48 é 358 BOTANICAL GAZETTE [NOVEMBER The number of cells was surprisingly the same in 1895 and 1896, though the difference in size was very great. The leaves of 1897 were exceptionally thrifty in all respects. The larger leaves contained more resin ducts. This plant was transplanted rather late, when the leaves CONCOMITANT DWARFING OF STEMS AND LEAVES OF CONIFERS. I. Picea alba; 2. Abies balsamea; 3. Pinus Strobus; 4. Tsuga Canadensis ; 5. Picea pungens; 6. Picea nigra; 7. Pinus Austriaca; 8. Taxus baccata. of deciduous trees were beginning to grow, probably about the first of April. The exact date is not known, and is not important because it would indicate a definite stage in development only for the one yeat and locality. The fact that the trees did not have their growth checked = Late a tice 1901] BRIEFER ARTICLES 359 before the first phase of the season’s growth is indicated not only by the full number of cells, but also by the presence usually of at least the normal number of needles (doc. ct#. p. 429). There are a number of trees of Picea excelsa on the campus of West Virginia University that were transplanted in 1899, several weeks earlier in the season than the Bloomington trees. While the size of the cells is not nearly proportional to that of the needles, the number formed during the spring following transplanting falls somewhat behind. The numbers in an average case are as follows: 1898 1899 1900 B piderniisiec cc. leaaet: 122 II4 140 Outer parenchymalayer..| 91 80 106 Mipdodetmii 5 Gao canes a8 18 25 The whole question of the growth of the needle will reward more experimental study, and Meissner’s interesting paper should stimulate it. The accompanying photograph is of the plants described by me in 1898, and will show the concomitant dwarfing of stems and needles better perhaps than tables of measurements did.— Epwin BINGHAM CopeLann, Monroe, Wis. THE INSTABILITY OF THE ROCHESTER NOMENCLATURE. SINCE the publication of my article in the BoranicaL GAZETTE for last March* three communications have appeared which discuss certain details there presented. Two of these letters, from Mr. C. L. Pollard? and from Professor L. M. Underwood,? question the significance of facts presented by me in parallel columns ; while a third letter, from Professor N. L. Britton,* presents at more length than was done by me the practical reasons why the first species of a complex genus should not necessarily be treated as the type. These articles bearing directly upon the vital question of stability in botanical nomenclature canst leave the general reader in some doubt as to the exact force of certain arguments I have already discussed. It may not be out of place, there- fore, to ask for a brief consideration of the points thus newly emphasized. *BoT. GAz, 31: 183. Igor. 3 [bid. 31 : 365. 190F. * [bid, 31 : 285. 1901. 4Science 13 : 588. I90t. 360 BOTANICAL GAZETTE [| NOVEMBER Mr. Pollard quotes from my article certain sentences embodying conclusions which he characterizes as misleading. The conclusions thus referred to were to the effect that in 1900 Professor Underwood, following logically and consistently the Rochester Code, used 25 per cent. of names different from those used by him in 1896, when he like- wise asserted that he was following the Rochester Code. Mr. Pollard’s contention is that this fact does not afford a proper basis for criticism of the Rochester Code, because most of the changes made by Professor Underwood represent very recent segregations among the ferns, com- parable with the divisions made within three years in the genus Anten- naria. Yet it does not seem to have occurred to Mr. Pollard that, in omitting the footnote to the paragraph quoted, he is possibly mislead- ing that large body of readers who habitually neglect to verify quotations and references. Had he carefully read and quoted the two sentences as they were originally published with the footnote: ‘The true ferns alone are here considered, and the genus Botrychium is purposely omitted, since that genus has been subdivided by Professor Underwood to such an extent that comparative figures would have little definite significance,” he would have found it unnecessary to inquire whether I do not “recognize the necessity for occasional segregations.”’ Every working systematist must recognize such necessity ; but the segregation of perplexing polymorphous types is a very different thing from the raking up of older and more or less obscure appellations for plants the names of which have long been established. The latter is a nomen- clatorial matter, the former botanical; and if anyone has read into my previous remarks the least opposition to such division of confused groups as shall lead to a clearer understanding of the forms, he has found that which [ in no way intended. But most of the ferns under discussion cannot be placed by Mr. Pollard under the same category as the recent segregates of Antennaria (and likewise Botrychium), for they do not represent new points of view unknown in 1896. Matteuccia is a name given in 1866 to Willdenow’s Struthiopteris, which was segregated in 1809 from Osmunda; our Dennstaedtia was separated from Dicksonia in 1857; Filix was pub- lished by Adanson in his well known Famille des Plantes in 17633 Phyllitis was re-distinguished in 1844; and it is certainly not a new idea to treat species of the complex genus Aspidium under the name Polystichum. Neither are Phegopteris Robertiana and Notholaena dealbata now treated as species for the first time. The former was published Igor] BRIEFER ARTICLES 361 by Hoffmann in 1795 as Polypodium Robertianum, the latter by Pursh in 1814 as Chetlanthes dealbata. In 1896 Professor Underwood had already published five editions of a work on North American ferns, and he was consequently in a position to know of the earlier conceptions and treatments of these plants. At that time he was publishing accord- ing to his interpretation of the Rochester Code, and the names thus published, we were assured, were those which would stand. Yet after having established (as he supposed) these names according to strict priority principles he has found occasion to alter 25 per cent. of them. Has his application of the Rochester Code brought uniformity ? In his “ Open Letter ” Professor Underwood shows very conclusively that “when its bald statements are unqualified,” “the deadly parallel” “seems to mean more than the facts will warrant.” For, avowedly with the purpose of showing that “the personal preference hit-or-miss system of Kew and Berlin” is bringing us unwarranted confusion in plant names, he places in parallel columns twenty-one fern names which appeared in the first edition of Gray’s Manual and the twenty-one more or less dissimilar names for the same plants in the séxrA edition. By the thoughtless reader, with “its bald statements . . . . unquali- fied,” this comparison might seem completely to dispose of my criticism of the recent changes of names made by Professor Underwood. Yet it should be borne in mind that such comparisons may “mean more than the facts will warrant.” Professor Underwood implies that the application of the combined Berlin and Kew rules is responsible for the changes between the first and the sixth editions of the Manuad. Can it be that he “forgot” that the first edition was issued in 1848, the Jfth edition (Pteridophyta by D. C. Eaton) in May, 1867, and the sixth edition in 1889, a// before the Berlin rules were formulated in 1897? It was three months after the fifth edition of the M/anua/ went to press that the Paris convention of 1867 was held and the DeCandol- lean Code was drawn up. Consequently the sixth edition (1889), by ‘Watson and Coulter, was the only one published after the adoption of the Paris Code of 1867. Furthermore, since even this edition was published eight years before the rules of the Berlin botanists, a com- Parison of the first edition (1848) and this sixth edition (1889) is no very logical proof that the Berlin rule of 1897 is “ unseaworthy.” The fact is, that the names in no edition of Gray’s Manual have been based on the combination of the Berlin rule for genera and the so-called Kew rule for species, which was recently advocated by me. 362 BOTANICAL GAZETTE [NOVEMBER This simple and definite method of obtaining a uniform system of names has been carefully tested, however, during the past half decade, and it has appealed as practical to many besides the “ trans-Carlines.” Let us now consider, on the other hand, the points raised by Pro- fessor Britton. The first rule of the Rochester Code reads: ‘‘ Priority of publication is to be regarded as the fundamental principle of botanical nomenclature.” The fifth rule that ‘ Puddcation of a genus consists only (1) in the distribution of a printed description of the genus named ; (2) in the publication of the name of the genus and the citation of one or more previously published species as examples or types of the genus, with or without a diagnosis” [italics ours]. A recom- mendation adopted by the reformers at Madison reads: “In determin- ing the name of a genus or species to which two or more names have been given by an author in the same volume or on the same page of a volume precedence shall decide.” By Professor Underwood, as already emphasized in the March GazettE, this principle of strict priority has been applied in the determination of the type of a complex genus. Professor E. L. Greene, likewise, has made important changes based upon its rigid enforcement. Professor Britton, too, has more than once maintained that priority of place (precedence) is final, saying, in regard to the now famous case of Buda and Tissa: “I accepted Z7%ssa rather than Buda for the simple reason that it stands first on the page in Adanson’s ‘Familles.’ That is priority, 1am sure” [italics ours].3 He has further said that this principle (priority of position) will be accepted by those who have recognized the necessity of adopting methods of procedure which will render the system of nomenclature stable.”® And again, “the number of cases in which change is desirable by reason of priority of place is not great.”? Thus, in 1890, Professor Britton clearly defined what he meant by priority. In 1892, “after a very careful consideration,” the Rochester Code was formulated ; and in 1893, after plenty of time for further deliberation on its fundamental prin- ciple, the code was augmented by the recommendation above quoted. Priority has been talked of until the subject has become a tedious one; and that it is the ruling principle of the reformers has been so — often avowed as to become axiomatic. It is not surprising, then, to find, in the ///ustrated Flora in 1896, the statement that “ its [ priority ’s] S Jour, Bot. 28: 295. 1890. *Jour. Bot. 28 : 371. 1890. 7 Jour. Bot. 28 : 372. 1890. _ f90r] BRIEFER ARTICLES 363 adoption is the only practicable way of securing stability to the origi- nal names.’’ This idea has permeated the writings of the reformers, and they have over and over again asserted in books and papers for amateurs and “the younger generation” that the names they advo- cate are alone the ones which can stand. That the principle of strict priority (the fundamental law of the Rochester Code) has been many times ignored by those who claim to follow it was sufficiently emphasized in the March Gazette. Indeed, when Professor Greene pointed out,’ in 1896, that the principle of priority demanded that the first species of a complex genus be taken as the type, he was merely showing the logical outcome of the prin- ciple. When again, in 1899, Professor Underwood followed the Same interpretation in his Review of the Genera of Ferns, he was merely following conscientiously the fundamental principle of the Rochester Code, and the law so often vigorously defended by Professor Britton, who “accepted Tissa rather than Buda for the simple reason that it stands first on the page in Adanson’s ‘Familles,’” saying, “That is priority, I am sure.” After such clear definition of the principle of priority by Professors Britton, Greene, and Underwood, we are now amazed to see from Professor Britton’s pen a convincing argument against the uniform selection of the first species as the type of a complex genus. Where Shall we look for that long-promised “uniformity” when, in writing of the principle as logically followed by Professor Underwood and more than once by Professor Greene and by Mr. Thomas Howell, Professor Britton now says, “‘Inasmuch as a great many genera have at their first publication been made to include more than one species, and in a large number of instances some of these, often the first in Position, have been used by subsequent authors as the types of addi- tional genera, this latter-day proposition affects an enormously greater number of cases than those which fall properly under the operation of the rule’ ?" And when, continuing his argument against the neces- Sary acceptance of the first species as the generic type, he says “dt is, therefore, clear that there is nothing logical in the proposed extension of the principle” [italics ours],* does he not directly contradict the con- clusions of Professor Greene and Professor Underwood? Further- more, how are such views reconciled with the fundamental principle * Pittonia 3: 128. 1896. Science 13 : 588. 1901. ° Mem. Torr, Bot. Club 6: 247. 1899. Science 13 : 588. 1901. 364 BOTANICAL GAZETTE [ NOVEMBER of the Rochester Code, and with the former statements of Professor Britton himself, who in 1890 emphatically argued of “priority of place,” that “that is priority, I am sure” ? When the Rochester Code and its Madison amendments were put forward we were repeatedly told that their ratson d’étre was to estab- lish our plant-names upon a permament basis. Therefore when the Check List was published it was naturally supposed that the names there included were final. At great inconvenience to all branches of Ameri- can botany we have been forced, consequently, to stumble through a most perplexing tangle of ever-changing “permanent” names. [If all this confusion and inconvenience were leading us by the shortest course —or by any course—to stability, no one but the short-sighted would complain. It is true that, in spite of their frequent changes from one policy to another, and notwithstanding the utter abandon with which they trifle with the most important principles, the reformers still claim to be bringing us uniformity. Howcan this be? Professor Underwood stands firmly for priority of place, claiming that the first species in the genus must be taken as the type. Professor Greene has taken the same ground, though openly arguing against some other principles of the Rochester Code. Now, after agitating for years the principle of strict priority and more than once defining his understanding of the term, Professor Britton has published an argument squarely opposed ‘to the uniform acceptance of the first species of a complex genus as the type. Can it be, then, that with such hopeless diversity of opinion on the part of two leading reformers at the same university, they still have sufficient sense of humor to tell us that they are establishing unifor- mity? If Professor Underwood, following consistently the spirit and text of the Rochester Code, believes in changing “99 per cent.” of names; and Professor Britton, abandoning the fundamental principle for which he has so long argued, changes names on a radically different basis, will they not give us systems of names more and more hopelessly nlike ? Professor Britton declares the position taken by Professor Under- wood a “latter-day proposition,” thus implying that the question of generic types was not seriously considered at Rochester and at Madi- son. Yet if the Rochester Code ig the result of a “ very careful con- sideration,” how can this vital matter have been overlooked? How- ever, the Rochester Code tells us that “ priority of publication is to be regarded as the fundamental principle of botanical nomenclature,” Sali ono Igor | BRIEFER ARTICLES 365 and Professor Britton has said “it is perfectly clear that as long as we allow ourselves a choice of names im any way, so long will authors ditfer in their acceptance and the settling of this important matter be deferred” [italics ours].* From these assertions the only logical conclusion is that doubtful cases are to be referred to the principle of priority. The decision between Buda and Tissa, and the determination of the generic type both “allow .... a choice of names,” and according to the first principle of the Rochester Code priority alone should settle them. The question of Buda and Tissa is one of decision between two generic names for the same plant; the question of the generic type asks which of two or more plants shall bear a given generic name. When these questions have such fundamental simi- larity, how can a reformer maintain that priority is to decide in one case and not in the other ? When Professor Britton now maintains that the first species of a complex genus is not necessarily the type, and that in such cases the Rochester code allows us to cling to the traditional genera, he at once places himself on record as likewise opposed to the fourth rule of the Rochester Code. This rule reads : IV. Homonyms.—The publication of a generic name or a binomial invalidates the use of the same name for any subsequently published genus Or species respectively. Let us look, for example, at the case of Mimosa, a name which Professor Britton and other reformers use in its traditional sense. The Linnzean Mimosa, published in the Species Plantarum, contained 39 species, the first on page 516, the last on page 523. Of these Species only six are now retained in the genus as finally defined “y entham in 1875 and now generally accepted. The first species now recognized in the Benthamian genus Mimosa is M. viva, no. 11 of the Species Plantarum. This species is the last one on page 517, and on that page it is preceded by six species, and on page 516 by four Species, all of which are now treated as members of other genera If Tissa has priority over Buda “for the simple reason that ie scans first on the page,” then surely by the same logic Mimosa bigemina and the other five species on page 517 have priority over M7. vrva. And still more clearly J. viva is preceded by M. Ledédeck (now treated as an Albizzia) and the other species on page 516. Mimosa was pub- lished on this page (516) of the Species Plantarum as the name of a ™ Jour. Bot. 28: 372. 1890. 366 BOTANICAL GAZETTE . [ NOVEMBER genus; then JZ. Leddeck, M. Inga, M. fagifolia, and M. nodosa were published, not only with citations of their previous places of publica- tion, but with brief diagnoses as well. Consequently the publication of the generic name Mimosa and the first of the species, M. Ledbeck, satisfies the Rochester requirement for the publication of a genus (see above). Furthermore, the species was treated as a Mimosa subsequent to its original publication in 1753, and it was first removed from the genus by Willdenow in 1806. Therefore, when the name Mimosa in the Benthamian sense is applied by Professor Britton to M/. viva (and its congeners) while on a preceding page the same generic name had been applied to M. Lebbeck, a plant of different generic affinity, he uses a name which is a homonym. Two fundamental principles of the Rochester Code are thus deserted by the chairman of the Rochester committee on nomenclature, while strongly defended by only one of the original members. And now from Nebraska, which has not long been notable as the seat of ultra-conservatism, comes the announcement, in regard to the recent work of the only consistent member of the original committee, that “it shakes one’s faith in the immutability of things to find old friends under unfamiliar names.” * The reformers wedded themselves to the principle of strict pri- ority. At the start they ignored the reasoning of those who foresaw where it would lead them. They rushed headlong and short-sighted into the scramble for hidden and obscure names. For a short time they worked harmoniously. Then came misunderstanding and lack of unity. One member of the nomenclature committee disclaims “that there is any obligation to elevate varietal names to specific rank when ‘the [aggregate] varieties themselves are thus promoted.” In so doing he opposes the rule of the Check List committee which says “that the original name is to be maintained whether published as species, subspecies, or variety.” Another member stands for absolute priority and takes a course in which Professor Britton finds ‘nothing logical.” The chairman of the committee now proclaims that priority of place (precedence) is not necessarily final. In so doing he opposes not only the first, but the fourth of the Rochester principles. To every logical and fair-minded scientist who will take the trouble to consider the question calmly and judicially, this woeful diversity in the practices of the committee must be apparent. No one can say 3 Science 13 : 833. Igor. ™%4 GREENE, E. L., Pittonia 4:253- 1901. Igor | BRIEFER ARTICLES 367 today where individual members of the Check List committee will stand tomorrow. ‘They have forgotten or ignored the fact that the Roches- ter Code was to give us permanent names. They have made of it a “personal preference hit-or-miss system.” In the words of one of their number, they are “openly at war with their own rules.” Is this uniformity ? Is this “the day of law”? Is this the high road to a stable nomenclature ? Do we sincerely want uniformity, or do we prefer the tangled results of individual interpretation ? If the former ideal still appeals to us, why not abandon this restless pursuit of the will-o’-the-wisp ? Why not honestly test the combined Berlin rule for genera and Kew rule for species? None of their opponents have given them a fair trial. Until they do can they really judge of their merits ?—M. L. FERNALD, Gray Herbarium, June, 1901. FLOWER VISITS OF OLIGOTROPIC BEES. III. Amonc the oligotropic bees mentioned in BoranicaL GAZETTE 28: 36, 215, and 30: 130, should be included: Andrena krigtana, which collects its pollen from Xvigia amplexicaulis; Entechnia taurea, which is an oligotropic visitor of Jpomoca pandurata: and Anthedon compta, which gets its pollen exclusively from Oe¢nothera biennis. Species of Melissodes, which usually collect the fine pollen of Compositae, have their scopae dense and quite closely plumose. On the other hand, Emphor, Xenoglossa and Entechnia, which collect the large pollen grains of Aidiscus lasiocarpus, Cucurbita pepo, and Lpomoea pandurata, have their scopae quite loose and thinly plumose. The close relationship of Anthedon to Melissodes, and the fact that the male has quite plumose hairs on his hind tibiae, show that the Scopae of the female have recently lost the barbs and have come to be composed of simple bristles. I have wondered why this was 0, and have expected to find some peculiarity in the pollen which the bee collects. Now in Oenothera biennis the pollen grains are large, tri- lobed, and connected by cobwebby threads. This condition of the pollen makes the barbs unnecessary if they would not greatly interfere with the collection of this kind of pollen. Andrena nasonit, mentioned in the first list, is not oligotropic. In the Fertilization of Flowers, p. 570, in discussing -— effect of conspicuousness of flowers in inducing insect visits, Miiller says: 368 BOTANICAL GAZETTE [NOVEMBER “The most important deduction to be drawn from them is, that in general anthophilous insects are not limited by hereditary instinct to certain flowers, but that they wander about getting their food on what- ever flowers they find it. For if each insect had its own species of flower, as most caterpillars have their own particular food plant, the abundance of insect visits to the plant would not depend at all upon its conspicuousness.” Then, after mentioning the case of Andrena florea and Bryonia dioica, etc., he says: “ But these insects do not form 1 per cent. of all the species that I have observed, and even of these cases the restriction is only complete in two.” In my neighborhood, excluding the inquilines, which do not make nests, 30 per cent. of the bees are oligotropic.— CHARLES ROBERTSON, Carlinville, Ill. oe ay CURRENT LITERATURE. BOOK REVIEWS. Latex and mucilage. TO THE already extensive literature of the laticiferou; tissues, Molisch contributes an important addition,* differing from its predecessors in giving chief attention to the constitution, both organic and chemical, of the latex itself. In his investigations he used living material and fresh latex, as well as that which had been fixed and stained. Molisch has confirmed and extended the earlier observations of Treub, Johow, and Schmidt on the exist- ence of a plasmic membrane and nuclei in the latex tubes, for demonstrating which he recommends Euphorbia splendens and Poinsettia pucherrima. He finds the membrane lining the latex vessel and constituting an inner living tube within which the latex is formed like the cell sap. Special examination of the nuclei shows that some nuclei are very different from those of ordinary plant cells, having characters not before known in nuclei of any plant or animal. Molisch calls them Blasenkerne. The granular nucleus seems to lie centrally or excentrically in a relatively large globular vacuole, but really the vacuole is in the nucleus, the nuclear sap filling the space between the nuclear material and the membrane. Nuclei of somewhat irregular form are also present. Various phenomena lead to the conclusion that the nuclear membrane is an independent, clearly differentiated organ. Besides the nuclei there are imbedded in the plasma leucoplasts of dif- ferent kinds. Some form the elongated starch grains; others (proteino- Plasts”) produce proteid granules, a phenomenon which has recently been observed also by Heinricher in Lathraea. Crystals of proteids or proteid-like substances are also produced, not anywhere in the contents of the latex tubes, but by the agency of special plastids or of vacuoles. Molisch also finds elaioplasts and vacuoles responsible for the formation of oil drops. Into the details of the chemistry of the latex we cannot follow the author. The latex he finds usually acid, rarely neutral, and never alkaline; calcium Salts and chlorids are variable in amount ; magnesium compounds are abun- dant and sometimes accumulate in extraordinary quantity; proteids and Carbohydrates are so abundant that one must look upon the latex tubes as Special reservoirs of these foods. : As the latex is an emulsion, the fine division and 1 ly great *MoLiscu, Hans: Studien iiber den Milchsaft und Schleimsaft der Pflanzen. 8vo, pp. viii 111, figs. 33. Jena: Gustav Fischer. 1901. M4. Igor] 369 37° BOTANICAL GAZETTE [NOVEMBER of its constituents probably facilitates the absorption of gases and metabolism generally, to an extent hitherto unnoticed. On the réle of the latex the author promises further publication, Because the mucilage tubes of the Liliaceae, Amaryllidaceae, and Commelynaceae are analogous to the latex tubes, Molisch has investigated them. He finds extraordinary nuclei in some of them—filaments I5004 long by 0.1-0.3 in diameter ; also proteid crystalloids, starch, glucose, and tannins, as in latex tubes, besides a new body, luteofilin, which occurs as sphere crystals in the mucilage of many monocots.—C. R A manual of bacteriology. IN 1897, Frederick D. Chester published in the Annual Report of the Dela- ware Agriculture Experiment Station a preliminary arrangement of the spe- cies of the genus Bacterium. This work, rearranged and enlarged to include all the groups of bacteria, has now appeared in a valuable Manual of deter- minative bacteriology.2 While not so voluminous as Migula’s great work on systematic bacteriology, this book is by far the most complete classification in English, conprising descriptions of some 780 forms. The system of classifi- cation adopted, by means of which related forms may be readily traced out or new species identified, is the same as that first proposed by Migula in Engler and Prantl’s Natirlichen Pflanzenfamilien (1896), with some minor modifications. In the synopsis of Bacterium and Bacillus, coloration by Gram’s method is used as an important differential test. This might be open to criticism, for variation of the Gram staining reaction within a so-called group is well known. It was brought out in a recent study of B. pyocyaneus by Roger C. Perkins as follows: “In reactions of the various organisms to Gram’s stain, my results did not coincide with those of Jordan and Ruzicka, who note complete decolorization in every case. Of ten varieties studied in this present series, seven decolorized uniformly and regularly when treated by this method, but three retained the color at every trial.” 3 On the other hand, Chester has given a subordinate place to formation of gas in the dif- ferent sugar bouillons, a comparatively constant reaction. In the section devoted to the terminology of descriptive bacteriology, the author has arranged and illustrated an excellent series of simple terms, capable of expressing definitely in one word the meaning of several sentences of the old style verbose and figurative cultural description. These terms are interesting additions to the bacteriological vocabulary. The criticism of species nomenclature is a point well taken, alt though no adequate suggestion is made as to how names of forms so closely related as the various kinds of * CHESTER, FREDERICK D., A manual of determinative bacteriology. 8v0, pP- vi-+ 401, figs. 73. New York: "The Macmillan Co. - $2.60. 3 Jour. Med. Research 281. 1901. 1901] CURRENT LITERATURE 375 fluorescent bacteria could be made indicative of that fact and yet avoid the trinomial term. What faults may be found in the book, however, are of minor importance in comparison with its value asa reference book, and a supple- ment to the text-book of every student in a bacteriological laboratory. — MARY HEFFERAN, The flora of Alabama.‘ Dr. CHARLES Mone has left behind him a most substantial monument, The bulky volume before us contains the botanical recerds of ‘forty years of sojourn and wanderings” through the state of Alabama. It may be added that the “wanderings” were by no means aimless, but were those of a keen and tireless observer, Such a mass of observations by a single man is the possession of no other state. It isa pleasure to note that the author was per- mitted to complete the organization of his notes of a lifetime into permanent and usable form, The book presents the patient study of a great and interesting area, not by the perfunctory cataloguing of species collected, but by the discussion of the broad biological features which have determined the flora and its distri- bution. The author evidently fully appreciated the newer aspects of the problems of floras, and has presented to us, in terms of Merriam’s life zones and Warming’s plant associations, the general ecologic and floristic features of Alabama. The general discussion occupies 137 pages, and is full of material for the student of phytogeography. After some preliminary historical material, in which the work of such pioneers as Bartram, Buckley, Gates, Peters, Beau- mont, and Nevius, are fully noted, the general physiographic features of the State are presented under topography and geology, river systems, and li- mate. Then follows an account of the general principles of plant distribu- tion, the significance of life zones and of plant associations and formations being explained. These principles are then applied to the flora of Alabama, which is presented in its general character and distribution. vl The ecologic relations are considered under the following titles : forest flora, open land or campestrian flora, water and swamp flora, organotopic flora (epiphytic, saprophytic, parasitic, and insectivorous plants), and intro- duced plants and their influence upon native plant associations. The distri- bution falls naturally under the two general heads of the Carolinian and Louisianian areas ; the former including the mountain region, the table-lands of the Warrior and Coosa basins, the region of the Tennessee river valley, and the region of the lower hill country; the latter including the region of *Mour, CHARLES: Plant life of Alabama. An account of the distribution, modes of association, and adaptations of the flora of Alabama, together with a sys- tematic catalogue of the plants growing in the state. Contrib. U.S. Nat. Herb. G: I-921. gis, I-13. 31 Jy. 1901. 372 BOTANICAL GAZETTE [NOVEMBER the central pine belt, the central prairie region, and the maritime pine region. The systematic catalogue occupies 682 pages, and is a model of pains- taking care in the way of bibliography, synonymy, and range. The thallo- phytes number 1722, and notably full is the list of fungi by F. S. Earle, and the list of lichens. The bryophytes number 166, the pteridophytes Iog, the gymnosperms 13, the monocotyledons 681, and the dicotyledons 1782. total enumeration of species and varieties is 4473. About ten new species are described, but very many described elsewhere are founded on Dr. Mohr’s material. The three endemic species are 7yichomanes Petersii, Croton Ala- bamensis, and Neviusia Alabamensis. The sequence is that of Engler and Prantl, and the nomenclature is that of the Rochester code. It is a matter of great regret that the author was not spared long enough to receive the congratulations of his associates upon the appearance of his monumental work.— Methods in plant histology.® As STATED in the preface, this work has grown out of a course in histolog- ical technique given by the author to his classes in the University of Chicago and to its non-resident students taking work in this subject through the Extension Division. A series of articles on the same subject published by the author in the Journal of Applied cade forms the basis of the present volume. The first part of the book, about one-third, is devoted to the discussion of apparatus, reagents, and most of the important methods of killing and fixing, staining, sectioning, and mounting plant tissues and the lower forms of plant life. The chapter on apparatus is short, and much could be intro- duced that would enable the student who has not a complete laboratory equipment before him to save time and material and also produce more perfect results. The chapter on reagents seems unnecessary, as most of the matter is repeated elsewhere in the book. The description of the paraffin method is most complete and very carefully prepared, while the treatment of the celloidin method is hardly adequate, and does not give the more recent improvements that contribute so much to the successful use of this method in plant histology. The treatment of the methods of killing, fixing, and staining is admirable. Part II is devoted to the study of types of the various groups of plants systematically arranged and illustrated by specimens easily collected by any one familiar with the main divisions of the plant kingdom. The directions for the treatment of the material used in these studies are excellent, and the ‘CHAMBERLAIN, CHARLES J.: Methods in plant histology. 8vo. pp- Vi--159- figs. 74. Chicago: The University of Chicago Press. 1901. $1.50 1900] CURRENT LITERATURE 373 specimens judiciously selected. The treatment of the Rule is the most comprehensive in the selection of material and in its preparation. The author, by his careful discussion of the various methods in the preparation of material emphasizes what is too often neglected in general histological studies, particularly of the lower forms, the careful preparation of material for examination. Much of the histological work with classes is misleading and useless because the teacher fails to appreciate this point. The student who learns to make a proper use of the best methods early in his course will be well repaid when at a later period he begins more critical studies. A convenient list of formulae for reagents is placed at the end of the volume. The technique of the book is good, and the illustrations well selected, a few of them being photomicrographs of excellent preparations. The work is a most acceptable contribution to the growing list of laboratory manuals.— M. s MINOR NOTICES. JAMEs R. Gow‘ has published a preliminary list of the flowering plants of Adair county, Iowa. WILDEMAN and DurRAND’? have issued a further publication on the Congo flora. It consists of the first fascicle of an enumeration of plants collected by Alfred Dewévre in 1895-6 in the state of Congo, and contains from Ranun- culaceae through Leguminosae, with descriptions of new species. A notice of the previous series may be found in Bor. GAz. 31:70. 1901.—J. M. C UND® has published a monograph of the genus Sorbus, which further rehabilitates a Linnean genus long included in Pirus. The author recognizes 58 species, and discusses them in great detail, with the help of text illustrations of venation, pollen grains, etc. He also includes subspecies and hybrids. Some ten or twelve North American species are involved.— }.M,C. THE FIRST part of the second volume of Wiesner’s Rohstoffe des Pflanz- enreiches has just been issued.2 It contains part of the seventeenth section on woods, newly elaborated by Dr. Karl Wilhelm with the assistance of Dr. S. Zeisel, who contributes the chapter on the chemistry of wood. The struc- ture of woods and their physical and chemical qualities are described in 51 pp., followed by a synopsis (go pp.) of the more important plants whose wood * Proc. Iowa Acad. Sci. 8: 1-8. 190 7Annales du Musée du Congo. Botanique. III. Reliqaiae.. Deweytoame. Fasc. 1. Bruxelles, May, Igo!. oo der Gattung Sorbus. Kongl. Svenska Vetensk.-Acad. Handl. 35 * 1-147. 9 WEISNER, JULIUS: Die Rohstoffe des Pflanzenreiches. Versuch einer tech- nischen Rohstofflehre des Pflanzenreiches. Second ed. Lieferung 6. : 1-160. Leipzig: Wilhelm Engelmann, 1901. 5. 374 BOTANICAL GAZETTE | NOVEMBER is used in the arts. A seventh chapter (incomplete) is to describe the micro- scopic characters of the most widely used woods. — B. THE LAST ISSUE of the Minnesota Botanical Studies (2:537-655. 1901) contains the following papers: E. M. FREEMAN, ‘A preliminary list of Minnesota Uredineae,” a little over 100 species being included ; DEALTON AUNDERS, “A new species of Alaria,” from the Californian coast; F. BuTTen, “A preliminary list of Minnesota Xylariaceae,” including 19 species; W. A. WHEELER, “A contribution to the knowledge of the flora of the Red river valley in Minnesota,” 325 species being listed, with eight excellent heliotype plates of plant formations; H. B. HUMPHREY, “ Observa- tions on Gigartina exasferata Harv.,” a histological study, with one helio- type plate; W. G. FANNING, “Observations on the algae of the St. Paul city water;’’ W. A. WHEELER, “Notes on some plants of Isle Royale;” D. LANGE, “Revegetation of Trestle island;” J. C. ARTHUR and E, W. D Hotway, “Violet rusts of North America,” with one plate; H. L. LYON, “ Observations on the embryogeny of Nelumbo ” with three plates, reviewed in the BOTANICAL GAZETTE for October.—J. M. C THAT FIRES are not always so detrimental as they seem is disclosed by a reading of the second edition of the phytogeography of Nebraska.” The first edition has been reviewed in this journal,’* and a statement as to the new material is all that is needed here. A comparison of the two editions shows that the entire book has been essentially revised and brought up to date, although the table of contents reads much the same in the two editions. Among the more important additions are a full discussion of methods for the determination of the frequence and abundance of species, a brief treatment of the primitive flora of the great plains, and the treatment of floral and vegetation elements and of accessory biological characters. Throughout the detailed chapters on the formations, much new material, the result of three years’ further labor, is added. In the review of the first edition, the impor- tance of this contribution to ecological workers was stated. Now that the book has been in actual use for four years, it is possible to speak yet more highly of its value. It is not too much to say that it is the most important valuable work that has yet appeared in the field of American phyto- eography.—H. C. CowLeEs. NOLES FOR-STUDENTS. S. YAMANoUCHTI™ has described and figured bodies in the dividing pollen mother cell of Lilium longiflorum, which stain deeply, are centers of radia- *PounD, Roscoz, and CLEMENTS, FREDERIC E.—The ee ot Nebraska, I. General Survey. 8vo. pp. 442, with four maps. Lincoln, Nebras The University Publishing Co. 1900. Second edition. ™ Bor. GAZ. 25 : 370. 1898. ™Einige Beobachtungen iiber die Centrosomen in den ee von Lilium longifolium. Beihefte Bot. Centralbl. ro: 310-303. pl. z. 1901 (gor] CURRENT LITERATURE 375 tion, and come to occupy a position at the poles of the nuclear spindle. He regards them as undoubtedly centrosomes.— J. M. C. Miss MARIA Dawson reports the results of three years’ experimentation to determine the economic importance of nitragin, the commercial culture of the nodule organisms of Leguminosae. Her conclusion is “that for peas grown upon ordinary garden soil, peat, clay, or loam inoculation with nitragin is useless and superfluous, whilst upon gravelly soils a small increase in crop results from its use.” *3—C R.B ELIZABETH DALE“ has studied the aerial tubers which are often very prominent in species of Dioscorea, notably D. safiva, in which they are said to become six or more inches in diameter. A study of their origin seems to indicate that they are stem structures, with conspicuous powers of propaga- tion. In fact, the author ventures the statement that ‘‘the abundant forma- tion of auxiliary tubers in m many species of Dioscorea seems as if it were connected with the fact that these plants do not appear to form seed readily,’’— C: DUusEN has studied the byrological collections of the Swedish polar expe- dition under Dr. A. G. Nathorst in 1899. The localities visited on east Greenland extend from Pendulum island (74° 40’ N.) to Cape Stewart (70° 30’ N.), about 150 miles north and south of Cape Parry. The island of Jan Mayen was also visited. In this work he has had the assistance of Dr. H. W. Arnell on Bryum and Plagiobryum, and of Herr C. Jensen on Dicranum and Sphagnum. Bryum subnitidulum, B. Dusenii, B. minus, B. Groenlandicum and B. Jan Mayense are described by Arnell as new species, and the sporo- phyte of B. obtustfolium is fully described for the first time.*—C. R. B. WAS OBSERVED by Sachs that when orthotropous roots, not too young, are inverted, they do not return to the vertical but become plagiotropous. émec finds a satisfactory explanation * of this in the alteration of the recep- tive apparatus described in his previous paper,” thus confirming his views as to the nature of the perceptivity of typical roots, as well as Noll’s idea, arrived at from theoretical considerations, that alterations in the response to directive Stimuli have their source in an alteration of the receptive structure. He has observed, coincident with this induced plagiotropism, a change in the “— 8raphic relations of the quality of the sensitive plasmic layer. The differ- ence between the original and the newly assumed structure is just the egy of Botany 15: 511-519. 1901. n the origin, development, and —_— nature of the aerial tubers in pha sativa. Ann. of Bot. 15: 491-501. p/. 26. 190 *5 Bihang till Svenska Vet. Akad. Handl. 27: 1-71. pls. g. 1901. ** Berichte deutsch. bot. Gesells. 19: 310-313. 1901. *7 Jahrb, £. wiss. Bot. 36:80. 1901. See also Bot. GAZ. 32: 145. 1901. 376 BOTANICAL GAZETTE [NOVEMBER difference between that found in orthotropous roots and normally plagi- otropous ones.—C. R. B : A SHORT TIME since reference was made in this journal to the question of nomenclature in phytogeography.® It is interesting to read in this con- nection the translation of Professor Flahault’s address at the Paris Congress of Igoo. hile many may object to the arbitrary usage of the various terms suggested by Flahault, all will agree that the general questions at issue are made clearer, and we shall await the more impatiently the final decision of a competent committee. It seems questionable to the reviewer, however, whether the nomenclature problem can be settled by a committee, or even a convention, An arbitrary set of terms often acts to prevent a real advance of knowledge, if strict adherence is given to them. Already the terms zone, region, and association have rather definite meanings in the minds of active phytogeographers. Even the much vexed word “forma- tion” is being more commonly used in the original and broader sense, as more inclusive than the word “association.” Perhaps other terms also will become more definite in a perfectly natural way.— H. C. COWLES. W. H. Lane in a late paper” has given a preliminary account of some experimental work on wild plants of a species of Anthoceros, in which he has demonstrated that by mutilating very young unopened sporophytes and bringing the cut surfaces in contact with soil under certain conditions of moisture, temperature, and light, bud-like outgrowths are produced from the vegetative cells that lie between the sporogenous region and the external layer. The cells in this region are the least specialized of all in the capsule. They become to some degree isolated in position by the decay of the sur- sounding tissue. The general sequence of the early divisions in the cells giving rise to the so-called prothallial growths are said to be ‘closely par- allel to the early stages of germination of the spores of Anthoceros.”’ His cultures were carried no further than to the production of a “bud-like stage,’ which produced rhizoids. The result would he more convincing if he had obtained flat prothallia bearing reproductive organs. This is the first discovery of apospory recorded for any liverwort, although found in a number of mosses and ferns.— FLORENCE M. Lyon THE IMPORTANT worRKs of Stahl and of MacDougal and Lloyd on myco- rhiza have been reviewed in this journal.2* Mention should also be made of the work of Hesselman” on the mycorhiza of arctic plants. Boreal forms * Bot. Gaz. 31: 361. 1900. 9 Bull. Torr. Bot. Club 28: 391-409. 1901. *°On apospory in Anthoceros laevis. Ann. of Bot. 15: 503-510. pl. 27. 1901. * BoT. Gaz. 30: 68. 1900. * Bihang till K. Svensk. Vet. Akad. Handl. 26:—. 1900. (See Bot. Centralb. 86: 239. 1901.) SET, 1 ERENT . Sr ey 1901] CURRENT LITERATURE 377 have been but little studied in this connection, and it might be supposed that the more xerophytic climatic conditions would work against the root fungi. Many plants, however, are found to possess mycorhiza in considerable abundance. Salix, various ericads, Dryas, Diapensia, and other species appeared much like more southern forms. W. Magnus has made an impor- tant cytological contribution?3 to the mycorhiza literature. He finds that the endotropic mycorhiza of Neottia is of two types, which differ in position, Structure, and function. Four to six layers of cells immediately within the exodermis contain fungal hyphae. In the outer and inner layers of these cells the hyphae are thin-walled, and the fungi are digested by the host plant; ultimately the fungal threads degenerate, forming masses which also contain parts of the cell protoplasm. These masses Magnus regards as excreted products. In contrast to the above the more centrally located hyphae are thick-walled and remain vigorous throughout, not being digested by the host; these hyphae have haustoria and are regarded as parasitic on the orchid. This study thus gives still another view as to the significance of endotropic mycorhiza. Magnus treats in detail the cytology of the hyper- trophied host cells, and at the close of the paper gives an excellent bibliog- raphy of the mycorhiza literature.— H. C. CowLEs. ITEMS OF TAXONOMIC interest are as follows: E. L. GREENE (Pittonia 4: 285-320. 1901) has described 17 new species of Viola, 12 of Cerastium, 5 of Rumex, 8 of Lesquerella, 3 of Draba, and 7 of Lacinaria—J. K. SMALL (Torreya 1: 73-75. Igo1.) has discussed Juncoides in the southeastern States, describing a new species; has described (édem 97) a new Crataegus from Florida; also (tdem 107-108) a new Chamaelirium; and (zdem 67) a new Alleghanian Rudbeckia.—E. P. BICKNELL (édem 102-105) has described 2 new eastern species of Lespedeza.—B. D. GILBERT (Fern Bull. 9: 53-54. 1901) has described a new Asplenium from Kamchatka.—F. S. EARLE (Muhlenbergia I:9-I7. IgoI) has written upon some fungi from Porto Rico, describing 11 new species and 1 new genus (Cercosporidium) of the Dema- tiaceae.—_ W. N. SuKSDORF (Bot. Monatss. 19: 91-93. Igo!) has described hew species of Sisyrinchium, Potamogeton, Juncus, Deyeuxia, Melica, and Equisetum, from Washington.—_THEO. Hoxtm (Ottawa Nat. 1§:ITo-IIl. 1901) has described 3 new Canadian species of Gentiana.—WILDEMAN and DuranD (Bull. Herb. Boiss. II. 1:839. 1901) have described a new ike (Bosqueiopsis) of Urticaceae from the Congo region._-SCHMIDLE and WE HEIM (idem 1007-1012, #/. 7?) have described a new genus (Riots) 0 of Pleurococcaceae.—H. MIEHE (Ber. deut. bot. Gesells. 19:434-441. pi. 2 ‘901) has described a new genus (Crafu/o) of marine Flagellates.—]. mie. SUMURA (Bot. Mag. Tokyo 15:67. 1901) has described a new Formosan slats (Alniphytium) of Styracaceae.—G. LopriorE (Malpighia 14: 435 and *3 Jahrb. fiir wiss. Bot. 35 : 205-272. 1900. 378 BOTANICAL GAZETTE [NOVEMBER 448. 1901) has described two new African genera (Argyrostachys and Sericostachys) of Amarantaceae.—H. and P. Sypow (Hedwigia Beibl. 40 62-65. Ig01) have described a new African genus (Hafalophragmium) of Uredineae.—FR. FEDDE (Engl. Bot. Jahrb. 31 : 30-133. Igo!) has taken up the Nuttallian genus Mahonia, recognizing 37 species, 7 of which are new.— SCHMIDLE (zdem 30: 246, 247, and 253) has described two new African genera (Myxoderma and Chondrogloea) of gerbe and a new African genus (Chaetonel/a) of Cladophoraceae.—J. K LL (Bull. Torr. Bot. Club 28 : 356-361. I901) has described 8 new ae aa trees from the southern states, and also a new genus (Arayodendron) to include Diospyros Texana ; and has also (¢dem 451-453) discussed Dasystoma flava and some related species, describing two as new.—R. M. Harper (dem 454-484) has described new species of Rhynchospora, Viola, Dicerandra, and Baldwinia, from Georgia—V. S. WHITE (dem 421-444. f/s. 37-go) has published a revision of the Tylostomaceae of North America, describing Io new species and including one new genus (Diéctyocephalos) described by L. M. Under- wood.—P, A. RYDBERG (idem 499-513), in continuing his studies of the Rocky mountain flora, has published new species of Trifolium, Vicia, Pri- mula, Cuscuta, Monarda, Castilleia, Pentstemon, Sambucus, and of g genera in the Compositae.—GEORGE MASSEE (Jour. Linn. Soc, 35: 90-118, Als. 4-5: 1901) has published the second part of his redescriptions of Berkeley’s types of fungi, including all the species of Discomycetes and Hysteriaceae of which type specimens exist at present in the Kew herbarium.—J. M. C es NEWS. Dr. ROLAND THAXTER has been promoted to a professorship of crypto- gamic botany at Harvard University.— Science. WE REGRET to announce the death of the young English phycologist William West, who died in India from cholera, at the age of twenty-six years. AN EXCELLENT biographical sketch and portrait of the late Dr. Charles Mohr appear in the September number of Plant World. It is written by Professor S. M. Tracy. Dr. A. F. W. Scuimper, professor of botany in the University of Basle, and widely known for his monumental Pflanzengeographie, died on Septem- ber 9, in his forty-sixth year. IN THE OCTOBER number of the Journal of Applied Microscopy there is an interesting illustrated account of the botanical laboratory and gardens of the Tokyo Imperial University, by Kiichi Miyake. A BIOGRAPHICAL SKETCH of Professor T. C. Porter, with an excellent artotype portrait, was published in the July number of the Budletin of the Torrey Botanical Club, having been prepared by Dr. N. L. Britton. Miss S. M. HALLOWELL, professor of botany at Wellesley College, has been given leave of absence for the year, and the work of the department will be under Miss Clara E. Cummings, assistant professor of botany.— Science. WE HAVE LEARNED that the herbarium of Theodor von Heldreich, pro- fessor of botany and director of the Botanic Gardens, Athens, is for sale. It contains approximately 20,000 species, and richly represents the floras of reece, Asia Minor, and Egypt. It contains also hundreds of types and authentic specimens of new species described by Heldreich in the works of Issier, PROFESSOR ALEx. P. ANDERSON, formerly in charge of plant physiology in the University of Minnesota, has been appointed Curator of the herbarium of Columbia University, to fill the position made vacant by the appointment of Dr. Howe to the Garden staff. Professor FRANK S. EARLE, formerly of the Alabama Polytechnic Institute, has been appointed assistant curator in charge of the collection of fungi at the New York Botanical Garden. AN ACCcouNT of the opening of the new botanical department at Glasgow University appears in the Annals of Botany for September last. It was a part of the celebrations on the ninth jubilee of the university, and the open- Tite 379 380 BOTANICAL GAZETTE | NOVEMBER ing address was made on June 13, by Sir Joseph Hooker, whose father had been for twenty years the professor of botany at Glasgow. He was followed by Lord Lister and Professor Bayley Balfour in proposing a vote of thanks for the address. AT THE DENVER meeting of the Botanical Society of America a com- mittee, consisting of Drs. Trelease, Britton, and Robinson, was appointed to investigate and report upon the condition of the National Herbarium. The fol- lowing resolution was passed: ‘‘ That it is the present policy of the society to accumulate invested funds until the annual income, interest and dues, is at least $500, and then to use such income yearly, or at greater or less intervals, as circumstances may dictate, for the best advancement of botanical knowl- edge.”’ THE SUBJECTS for the Walker prizes in Natural History for the next two years have been announced. For 1go2 the botanical subject is ‘‘ Nuclear fusions in plants,” besides a general subject entitled “ The reactions of organ- isms to solutions, considered from the standpoint of the chemical theory of dissociation,” which may just as well be competed for by a plant physiolo- gist. For 1903 the subject is “ A monograph of any genus or group of thal- lophytes.” For detailed information as to the conditions of competition address Glover M. Allen, Secretary, Boston Society of Natural History, Bos- ton, Mass. THE EXECUTIVE COMMITTEE of the 11th Congress of Naturalists and Physicians has asked that the following announcement be made: The Con- gress will convene at St. Petersburg on January 2, 1902, and continue until January 12. General sessions will be held January 2, 8, and 12; and sec- tional meetings on the other days, botany constituting a distinct section. Those wishing to be present as members should send their names and addresses, with the membership fee (3 roubles), also indicating their section, - to the Executive Committee of the Congress, University, St. Petersburg, not later than December 15. ait A Wholesome Tonic Horsford’s Acid Phosphate Taken when you are tired and completely worn out, can’t sleep and have no appetite, it imparts new life and vigor to both brain and body by supplying the needed tonic and nerve food. 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IT FLOATS. a nothin IN URIC ACID DIA- BUFFALO LiTHIA WATER {S245 fige 3 es THIS WATER DISSOLVES URIC ACID AND PHOSPHATIC SEDIMENTS. ETC., ETC, John = Shoemaker, M.D., LL.D., Prozssor of Matera Medica and Therapeutics in the Medico-Chirurgical C ollepe of Philadelphia, etc., in the New York Medical Journal, june, 22, 1899: “ is doubly efficient in Rheumatism and Gout, It The BUFFALO LITHIA WATER dissolves Uric Acid and Posphatic sediments, as well as other products difficult of elimination, while at the same time it exerts a m moderately stimu- tt effect upon the renal cells, and bande Fist Maran the swift rem oval of insolua = materials from avoided b Na elimination. niquestionably. troup the ey tear 8 Uric Acie and gee products o 3 f faulty tissue change is of conspicuous benefit, yet to nape oie formation a het still more importa as when it corrects This service is performed by the BUFFALO LYTHIA WATER lisestive failures which are responsible for the Saeco of deteterious materials.” The late Hunter McGuire, M. D., LL. D., skeen President and Professor af Clinical re — College a Medicine, kichmond, , and Ex-President of te American Medical Assoc “ : as an alkaline diuretic is invaluable. in Uric Acid IL yatahios © and indeed j in er a Uric Acid Diathesis, it is a remedy of extraordinary potency. erly goo te it in cases of Rheumatic . which had resisted ee ordinary yt with onineriel I have used it also in my own case, ga r from this ayo ve benefit. from it than from any other re ramiciy. Dr. P. B. Barringer, Professor of Physiology and Surgery, University of Rete ge a = “In more than twenty yeats of practice | have used Lithia as an anti- -uric acid agen? many times, and have tried it in a great variety a forms, both in the NATURAL WATERS and in Macnp scar As the “a re arte experience, | have-no hesitation in stating that for ark results [ have fou 9 WATER body. Ny experience with it as a solvent body. M signe ie it as a solven’ of old cxisting deposits ones has been Jp limited, Sod 1 T heeltate to are it Papi — er for o their disadvantage: but fo wr first class of conditions above set forth | feel that + BUFFALO LITHIA WATER ALONE. et Dr. Thomas re Syne gpa of Paris (Formerly of Baltmore), Suggestor 9 Lithia as a Sol- _— ric Acid, “Nothing I could say wou Sisk add 7 . 1 have frequently _ to the weli- beninets reputation of the fesults in URIC ACID DIATHESIS, RHEUMATISM, and GOL GOUT, and with this object have / : rope. Lithia is aluable as it exists in the carbonate, the | h a Seeey are's mode of solten and division fo ie = Sond‘ BURFALO LYTHIA WATER, "ics! we and Spondumne Mineral formations.” oe ¥ iw Mallet, Professor of C . University of virginia. Extract from report of analysis <— et Galcell aL ros Uaie AEID AND 1 that the coined of the w sito Al THE LABELS ! a THE GENUINE g A BAERS ‘PIAN 0s 2; COCOA Me ATE gs cc PUT UP IN PACKAGES LIKE THESE world for Pure, Rich and W Sympathetic Tone Com- bined with greatest Power and Durability. WEBER WAREROOMS: 108 Fifth Avenue, New ‘ 266 sieges Avene, Chica { Tremont Se Boston. ieee ; tea-kettle si sift up low: es .She will. JOHN M. COULTER anp CHARLES R. BARNES, ES WITH OTHER MEMBERS OF THE BOTANICAL STAFF < . _ OF THE UNIVERSITY OF CHICAGO a. apes 2 ee ee = APPLES MAKE 2 > CIDER = " A a ta) The soap made © PEARS, in Great Britain, is incom- parably the purest and best for the toilet and bath. All sorts of stores sell it, all sorts of people use it. AM rights secured. * Wotanical Gazette A Monthly Fournal Embracing all Departments of Botanical Science Subscription per year, $4.00. Foreign, $4.50. Single Numbers, 40 Cents The aibaaipoos price must be paid in advance. 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Thus DeBary, the exponent of the “bundle system,” states that “collateral bundles” are with rare exceptions characteristic of the stems and leaves of the phanerogams, but are likewise found in the Osmundaceae,* and that in their arrangement in the stems of the Osmundaceae they follow the ‘“dicotyledon type.”? Later we find Van Tieghem, the first enunciator of the ‘‘stelar theory” expressing himself as follows :3 La tige des Osmundes et des Todées différe de celle des autres Fougéres. La stéle axile et sans moelle du jeune Age, au lieu de se diviser en restant 8réle, demeure simple en s’élargissant progressivement 4 mesure que la tige 8rossit; elle prend une moelle de plus en plus large, a la périphérie de laquelle sont rangé en cercle un certain nombre de faisceaux libéroligneux 4 bois séparés, mais a libers confluents, entourés d’un péricycle commun et d'un endoderme général. Enun mot la tige de ces plantes demeure monos- télique a tout age, comme celle de la plupart des Phanérogames. * DeBary: Vergleichende Anatomie der Vegetationsorgane der Phaneroga- men und Farne 331. * DEBARY: of. cit. 246. 3 VAN TIEGHEM: Traité de Botanique 1373. 381 382 BOTANICAL GAZETTE [DECEMBER Plainly enough, therefore, these eminent botanists, starting from very different conceptions, have arrived at the same con- clusion, namely, that the central cylinder of the Osmundaceae resembles that of the phanerogams. _ Itis important to note, however, that heretofore all anatomical researches in this family have been confined to the tropical genus Todea and the cosmopolitan Osmunda regalis; and that hence the conclusion just stated has been based on the phenomena presented by these alone. When Van Tiegham proposed his ‘“stelar hypothesis” several cryptogams besides the Osmunda- ceae were cited as exceptionally possessing medullated mono- stelic central cylinders. Since then more extended researches have been made which have yielded important results. Thus it has been shown that the central cylinder of Ophioglossum and of Botrychium instead of being medullated monostelic is in reality “‘gamodesmic;”* that the central cylinder in the entire family Equisetaceae, some of whose species were included in the exceptions, is of the same kind;5 and that the central cylinder of the genus Helminthostachys is also of the ‘* gamo- desmic”’ type.© It is true that Strasburger holds’ that the internal endodermis and endodermal sheaths about individual bundles are of intrastelar origin, and not of cortical as is the external endodermis, and that therefore these exceptions still stand; but this objection may be advantageously left for sub- sequent consideration. Of the apparent exceptions, the family Osmundaceae has remained untouched, and I have undertaken the present research on this anomalous case, with the primary object of furnishing data that will help determine the proper morphological interpretation of its vascular system. The family Osmundaceae is a very limited one in point of numbers, consisting of but two genera, Osmunda with eight species, and Todea with six, and therefore constitutes a very *PoIRAULT: Ann. Sci. Nat. Bot. VII. 18: 113. 1893. SJEFFREY: Mem. Boston Soc. Nat. Hist. 5:155. 1899. *FARMER: Ann, Bot. 13: 421. 1899. 7 STRASBURGER: Histologische Beitrage. 3:—. 1891. Igor] ANATOMY OF THE OSMUNDACEAE 383 small part of the fern flora of the earth. But this does not seem to have always been the case,® for the Marattiaceae, although overwhelmingly predominant in the Coal period, con- stituted but 4 per cent. of the total filicineous flora in the Lower Jurassic, the remainder being composed of Osmundaceae and Cyatheaceae, with the related families Matonineae and Proto- polypodiaceae. As to distribution, the first genus is confined to the northern hemisphere, and the Todeas are with one exception found only in Australasia. Five Osmundas belong exclusively to restricted areas in east Asia and the adjoining islands; (O. Claytoniana occurs in the Himalayas and North America; 0. cinnamomea in eastern Asia, North and South America; and QO. regalis in every continent except Australasia. Of the Todeas, 7. barbara is a native of Australia, New Zealand, and South Africa; and the remaining species, the so-called ‘‘filmy” Todeas (Leptopteris of some authors), belong to oceanic islands in the eastern south-tropical region. Of these species I have had the opportunity of studying five, namely, O. regalis, O. cinnamomea, O. Claytoniana, T. barbara, and 7. superba. Nevertheless, in the following pages most atten- tion will be devoted to O. cinnamomea, not so much because its anatomy has not previously been described, as because the writer, for reasons which will become apparent, believes it retains a more primitive type of skeletal axis than any of the family so far investigated. The material of the species of Osmunda stud- ied was collected from several different localities, and in large quantities. Of O. cinnamomea specimens from fully a hundred and fifty plants were preserved and examined, and of each of the others perhaps one-third of that number. The more important Points were verified from specimens taken from three different localities, Observations have been mainly restricted to the mature root, Stem, and leaf trace. Some young plants of Osmunda were Studied, and the growing points of the older stems have been *Scorr : Studies in fossil botany 304. 1900. 9DIELs: Engler and Prantl’s Natiirlichen Pflanzenfamilien 14: 377. 1900. 384 BOTANICAL GAZETTE [DECEMBER sectioned. But the mature stem, especially the region at which it branches, has proved to be of chief interest from the stand- point of questions of comparative anatomy. THE STEM. GENERAL ANATOMY.—The mature stems are very stout rhi- zomes, exceptionally so in 7. barbara, which grow in a direction somewhat oblique to the horizontal. The leaves are in a closely set tuft at the anterior end, for they are annual and the inter- nodes are very short. The broadly winged, overlapping bases with their sclerenchymatous sheaths resist decay long after the remaining portion of the leaf has perished, and these, together with the roots, which are very numerous, greatly add to the bulk of the stem. The stem usually bifurcates once into two branches of equal size, which lie in a horizontal plane. A few specimens of O. regalis were found, however, in which one of the forks was much larger than the other, but the larger almost immediately divided again, so that there were three branches of about the same size lying in the same plane. The forking bears no relation to the number of leaves produced, counting from the cotyledons, nor to the age of the plant. Occasionally there is no branching at all, though maturity has long since been attained, while in rare cases it has taken place comparatively early in the life of the fern. The rhizome exhibits a very characteristic appearance in cross-section (fig. r). The outer portion, the thick external cortex (ex. c.), consists of very resistant, dark-brown scleren- chyma, in O. ctnnamomea of a rich red-tinted brown, in O. regalts and the Todeas of a black, and in O. Claytoniana of a dull brown hue. The cortex is marked by leaf-traces (/¢), which form a close spiral, and at the nodes by the escaping roots (7). In _O. cinnamomea sclerification of the cortical tissue is later in taking place than in the other species. The internal cortex (7. ¢.) is parenchymatous, comparatively narrow, roughly pentagonal, and its cells are heavily loaded with starch grains, Passing the pericycle and the bast region, which form a complete ee ae 1901] ANATOMY OF THE OSMUNDACEAE 385 sheath, the wood (x) of the stele is seen to be broken up into bundles of various shapes arranged ina circle, and sepa- rated from one another by the so-called medullary rays. These medullary rays extend out from a large pith. The pith or medulla in O. Claytoniana and T. superba is apparently homo- geneous. In O. vegalis it is often discolored and may contain one or more strands of brown sclerenchyma; in O. cinnamomea it is very frequently characterized by some brown sclerenchyma- tous tissue, and in 7. barbara there is a large axial strand of this supporting tissue. HistoLocy.— But we turn now to acquire a more intimate acquaintance with the stem as revealed by a study of its histo- logical features. For this purpose several sets of transverse and longitudinal series were prepared, and a great many microtome sections examined. The material cut included stems of various ages. As development proceeds rather slowly, all the tissues are mature only at a considerable distance from the apex of the plant. - The cortical part of the stem has little of interest for us other than in the respects already mentioned. The scleren- chyma consists of elongated, thick walled cells, with a small lumen containing starch grains. The walls are brownish, and marked by simple pits, which are round or slit-like. According to Strasburger," the endodermis is not the innermost cortical layer, but I am unable to verify this. He has made the state- Ment that the innermost cortical layer at a certain stage divides by tangential walls to form several layers of cells; of these, the outermost becomes differentiated as the endodermis, and the remaining layers lie between this and the phloem, filling the Place of a pericycle. The somewhat elongated cells of the endodermis are marked in every case by the characteristic cuti- cularization of the radial walls, which in transverse section shows as the “radial dot” (fig. 6, ¢,¢). The “radial dot” is dis- tinctively brought out by treatment with phloroglucin and hydrochloric acid, and also with dilute sulfuric acid. In O. *° STRASBURGER, of. cit. 449. 386 BOTANICAL GAZETTE | DECEMBER Claytoniana the radial markings are generally not as distinct as in the rest of the species studied, and the cells are reduced in size in comparison with those of the layers in contact (fig. 8, ¢). The contents in this species, too, are meager, consisting of granular protoplasm, a nucleus which as a rule stains a deeper red with saffranin than those of surrounding cells, and a few starch granules as shown by treatment with iodin solutions. Sometimes the endodermal cells of O. cinnamomea are likewise apparent by the lack of contents, in contrast to the heavily-laden cells, both ectad and centrad. Generally in this species, as in the remaining ones, 7. superba excepted, the cells are filled with tannin, so that the endodermis stands out very distinctly. The pericycle is entirely parenchymatous and consists of several layers—in O. Claytoniana and Todea of two or three, in O. cinnamomea of three or four, and in O. regalis of one to three. | The cells are elongated, cylindrical, provided with large nuclei, and filled with finely granular contents, part of which is starch. Haematoxylin imparts to this tissue a light blue color. Tangen- tial sections show that the orientation of the cells is very irregu- lar (figs. 3 and 9, p). Immediately opposite the point of origin of a leaf trace, and for a short distance below, the long axes of the cells run parallel with the long axis of the stem, but for the most part in the remaining regions of the stem there is consid- erable disturbance, though only in tangential planes. This dis- turbance is commonly so marked that the long axis of the cell is at right angles to the stem axis, and between this and the paral- lel position there is every gradation. Therefore in transverse section these cells are either round or more or less tangentially lengthened (jig. 8, pf). This variation in orientation is of interest, as it is connected with a similar phenomenon in layers lying nearer the cauline axis, namely in the phloem region. XYLEM.— Before dealing with the phloem, however, it will be convenient to describe the xylem. The wood elements are of two kinds, namely, small ringed and spiral elements consti- tuting the protoxylem, and scalariform tracheids which are of later development constituting the metaxylem. Occasionally a See 1901] ANATOMY OF THE OSMUNDACEAE 387 parenchymatous cell is found among the tracheids. A transverse section shows, as mentioned before, a ring of variously shaped bundles; and by tracing these up and down, or by boiling a piece of stem in potash and then removing the softer tissues, there is shown to be a network forming the wall of a hollow cylinder, the strands being the “bundles” of DeBary, and the meshes the Spaces occupied by the “ medullary rays.” Though there is a great deal of regularity in the apparent construction of this net- work, as proved by DeBary and Zenetti in O. regalis, yet a study of development shows that the “bundle theory” is inadequate for giving the right conception of the vascular system. In the young stem of the Osmundaceae the wood forms a completely closed cylinder, and Van Tieghem, basing his conclusions on Todea and QO. vegalts, has stated this to be the case for the Whole family. I am able to state that the phenomena in the young stem of O. cinnamomea and O. Claytoniana are in accord- ance with his general conclusions in this respect. Now directly above the point at which a leaf trace leaves the stele the wood is not developed for some distance. This gap is filled by parenchyma chiefly, except at the outer part, which is occupied by sieve tubes. There are exceptions in O. cinnamomea to be described later. Thus a transverse section of the stele, just above a node, shows a ring of wood broken at one place, the break being occupied by the tissues just referred to; in other words, the stele here has one medullary ray. fig. 23 shows a transverse section of the stem of QO. Claytoniana through this region. Still further up the internode the ring is complete again. There is the same sort of gap above the second node. However, as the nodes become more frequent, that is, as the internodes become shorter, a leaf gap extends through more than One internode, and in a transverse section there is more than one medullary ray, until in the full grown stem, where a leaf gap extends through several internodes, a transverse section shows Several gaps cut across, or in other words shows several medul- lary rays. It is therefore evident that the number of medullary rays seen in any transverse section depends on the frequency of 388 BOTANICAL GAZETTE | DECEMBER the nodes and the length of the gaps. In well nourished stems the number is greatest in O. Claytoniana ( fig. 17), there usually being about twenty, and in 7. darbara (fig. 24) the fewest. In this species the gaps are quite short, so that while the wall may be thin in many places at any given level, there are not more than two to six medullary rays seen in the cross section (fig. 24). The persistent portions of the cylinder of wood, the ‘bundles,’ present various contours in cross section, the shape of any particular portion lying between two adjacent gaps, that is, of any strand, varying with the level at which it is cut. Just below where the leaf trace is given off, the wall is hollowed out on the side towards the pith, so that the transverse section of the strand presents a horseshoe shape (fig. 177). The middle of the inner surface of the strand at this level is occupied by protoxylem, which consists of about a half dozen small ringed and spiral vessels. Following the strand down, it is seen that the arms of the horseshoe thicken on the sides facing one another, especially towards the ends of the arms ( fig. 75,5). Finally, the opening between the ends is fully closed and a small group of parenchymatous cells lying exactly centrad of the protoxylem is thereby enclosed (fig. 17,5). The parenchyma is more and more encroached upon by the xylem, until lower down it is seen no more. Not far below where the parenchyma vanishes, the protoxylem in that strand likewise disappears. Somewhat above the level at which the parenchyma is enclosed the strand begins to thin out on the outer side, a sharp trough- like indentation appearing, but not in the same radius as that in which the protoxylem lies. This trough continues to deepen until a few nodes down the strand is cut through, the point at which the break occurs being, indeed, the apex of a leaf gap. Thus neither the outer nor the inner surface of the cylinder of xylem is smooth; the lower part of a leaf gap can be traced as a hollow on the inner surface just below where the leaf is given off, ending as a blind tube amongst the tracheids, while the upper end of the gap may be traced as a furrow on the outer sur- face of the cylinder, gradually becoming more and more shallow ee dl Se 1901] ANATOMY OF THE OSMUNDACEAE 389 The protoxylem occurs in small groups of six to eight cells each, and a transverse section of the central cylinder shows from five to seven of these groups. Each group of protoxylem elements passes out in its entirety into a leaf trace, and on fol- lowing back from the leaf trace each vanishes as already described. The protoxylem is therefore not continuous through- out the stem, but is in small, discontinuous strands. This fact has been recorded for O. regalis by Zenetti.™ Lying externally to the wood are from four to six layers of elongated parenchymatous cells, rich in protoplasmic contents and in small starch grains. They are continuous with the parenchymatous cells of the medullary rays and do not mate- tially differ from them. Those occupying the middle of the medullary rays have more meager contents, and towards the stem axis they became larger. That there is a “ xylem sheath” characterized by cells of greater size and richer contents such as Zenetti describes for O. vegalis, | cannot affirm, and certainly there is not such a sheath in QO. Claytoniana. PHLOEM.—The tissues that have just been described are bordered by the phloem, which consists chiefly of sieve tubes. Parenchymatous cells are sometimes met with in isolated posi- tions in the metaphloem, and between the metaphloem and the Protophloem they constitute a more or less broken layer, most Pronounced in O. Claytoniana, and least constant in O. regalis and I. barbara.’ The sieve tubes are strongly developed and are of the “type vigne” of Lecomte. They are large, have thin walls of unmodified cellulose lined with a delicate layer of protoplasm, and are devoid of nuclei. They are provided with oblique terminal walls and are furnished with sieve plates both simple and compound. The sieve plates are covered with ‘‘olobules brillants,” and by treatment with proper reagents callus plugs may be demonstrated in them. The sieve tubes of the proto- Phloem are smaller than those of the metaphloem and their terminal walls are not as oblique. As there has been considerable difference of opinion regarding ™ ZENETTI: Das Leitungssystem im Stamm von O. rega/is. Bot. Zeit. 53:63. < 399 BOTANICAL GAZETTE | DECEMBER the disposition of the phloem in OQ. regalis, it will be well to define a sieve tube. Both Russow and Janczewski have studied sieve tubes very carefully, and Poirault has more recently rein- vestigated the subject in the vascular cryptograms. The investi- gator last named summed up his observations on sieve tubes in the roots of vascular cryptogams in the following terms :” Les tubes criblés peuvent se rapporter 4 deux types: le premier carac- térisé par des cloisons transverses perpendiculaires aux faces principales et ne portant qu’un seul crible (¢vfe Courge, Lecomte); le second reconnaissable a ses cloisons transverses trés obliques portant d’autant plus de cribles que leur obliquité est plus grande (¢yfe Vigne, Lecomte). On trouve, en outre, sur les faces longitudinales des ponctuations isolées ou réunies en trés petits groupes, constituant rarement des cribles aussi développés que ceux des faces transverses. Le contenu de ces tubes est un liquide hyalin tenant en suspension de nombreuses sphérules réfringentes, rassemblés surtout au niveau des cribles et des ponctuations isolées. I] n’ 'y a pas de noyau. La membrane est cellulosique. He further adds that two substances occur as a rule — (1) the “globules brillants’’ already mentioned, and (2) “les bouchons calleux qui font corps avec la membrane et peut-étre la traversent entiérement.” In dealing with the sieve tubes of the stem and petiole he does not point out any other peculiarities, but deals at length with the callus plugs, and the perforation of the sieve plates.% The observations of Russow, Janczewski, and Poirault agree for the most part, except in reference to the callus plugs. The following criteria would seem to be distinctive in deter- mining the presence of sieve tubes in the Osmundaceae; the existence of sieve plates, the absence of nuclei, and the presence of ‘‘globules brillants.” Less distinctive and rather as a con- firmatory test I have sought for callus. Russow made this test one of paramount importance, but it seems best in dealing with the vascular cryptogams to give it a second place, and for the following reasons: (1) the callus, so-called, in the seive tubes ™?POIRAULT: Or sur les cryptogames vasculaires. Ann. Sci. Nat. Bot. VIL. 18: 138. 1 3 POIRAULT : of. cit. 191. ae ee, Sik Igor | ANATOMY OF THE OSMUNDACEAE 391 of the vascular cryptogams may not be identical with that found in phanerogams ; (2) it occurs in minute quantities only, and in some plants (e. g., the Ophioglossaceae) probably does not occur at all; (3) its presence is determined by delicate microchemical means, and then only by limited color reactions. Janczewski*'+ claimed to have found callus in Preris aquilina alone of all the vascular cryptogams he examined, and states that it does not occur in O. regalis. The reagents he used were Schulze’s solution (or chlor-zinc-iodin) and rosolic acid. On the other hand, Russow found callus in all of the sieve tubes he examined. The reagent he used was a mixture in variable pro- portions of chlor-zinc-iodin and potassium iodid-iodin. It should be stated that in the vascular cryptogams callus occurs in _ the wall of the sieve plate, appearing as if it were a part of the wall. After staining with a suitable iodin solution, the callus shows in- face view as one or more round brown spots, and in section as rods or granules occupying the entire thickness of the lamella. Poirault has largely corroborated Russow’s observa- tions. He disagreed with Russow’s generalization that it is a constant feature of sieve tubes, for he states that he has been unable to find any trace of callus in Angiopteris and Ophioglos- sum.'S In view of these investigations, therefore, it becomes a matter of interest to know if the sieve tubes of Osmundaceae Show the phenomena of callus as described by Janczewski for Pteris, and by Russow and Poirault for many others of the vas- cular cryptogams. Accordingly tangential and transverse sections about five microns thick, of the three Osmundas studied and of 7. barbara were cut from mature pieces of stem embedded in celloidin. In the present research the writer has tried several stains, such as Tuthenium oxid, Hofmann’s blue, rosolic acid, and Russow’s mixture. These have been applied to sieve tubes of plants from widely separated groups, such as Vitis (summer and winter sieve tubes), Tilia, Pinus, Pteris, and the mixture of chlor-zinc-iodin “ JANCZEWSKI: Tubes cribreux. Ann. Sci. Nat. Bot. VI. 14:50. 1882. *S POIRAULT : of. cit. 192. 392 BOTANICAL GAZETTE [DECEMBER and potassium iodid-iodin proved to be by far the most satis- factory reagent for the demonstration of callus. The two con- stituents of this reagent were prepared fresh, and then mixed in different proportions until one giving the best results was obtained. The proportions vary with the different kinds of plants tested. In using this stain, though the presence of the celloidin is not a serious objection, it is preferable to dissolve out the celloidin, wash in alcohol, then in distilled water, and examine in stain on the slide. In face view it is difficult to make observations on account of the “globules brillants,” hence the most reliable observa- tions can be made on sectioned plates. Almost at once after applying the stain, the callus plugs become evident, staining a dark red-brown (fig. 7). They appear as more or less fine rods, completely traversing the sieve plate, and their number ina sieve plate depends on its size. The cellulose is slower in staining; at first it is light blue or a violet, and later a deep blue. Hence the callus plugs are to be seen most clearly in the early stages of the staining process. The stain unfortunately is not perma- nent. Callus was clearly demonstrated in the species under investigation, but on account of the size of the cells and of the sieve plates, 7. darbara proved the best subject for the purpose. As one of the characters of the sieve tubes of the Osmundaceae, we record, therefore, the regular occurrence of callus plugs in the sieve plates. The “globules brillants” are exceedingly abundant in the sieve tubes, and especially in the older ones (fig. 7). While they adhere to the protoplasm of the cell and may be found in any part of the cell they are by far most abundant about the sieve plates, dotting their surface, filling the pits, and surround- ing the entrance to the pits. They are evidently not homoge- neous, but appear to consist of two substances, one of which is more refractive than the other, for by slight focusing up and down they change from a dark looking, opaque granule to a light semi-translucent spherule. Jodin solutions stain them brown, not appreciably different from the callus plugs. Occasionally Igor] ANATOMY OF THE OSMUNDACEAE oy OS irregular fragments of matter are found in the cell, which also stain a similar brown. The relation between these fragments, the ‘‘globules brillants,” the callus, and the disappearance of the nuclei calls for further investigation. The sieve plates are very numerous, and vary in size and form. The walls of the pits are abrupt, and the number of pits varies with the size of the plate. The larger plates are irregu- larly oblong and the smaller ones are round. In Todea they are largest and most numerous. Having described the sieve tubes at some length, we shall now examine their distribution. The phloem forms a continuous sheath, the outer portion of which is the protophloem. To this peripheral part of the phloem I find that DeBary and Zenetti alone make specific reference. The former states* that this region of the central cylinder is characterized by ‘“ quergestreckte Zellen,” but he offers no opin- ion as to the nature of the tissue in question. Zenetti divides the zone into strands of typical protophloem and connecting portions of ‘‘quergestreckte Zellen.”” The typical protophloem is in short strands one cell thick, lying ectad of strands of xylem which are about to give off the leaf traces. The ‘‘quergestreckte Zellen” he cannot recognize as sieve tissue, because of the ‘ quer- gestreckt”’ form and the position of the cells. This tissue in Osmunda regalis, consequently, forms a cylinder interrupted by the strands of protophloem only, but Zenetti found it to be gen- erally two cells in thickness and opposite the medullary rays often several cells deep. I have ascertained that the ‘‘quergestreckte Zellen” are devoid of nuclei; their walls are of cellulose, staining violet or blue with iodin solutions. They are rounded, elongated ele- ments, with more or less oblique terminal walls; and are char- acterized by the possession of sieve plates which show the callus reaction when treated with Russow’s reagent; the cells contain an abundance of ‘globules brillants’’ which are aggregated especially about the plates; the protoplasm is reduced to a thin parietal layer. The characters of the typical protophloem cells *° DEBARY :. of. cit, 360. 394 BOTANICAL GAZETTE | DECEMBER ”” are the same as those of the ‘‘quergestreckte Zellen regard to orientation (fig. 6). Transverse sections show that the long axes of the so-called ‘‘quergestreckte Zellen” are tangentially placed, and never radi- ally to any degree. To determine the slant of the long axes, therefore, with reference to the axis of the stem, tangential sec- tions must be made. If such be examined it is seen that some are exactly at right angles to the axis, others are almost or entirely parallel, and between these extremes there is every gradation. This at once explains the difference in “width” of the ‘‘quer- gestreckte Zellen’”’ in transverse section. It is further to be noted from the tangential sections that the ends of typical pro- tophloem cells never abut against the long sides of the ‘ quer- gestreckte Zellen,” but there is a gradual change in the direction of the latter so that their ends communicate with the proto- phloem and it is quite impossible to say where the typical proto- phloem ends and the “quergestreckte Zellen” begin. The root and leaf have been examined for ‘‘ quergestreckte Zellen,” for if these elements constitute a characteristic textural feature of the Osmundaceae they would naturally occur elsewhere than in the stem. They are not present at all in the appendicular organs. Further, in the young sporophyte, where the leaf gaps are far apart, they are absent from considerable portions of the stem. The real nature of the ‘“quergestreckte Zellen” will be discussed after observations on their development and their rela- tion to the leaf traces have been described. The ‘‘quergestreckte Zellen” and the typical protophloem cells form a continuous sheath in all the species studied. In O. Claytoniana the elements of this sheath are very much smaller, and so it is easier to distinguish them from the metaphloem. In 7. barbara their histological characters are best studied because of their relatively large size. Frequently in O. cinna- momea and O. regalis it is difficult to decide in the mature stem whether or not certain cells belong to this sheath or to the metaphloem. But evidently in all of the species the sheath is rarely more than two cells in thickness, and often, especially in except in 1901 | ANATOMY OF THE OSMUNDACEAE 395 O. Claytoniana, there is but a single layer. Opposite outgoing leaf traces the sheath is reduced to a single stratum. The metaphloem forms a hollow cylinder consisting of large sieve tubes such as have already been described. They are thin-walled, and especially in O. regalis in the older parts of the stem have often collapsed. The sheath is one or two cells thick Opposite the strands of xylem, and several cells in thickness opposite the medullary rays (fig. 8, ph). Most of the tubes run parallel with the long axis of the stem, but here and there “quergestreckte’’ examples occur. This cylinder of metaphloem has a smooth outer surface, but the inner surface is rendered very uneven on account of the wedge-like proliferations of the sieve tissue opposite the leaf 8aps. Since this is a phenomenon common to all the species Studied, we naturally seek an explanation of this peculiar dis- position of the phloem. In his memoir on sieve tubes Janczew- ski,” who could hardly have been prejudiced by any stelar theories, noted that isolated sieve tubes occur occasionally here and there in the medullary rays of O. rvegalis. The writer has found undoubted cases of the same thing in O. cinnamomea. Two such eminent botanists as DeBary and Strasburger have disagreed as to the topographical distribution of the layer of metaphloem sieve tubes in QO. regal’s. The former states that the sheath is continuous, while the latter states that he puts himself in Opposition to DeBary on this point, for he considers the phloem to be interrupted opposite the medullary rays. Strasburger does not say for what reason he considers the cells Opposite the medullary rays not to be sieve tubes, My own observations on OQ. regalis are precisely in accord with those of DeBary and Janczewski. The cells opposite the medullary rays differ in no way from the sieve tubes opposite the xylem strands. I have found the same to be true of the other species studied, With the additional observation that isolated sieve tubes occur Sometimes in the tissues filling the leaf gaps of O. cimnamomea. * JANCZEWSKI: of. cit. 66. * DEBARY: of. cit. 360. 19 STRASBURGER : Of. Cit. 449. 396 BOTANICAL GAZETTE [DECEMBER To this last observation I have two others to add, namely, the occurrence of an internal phloem in which the sieve tubes form a more or less continuous ring (figs. 2z, and 22), and in rare cases the union of external and internal phloem through a _ leaf gap. Inacertain rich, moist situation about a dozen well nourished plants of O. cinnamomea grew, of which, on examina- tion, five showed the phenomenon of a continuous layer of inter- nal phloem. Search in an adjoining locality resulted in finding specimens which showed the same feature. To extend the range of observations, I visited a peat bog some twenty miles distant from Toronto, where I knew the cinnamon fern grew, and secured specimens characterized by the same peculiarity. ig. 27 shows a transverse section of a stem found in this last locality. The sieve tubes of this internal phloem are as typical as those of the external, and except for their position not be distinguished from them. They do not always form a continuous ring as do the sieve tubes of the external phloem, but are often in more or less detached groups, embedded in small celled parenchyma. The layer of sieve tubes is from one to three cells thick. It should be added that internal phloem occurs only near where the forking of the stem takes place. O. cinnamomea shows likewise two other features which are constant throughout every part of the stem, and at once distin- guish it from other species: (1) an internal endodermis, and (2) several layers of parenchyma between this and the xylem. INTERNAL ENDODERMIS.—The internal endodermis possesses the characteristic radial dot, though sometimes not as clearly distinguishable as in the external endodermis (17. ¢., fig. 2} Its cells are usually larger than those of the latter, but are filled with similar contents, most frequently tannin (fig. 75). It is further to be noted that it bends outwards opposite the leaf gaps (figs ro, etc.), and not infrequently connects through them with the external endodermis. I have examined scores of stems of the cinnamon fern, and in every specimen there was an internal endodermis. On the contrary, it seems to be invariably absent from the other species studied. As the central cylinder of the tgor] ANATOMY OF THE OSMUNDACEAE 397 family Osmundaceae has heretofore been classed as monostelic, the existence of an internal endodermis in one of the species is therefore a matter of considerable moment, especially if it be regarded as a real phloeoterma. Between the internal endodermis and the xylem there is a cylinder of elongated parenchyma, rich in starch and protoplasm, and from two to seven cells in thickness. This layer is continu- ous with the medullary rays. In 0O. regalis, O. Claytoniana, T. barbara, and T. superba a similar but thinner layer it found as a rule, and the cells are always smaller and richer in contents than those of the medulla on which they border. THE MEDULLA.—The medulla is very large in this family, particularly so in O. Claytoniana, and consists of large-celled parenchyma. Most of the cells are partly filled with large Starch granules, but frequently some of them contain tannin, especially in 7. darbara. A brownish fluid may occur in inter- cellular Spaces, and in O. regalts within the cells themselves. In these regards there is often a striking resemblance between the parenchyma of the medulla and that of the internal cortex in the Same plant. But there yet remains to be described a still more significant phenomenon, namely, the occurrence in the pith of brown sclerenchyma of the same kind as is found in the external cortex (figs. rg and 20). This is probably a primitive feature, and in this, as in many other respects, O. cinnamomea proves to be most interesting. Out of forty-four pieces of stem, chosen at random, and representing a corresponding number of different plants of this species, twenty-five of the examples showed brown sclerenchyma in one or both ends. It occurs as a central strand, varying in size from a few cells to almost the limits of the pith, Or as several small strands irregularly arranged. Fig. 14 is a Photograph of the transverse section of a stem in which there is a large axile strand, and fig. 75 of one in which the scleren- chyma is entirely absent from the pith. Further it has the peculiar habit of being present at one level, but perhaps bot at another; so it is likely to be found in nearly every plant if the Stem be sectioned from end to end. 398 BOTANICAL GAZETTE [DECEMBER This same habit is characteristic of its appearance in O. regalis (fig. 20), but more often it is not present at all. That brown sclerenchyma occurred in the pith of O. rvegalis did not escape the observant DeBary,” but elsewhere I find no reference to this fact. Strangely enough, however, out of thirty-five or forty plants harvested from one locality there was not a trace of sclerenchyma to be found in the medulla of any of them, while in one region not far distant 25 per cent. showed this phenome- non, and in another a still higher per cent. Parenchyma is the sole constituent of the medulla of O. Clay- tontana (fig. 17). This is probably true of 7. superba too. fig. 25 is a cross section of 7. barbara taken too near the growing point to show sclerenchyma, but farther down the medulla was occupied by a large strand of this tissue (fig. 2). Thus medullary brown sclerenchyma is usually present in O. cinnamomea, in O. regalis not uncommonly, and in O. Claytoniana not at all. In 7. darbara it also occurs, but apparently not in T. superba. It is perhaps significant that such series can be arranged, but of greater importance is the fact that the occur- rence in the Osmundaceae of brown sclerenchymatous tissue, apparently within the cauline central cylinder, has no parallel among existing ferns. THE FORK,—There yet remains to be described the anatomy of one particular portion of the stem, the part in the region of bifurcation. It has been stated that it is peculiar to the stem of the Osmundaceae to fork once, and that in a horizontal plane. We shall treat first of the phenomenon in O. cinnamomea. Trac- ing the main stem forwards, it is seen to become flattened and then to become constricted in a median vertical plane. Imme- diately anterior to the point of bifurcation of the vascular axis, there is a wide ramular gap in the céntral cylinder of each branch (fig. ro). Sections of the main axis immediately below the fork show two bands of phloem, one on the upper and one on the lower internal surface of the central cylinder (fg. 13). Sections passing through just in front of the region of bifurcation 2° DEBARY: of. cit. Pp. 290. ——— 1901] ANATOMY OF THE OSMUNDACEAE 399 show similar bands of phloem along the inner wall of the central cylinder of each branch (fg. zr). Cases have been described above, in which there is a complete cylinder of inter- nal phloem instead of the two isolated bands just referred to (jig. 2z). The internal and external phloem connect through the ramular gaps (fg. rr). Likewise the internal and external endodermis are in textural continuity through these gaps, so that there is free communication between the cortex and the pith (fig. 20). Sometimes the cortex lying between the two branches con- tains brown sclerenchyma which is continuous through the ramu- lar gaps with strands of the same tissue occurring in the medulla of the branches. Frequently in less vigorous plants a transverse section of the main axis posterior to the point of ramification shows a diamond-shaped piece of cortex surrounded by endo- dermis (fig. 72). Posteriorly this included piece of cortex becomes continuous with the medulla of the main axis (jig. 73), and anteriorly with the general cortex (fig. 17). Twenty-five forks of O. cinnamomea were selected at random and sectioned. Twelve of them presented the phenomenon of typical wide ramular gaps. Six of them were of the reduced kind just described. In five cases there were gaps in the xylem only, cortex and medulla never becoming continuous; and in two even the xylem did not open up (fg. 76). For reasons to be outlined later, the writer believes the wide gaps to be the most primitive. O. regalis presents a much degenerated form of ramular gap, for here only the xylem opens (fig. 79). In O. Claytoniana the degeneration is carried still farther, for as a rule there are no branch-gaps at all (fig. 28). In 7. barbara the xylem alone may Open up. The phenomena of the fork may be thus summarized : (1) Complete ramular gaps occur only in O. cénnamomea. (2) Internal phloem occurs only in O. cimnamomea. it is found in the branches just above, and in the parent axis just below the point of bifurcation of the central cylinder. 400 BOTANICAL GAZETTE [ DECEMBER (3) The internal phloem may form an entire cylinder. (4) Where gaps are complete, the cortical and medullary tissues connect through them. (5) Thus sclerenchyma of the cortex is sometimes continu- ous with sclerenchyma in the medulla of the main axis, and of the branches. (6) O. cinnamomea presents the following forms of ramular gaps arranged in order of degeneration, (2) complete gaps, (4), phloem and xylem only open, (c) the xylem alone opens, (@) no gaps at all. (7) O. regalis and T. barbara show gaps in the xylem only, and in O. Claytoniana there are usually none at all. 0. Claytont- ana, therefore, presents the extreme case of degeneration. THE. LEAF TRACE. The leaf traces pass very obliquely up through the external cortex. A section of a leaf trace shortly before it passes into the petiole presents some noteworthy characters. In the first place there is no pith, but a solid horseshoe-shaped mass of xylem with the convex side turned outwards (jig. 5, *). The xylem is made up of large scalariform tracheids with a protrud- ing mass of a few small vessels constituting the protoxylem. The protoxylem is situated on the inner face of the single strand of xylem (fx), and is continuous with that of the stem. In Z. barbara it frequently breaks into two or three groups. Surrounding the wood is a layer of parenchyma, which on the concave side of the xylem quite fills the space between the arms of the horseshoe. The phloem consists of a crescentic band of sieve tubes, one to three cells thick on the external side of the leaf trace (p2), and a smaller band on the opposite side (gk). The protophloem consists of small elements which form a ring, broken only on the concave side of the xylem. Here the ring is completed, however, by the inner band of metaphloem. In Q. cinnamomea and T. barbara isolated proto- phloem cells have been observed by the writer on the side of the inner band of metaphloem towards the stem axis. On the convex 1901] ANATOMY OF THE OSMUNDACEAE 401 side the protophloem is separated from the metaphloem by paren- chyma. There are no “quergestreckte Zellen.” The pericycle consists of two or three layers of cells, and is bounded by a well developed endodermis continuous with that of the stem. With reference to the attachment of the leaf trace to the cauline vascular axis Zenetti has given a very careful and accurate description.” Strasburger has held”? that the stele of the petiole of 0. regalis is a collateral bundle. He has considered the inner band of metaphloem to be a parenchymatous tissue. However, the cells of this band prove to be characteristic sieve tubes, and are continuous with sieve tubes in the stem opposite the medullary rays. The leaf traces, therefore, are undoubtedly concentric. Several botanists have arrived at the same conclusion for Q. regalts 73 24 In summary, the most important features of the leaf trace are: (1) the absence of a pith, (2) the endarch xylem strand. (3) the concentric type of stele, (4) the absence of ‘ querge- streckte Zellen,” and (5) the cylinder of protophloem completed on the inner face by a band of metaphloem. THE ROOT. The roots have a definite relation to the leaves, both in posi- tion and in numbers. Two roots invariably originate from the base of every leaf trace, or from the central cylinder immediately below. They come off at the same level, one opposite each arm of the horseshoe-shaped strand of xylem ( fig. 18) in every case where there are just two roots to a leaf. They grow almost directly outwards, and so in a transverse section of the stem are cut longitudinally. In such a section it is seen, likewise, that the cortical tissues of the stem and root are entirely independent of each other, and that, therefore, the root is of endogenous Origin. This fact is true of the secondary roots also. * ZENETTI: of. cit. 69. * STRASBURGER : of, cit. 448. 3Scorr; op. cit, 319. * ZENETTI : of. cit. 66. 402 BOTANICAL GAZETTE [DECEMBER The cortex is exceedingly thick, forming by far the main bulk of the root, and consists of large celled sclerenchymatous tissue. The cortical cells diminish in size towards the periphery, and become thicker walled. In TZ. darbara, however, there is a discontinuous ring of exceedingly thick walled brown scleren- chymatous cells immediately surrounding the vascular axis. The endodermis, which is continuous with that of the stem and leaf, is very pronounced in all of the species, and is at once noted by the radial dot, and by the fact that its cells are filled with tannin. In the second particular, exception must be gener- ally made of O. Claytoniana. The stele is comparatively small, and is typically protostelic, since there is no pith. The wood has a narrow elliptical form, consisting mainly of very large scalariform tracheids. At each end of the ellipse there are a few small protoxylem elements, which are especially evident in the young root, and which have no connection with the protoxylem of the stem or leaf. The root, therefore, is diarch. There are likewise two bundles of phloem alternating radially with the bundles of xylem. In all of the Osmundas, however, | have observed triarch steles in the larger roots, which exception is of comparative frequence in 0. cinnamomea. The phloem consists of two flat bundles or bands. These bands are made up chiefly of thin walled sieve tubes which are of the same kind as‘occur in the stem. None of them are ‘‘quergestreckt.” The phloem is separated from the xylem by three or four rows of parenchyma, and from the endodermis by a two rowed parenchymatous pericycle. DEVELOPMENT OF THE TISSUES FROM THE GROWING POINT. In discussing this subject there are two points in particular which will receive special consideration: (1) the statements of Strasburger and Zenetti regarding the origin of the endodermis, and (2) the real nature of the “‘quergestreckte Zellen.” The determination of the relation of the tissues to the apical cells seems of little concern, and moreover in the study of the apical region of the growing point there are serious difficulties. 4 1go1 | ANATOMY OF THE OSMUNDACEAE 403 Having described these for 0. regalis, Professor Bower aptly remarks :?5 The meristem being thus at times irregular, and the subdivisions of the segments being variable, it is to be expected that the study of it (the apical region of the growing point) in longitudinal section would present difficulties, and I have not been able to trace any definite and characteristic mode of segmentation. Longitudinal sections cut from a considerable number of stems show that a conical apical cell is present. The relations of the sur- rounding tissues, and their reference to regularly succeeding segments are difficult to recognize. To these observations on the extreme apical end of the growing point we have nothing to add, but pass further down the stem. A short distance from the apex of .the stem, the various tis- sues, though in embryonic form, become apparent. The cylinder of wood, whose thin walled, unlignified cells are still provided with protoplasm and nucleus, can be distinguished from the pith, the parenchyma in the leaf gaps, and the immature phloem. The pericycle is rich in protoplasm, and its cells are radially arranged. At an earlier stage still, even before there is any evi- dent differentiation in the vascular tissues, the leaf traces can be Seen coming off from the cauline vascular axis. When the protoxylem can be first demonstrated by phloro- glucin and hydrochloric acid, the endodermis (both internal and €xternal in the case of O. cinnamomea) is also demonstrable by the same reagents, though not before. Zenetti has claimed” that at the time the protoxylem is formed, the endodermis, peri- cycle, “‘quergestreckte Zellen,”’ protophloem cells, and some Cortical cells are all in the same radial rows; and that, there- fore, all have originated from the same mother layer. Stras- burger has asserted?’ that the tissue lying in the stem between the phloem and the endodermis and occupying the position of a pericycle arises by tangential divisions with the endodermis out of the innermost cortical layer. Therefore, not the entire Phloeoterma, he claims, but the outer division product is that which gives origin to the endodermis. Bower: The comparative examination of the meristems of ferns as a phylo- §enetic study. Ann. Bot. 3: 323. 188 *ZENETTI: of. cit. 64. 27 STRASBURGER : of. cit. 449. 404 BOTANICAL GAZETTE [DECEMBER Now, at the time the protoxylem elements appear, I did not find, in the species examined, the cells of the endodermis cor- radial with those lying centrad. It is true that in younger stages the cells in this region are in radial rows; but nearer still to the punctum vegetationis this is approximately true of all the cells of the stem. At this earliest stage one would hesitate to say, because certain cells were corradial, that they were therefore division products of the same mother cells; so Zenetti’s con- clusion, based on this sole argument, scarcely seems conclusive, even granting the correctness of his observation. If, too, such a conclusion were correct there would be the curious anomaly of certain phloem and cortical tissues having a common origin. Evidently the study of transverse sections cannot settle the matter. To attempt to follow these layers upwards is obviously | only possible in median longitudinal sections. But in the stems of the Osmundaceae the leaf traces are exceedingly numerous, and at the growing point are closely packed together, and appear before the tissues of the cauline central cylinder become at all differentiated. Hence, no matter what be the plane of section, the endodermis cannot be traced continuously very far anteriorly to the point at which it is differentiated, for a leaf trace is certain to intervene; andI found it quite out of the question to pick out an undifferentiated endodermis on the side of the leaf trace turned towards the apex. Therefore, every attempt failed to refer the endodermis and the rows of cells ‘occupying the place of the pericycle” to the same initial layer. The typical protophloem, and the “quergestreckte Zellen e begin to be differentiated simultaneously with the appearance of the protoxylem. They are best examined in tangential sections. Their walls at this time become pitted, and their contents much less granular than those of the surrounding cells. Here, as in the maturer parts of the stem, there appear to be no differences between the typical protophloem and the “quergestreckte Zel- len.” Their relation to the leaf traces seems to explain their irregularity in orientation. Immediately below the point of ori- gin of a leaf trace they are arranged with their long axes parallel 1901 | ANATOMY OF THE OSMUNDACEAE 405 to the long axis of the stem, and there is a gradual transition to the tangential position. More than this, the laterally placed protophloem cells of the leaf traces can be directly traced into the ‘‘ quergestreckete Zellen” of the stem. There seems little doubt, therefore, as to their nature. To summarize observations : (1) The “ quergestreckte Zel- len” are sieve tubes, as has been demonstrated above ; (2) they become differentiated at the same time as the typical proto- phloem, and (3) occupy the same relative position; (4) they resemble the protophloem cells in form ; (5) their orientation is not uniform; (6) they pass imperceptibly into the longitudi- nally orientated protophloem cells of the leaf traces. Hence there seems no reason to regard them as anything else than pro- tophloem, CONCLUSIONS. The question now remains, how to interpret the vascular sys- . tem of the Osmundaceae. To do this more intelligibly, it will be well to recapitulate the main fibrovascular theories. We shall begin with that of Sachs and DeBary. These botanists regarded the bundle as the unit, and the vas- cular system as a more or less simple complex of bundles embedded in ground or fundamental tissue. Developmental Studies have shown that this theory is inadequate, for the unit is wrong, The hypothesis which at present obtains was supplied by Van Tieghem and Strasburger. In this conception” the stele is the unit. The primitive form of stele, the monostele, such as occurs for example in most roots and in the stems of lycopods, is a solid central strand of xylem, surrounded by a sheath of phloem, and marked off from the cortex by the differentiated internal Cortical layer, the endodermis. Of this there are many modifi- Cations, of which mention is made of the most important. By the repeated bifurcation of the monostele, the polystelic type is VAN TIEGHEM: Traité de Botanique 673, 765- *® VAN TIEGHEM: Sur la polystélie. Ann. Sci. Nat. Bot. VII. 3: 275. *®VaN TieGHEM: Eléments de Botanique 1:84, 179- 406 BOTANICAL GAZETTE [DECEMBER presented, as in Primula and Pteris, each segment being in every respect a stele. If these steles fuse laterally, thus forming a ring with internal and external phloem, the gamostele is pro- duced as illustrated by Marsilia. Again, when parenchyma seg- regates in the axis of the monostele, and the vascular ring is broken into strands by ectad extensions of this pith (the medul- lary rays), we have the medullated monostelic type, such as is common in phanerogams. It is to be noted that the medullary and cortical tissues are considered by both these botanists to be of morphologically different value. Now by the bending in of the endodermis of the medullated monostele between the bundles, and the fusion of thé ends of adjacent groups on the centrad side of the bundle, so that each bundle has its endoder- mal sheath, and medulla and cortex become continuous, the schizostelic or astelic type results. Of this phenomenon Ranun- culus and Equisetum afford examples. A modification of this type, the gamodesmic-schizostelic, is produced by the lateral fusion of these endodermal sheaths, so that there is a common internal and a common external endodermis. If the internal endodermis degenerates, as it does in Z. arvense, then there is evidently a simulation of the medullated monostele. It is fair to add that Strasburger dissents 3* from the last two types des- cribed, the astelic and the gamodesmic, for he regards the endo- dermal sheaths about the bundles in the first of these, and the internal endodermis in the second, as not morphologically phloeotermal, but originating from specialized stelar cells. The researches of Gwynne-Vaughan* and Jeffrey # have shown that the phenomena said to lead up to polystely do not occur in Primula and Pteris. If the polystelic conception falls, obviously gamostely goes too. Further, astely has been shown, where it occurs in Equisetum and Ranunculus, to be preceded by the gamodesmic appearance. Later the internal and external 3* STRASBURGER : of. cit. Pp. 442. ; = OWE Vepenae < Polystely and the genus Primula. Ann. Bot. 11: 307- % JERFREY : Morphology of the central cylinder of angiosperms. Trans. Canad. Inst. 6 :—. (1-40) 1900. Igor | ANATOMY OF THE OSMUNDACEAE 407 endodermis may fuse between the bundles, but in no case is there an inward looping of the endodermis. Finally, the stelar origin of the pith of the medullated monostele has been disputed, and the question raised as to whether the medullary and cortical tissues are in reality morphologically different. In other words, is the medullated monostelic type primitive, as its simplicity might indicate, or has it resulted by degeneration from more complex types ? It is interesting to note that Potonié had discussed this last question from the standpoint of fossil botany, and concludes™ that it seems evident in the case of certain groups, such as the cycads, that the simple results from the complex (for example, the cycads from the Medulloseae). Hence for these groups at least he is inclined to reject this idea of segregation of parenchyma in the center of the protostele to form the medullated monostele, but holds that the medullated monostelic type has probably arisen by degeneration from his “pericaulom.” Since this peri- caulom was produced, according to his theory, by the lateral fusion of leaf bases in the stem surrounding the originally solid stele, the ‘‘urcaulom,” the medullated monostele has been derived from a form of central cylinder such as Van Tieghem has described as polystelic, preceded or accompanied by the dis- appearance of the enclosed urcaulom. The paleontological evidence, however, appears not to be conclusive, for in the very group that Potonié cites, the cycads, so eminent a paleobotanist as Dr. D. H. Scott takes a directly opposite view. He points out that the vascular system of the Medulloseae was typically polystelic, while in the recent cycads there is but one vascular cylinder, and that hence “we should involve ourselves in unneces- sary complications if we endeavored to derive the simple, primary Structure of the cycadean stem from the more elaborate organi- zation of a Medullosa. It is far more natural to suppose that the monostelic cycads arose from monostelic ancestors.” PoToniE: Die Metamorphose der Pflanzen im Lichte palzontologischer Thatsachen 22. 33Scotr: Studies in fossil botany 395. 1900. 408 BOTANICAL GAZETTE [DECEMBER In 1897, Dr. E. C. Jeffrey put forward another view of the vascular system, based upon a study of the young sporophyte. Here, too, the stele is the unit. According to this conception there are two primitive types of vascular axes; the first the same as Van Tieghem’s primitive type, and designated “ protostelic ;” the second, one in which there is a hollow cylinder, or ‘siphono- stele,” whose external wall abuts on the cortex, and whose internal wall encloses the medulla, and which possesses internal as well as external phloem. This is the ‘‘amphiphloic siphono- stelic” type, called by Van Tieghem the ‘‘polystelic.” The commonly called “‘astelic’’ modification results from the amphi- phloic type by a degeneration of the internal phloem, and the medullated monostelic type of Van Tieghem is derived from the astelic by the loss of the internal phloeoterma or endodermis. A study of development from the seedling is likely to show how these and other modifications in the stellar structure have been derived from the primitive types. Attention is also called to certain portions of the wall of the siphonostele in which the vascular tissues do not develop. These places lie above the points of exit of branch traces, and of leaf traces, and are known as ramular and foliar gaps respectively. Through these gaps the tissues outside and inside connect. In transverse section, the connecting tissues seen constitute the medullary rays, and the segments of the woody cylinder with adjacent phloem and parenchyma the bundles. A fact of great phylogenetic impor- tance in dealing with ‘“‘gaps” was further pointed out, namely, that in small leaved plants, as in the Lycopodiaceae, Equise- taceae, etc., only ramular gaps occur. These plants are grouped in the division Lycopsida, and their steles are said to be cladosiphonic. In all other vascular plants there is a gap for every leaf. These constitute the large leaved plants, the Pteropsida, and their steles are said to be phyllosiphonic. As a matter of theory, it is suggested that the siphonostele arose from the protostele for mechanical causes, in the Lycopsida to support the branches, and in the Pteropsida to support the 3©JEFFREY: Trans. Brit. Assn. Toronto. 1897. Igor] ANATOMY OF THE OSMUNDACEAE 409 leaves. Potonié also explains the origin of his second primitive type the ‘pericaulom,” the homologue of the siphonostele, on mechanical grounds. In the light of these theories we can now apply ourselves to an interpretation of the anatomy of the vascular system of the Osmundaceae, and likewise note if the facts already dealt with throw any light on the theories. First, we are in a better position now to decide whether the internal endodermis of O. cinnamomea is phloeotermal or not. It has been noted that in similar cases, that is, in gamodesmic stems, Strasburger has denied the phloeotermal character of the internal endodermis. With regard to the internal endodermis the following facts have been observed: I. There is present the characteristic cuticularized ‘“ radial dot.”’ 2. The structure and contents of the cells are materially the Same as of the external endodermis. 3. The sheath is continued into the portions which in some individuals present the phenomenon of internal phloem, just as in any form called by Van Tieghem and Strasburger gamostelic. In the gamostelic type the phloeotermal character of the internal €ndodermis has been admitted. 4. It generally connects with the external endodermis through ramular gaps, and by no means rarely through foliar gaps. When this occurs, there is no point at which it could be said that the one stops or the other begins. Having verified these facts in a great many cases, lam there- fore of the Opinion that the internal and the external endodermis are homologous tissues. Second, are the medullary tissues morphologically equivalent to the cortical ? Again we recapitulate observations. 1. They do not differ in structure or in contents. 2. The medulla very often contains brown sclerenchyma, at least in three species studied, a tissue which, in other ferns, never constitutes a part of the stele. 3. Medulla and cortex connect more frequently than not 410 BOTANICAL GAZETTE | DECEMBER through the ramular gaps in O. cimnamomea, and occasionally through foliar gaps; and neither is there a transition in the nature of the connecting tissues, nor any line at which we can say, the cortical tissues lie externally to this and the medullary tissues internally. 4. The cortical and the medullary brown sclerenchyma some- times fuse through ramular gaps in O. cimnamomea. 5. Portions of stem of O. cimnamomea have been found which are of the ‘‘gamostelic” type of Van Tieghem. The medulla in gamosteles is granted to be morphologically a cortical tissue. The conclusion is evident for O. cimnamomea at least, and if it be granted that the medullary tissues of this species are morphologically equivalent to the cortical tissues, then biological principles alone would demand a like conclusion for the other species. Third, of what type is the vascular system of O. cinnamomea? Again the facts must form the basis for a decision: 1. The young stem of O. cinnamomea possesses an entirely closed hollow vascular cylinder, sheathed with phloem and broken only immediately above the exit of a leaf trace ; and et a level higher up the cylinder is entirely closed again. There is a medulla and an internal endodermis. 2. In older plants the leaves are more frequent, and the gaps extend through several internodes; but yet the cylinder is the unit. The cylinder of phloem is quite rarely broken, except - where branching takes place. 3. There is an internal endodermis which is persistent throughout the entire central cylinder of the stem. 4. As a rule the internal endodermis bends out opposite leaf gaps. 5. There is an internal phloem in portions of some plants. 6. Not only does the cylinder of external phloem remain practically unbroken, but opposite leaf gaps there is on the inner side a proliferation of sieve tubes. In O. regalis Janczewski found isolated sieve tubes in the parenchyma filling the leaf gap; and the same thing is true of O. cinnamomea. 1901] ANATOMY OF THE OSMUNDACEAE 4la According to Van Tieghem’s stelar theory, the last two facts can be explained only by considering the central cylinder of the Osmundaceae to be “gamostelic.” The centrad exten- sions of the phloem opposite the medullary rays could then be explained by assuming that steles had united laterally, with the disappearance of phloem on the medullary side, but with the partial persistence of phloem on the radial planes. This would also explain the occurrence of internal phloem, the union of internal and external endodermis, and the homology of medul- lary and cortical tissue. But from the study of development there is not a shred of evidence to prove that there has been a union of steles. In fact, such a study shows distinctly that there is but one stele in the stem of O. cinnamomea from the very first. Van Tieghem’s observations on O. regalis have already been quoted (see INrRopUCTION); so we cannot describe the cauline vascular system as “gamostelic,”’ if this name implies a union of steles. There remains yet another interpretation, namely, that the vascular system of the stem of O. cimnamomea is a siphonostele in which some degeneration from the primitive type has taken place. It has been pointed out in a description of the concep- tion of the vascular system held by Dr. E. C. Jeffrey that the most primitive siphonostele is the amphiphloic siphonostele. In this there is an internal phloem and phloeoterma, and in its phyllosiphonic form there are wide leaf gaps and branch gaps through which jnternal and external phloem, internal and external phloeterma, and medulla and cortex connect with each other. In O. cinnamomea the gaps in this primitive type have closed somewhat, so that medulla and cortex rarely connect except through ramular gaps. Also the phloem forms an almost unbroken cylinder, and the centrad proliferations opposite the medullary rays are the vestigial relics of connection between external and internal phloem. The internal phloem has also disappeared in greater part. With such a conception of the cauline vascular system of 0. cinnamomea, the centrad accumulation of sieve tubes opposite 412 BOTANICAL GAZETTE [DECEMBER the medullary rays, the occasional presence of sieve tubes in the medullary rays, the fact of the internal phloem, the connection of medulla and cortex through ramular and foliar gaps, the presence of sclerenchyma in the medulla, the bending out of the internal endodermis into the leaf gaps, and the facts of develop- ment, all become intelligible. Fourth, which of the species studied possesses the most prim- itive type of central cylinder ? After a fairly comprehensive study there is one feature that stands out prominently, the great similarity and uniformity of vascular structure in the various species of Osmunda and Todea. According to Solms-Laubach the stems of fossil remains of this family, of which none earlier than the Tertiary have been found, do not present any striking differences from the living represen- tatives. Paleobotany, therefore, offers no solution to the prob- lem. In spite of the conservatism of the central cylinder, there are, however, minor anatomical differences. On the basis of these alone, without referring to the young sporophytes, I think there is sufficient warrant for placing O. cénnamomea at one end of the series, possessing as it does an internal endodermis, inter- nal phloem, and wide ramular gaps. It is difficult to say which species is to be placed at the other end of the series. In view of the fact that O. Claytoniana never has sclerenchyma in the. medulla, that there are small or even no ramular gaps, no inter- nal sclerenchyma, and even a degenerated external endodermis, we may not be far astray in putting it in. the position farthest from O. cinnamomea.. Now of these two, which retains a cen- tral cylinder more nearly primitive? If O. regalis has a medul- lated monostelic central cylinder, as has hitherto been claimed for it, then O. Claytoniana has also, and therefore, according to Van Tieghem, a more primitive form than that of O. cimmamomed. Assuming the correctness of this for the moment, it will be in order next to see if such phenomena as presented by 0. cinna- momea could be derived according to Van Tieghem’s hypothesis from such a simple medullated monostelic form as that of 0. Claytoniana. 1go1 | ANATOMY OF THE OSMUNDACEAE 413 The phloem sheath must have broken into bundles, and the endodermis must have looped in between the bundles, and con- nected around them on the centrad side. With the formation of this astelic type some of the cortex would have been included in the medulla, in evidence of which the sclerenchyma in the pith would stand as proof. Then next the bundles must have fused laterally to produce the gamodesmic type in which there is an external and an internal endodermis. Granting that the central cylinder could be so plastic in a single species, there are left yet to be explained the continuous sheath of phloem, the proliferation of sieve tubes opposite the medullary rays, the occurrence of isolated tubes in the medullary rays, the occur- rence of internal phloem, and the phenomena of the ramular gaps. Further, there are no facts in development that point to such a series of changes. Turning now to the other alternative, namely, the possibility that O. cinnamomea has the more primitive form of central cyl- inder, it will be granted that by the degeneration of internal phloem, endodermis, and medullary sclerenchyma, and by the closing of the ramular gaps the central cylinder such as we find in O. Claytoniana would result. In proof that such degeneration could have taken place, it is to be noted (1) that in O. cinnamomea itself, it has been pointed out that the amphiphloic condition is localized, that the internal endodermis has already begun to degenerate, that medullary sclerenchyma is not a constant fea- ture, and that closed steles above the point of branching are not at all uncommon; and (2) in further proof, analogous cases of degeneration within the same genus are frequent. Thus within the genus Equisetum two species such as £. arvense and &. hiemale may be chosen, the first long considered medullated monostelic and more primitive, the second gamodesmic and considerably modified. But a study of development and of nodal por- tions of the stem has shown that &. arvense has a reduced cen- tral cylinder, the product of degeneration from .. gemedesic type, and that therefore &. hiemale is nearer the primitive. Sim- ilar cases of degeneration have been pointed out by Van 414 BOTANICAL GAZETTE [DECEMBER Tieghem, Poirault, and Jeffrey, in the genera Ophioglossum, Botrychium, Equisetum, Ranunculus, etc. Very lately Boodle, 37; 38 has called attention to an interesting series of central cylinders in the family Schizaeaceae. Aneimia Phyllitidis has a ring of separate bundles, each with a band of xylem surrounded bya phloem, pericycle, and endodermis of its own; A. Mexicana has a complete ring of xylem in the internodes with external and internal cylinders of phloem and endodermis ; Schizaea has a ring of xylem surrounding a central pith, but no internal phloem or endodermis. It is likely that here, too, the Schizaea type is derived from the Aneimia type by degeneration. In the Hymen- ophyllaceae likewise, every grade is found from the case in which the phloem of the solid stele forms a complete ring to that in which it is developed on one side only. After examining a number of comparatively young specimens of O. Claytoniana, 1 am somewhat doubtful if the study of the development of this species will throw any further light on the subject of morphology; but for O. regalts 1 am more hope- ful. Nevertheless, aside from further developmental proofs, I incline to the view that O. cinnamomea possesses the most primitive type of central cylinder. I again recapitulate the reasons : 1. The opposite view demands a very plastic central cylinder in one species alone, not differing very greatly in habit from the others. 2. There would still remain phenomena that the opposite view could not explain. 3. There are no facts of development even in analogous cases to support the opposite opinion. 4. The view adopted here demands only slight changes, and those are of degeneration, to explain all the phenomena. 5. There are precisely similar analogous cases of degenera- tion. 6. Within the species O. cinnamomea itself, every phase of - 3 BOODLE: Stem structure in Schizaeaceae, etc. Brit. Assn. Dover, 1899. ea BooDLE: On the anatomy of the Hymenophyllaceae. Ann. Bot. 14: 455- 1901] ANATOMY OF THE OSMUNDACEAE 415 degeneration except the entire disappearance of internal endo- dermis is. observable in suitable specimens. When we attempt to orient the other species amongst them- selves, the task is more difficult, and of little importance. As already indicated, a closer study of development may afford more precise proofs. In the mature stems we have seen that O. regalis occasionally has sclerenchyma in the medulla, that there are ramular gaps, though usually small, and that the external endodermis is well developed. In OQ. Claytoniana, on the other hand, sclerenchyma is never found in the medulla, ramular gaps are infrequent, and the external endodermis shows indications of degeneration. In neither of these species is internal endodermis or internal phloem present. The probability, therefore, is that in the genus Osmunda there is a series, O. cinmamomea possessing the most primitive type of central cylinder and O. Claytoniana the most degenerate, O. vegalis occupying a middle position, but nearer to the latter. It is merely interesting to note in passing that Professor Campbell concluded from his study of the pro- thallia of O. Claytoniana and O. cinnamomea, that the gametophyte of the former was more specialized in many particulars, in other words, was less primitive in type than the latter. fifth, does a study of the vascular system help to determine the phylogenetic position of the Osmundaceae ? It was stated at the beginning of this paper that botanists have regarded the: Osmundaceae as possessing an anomalous form of central cylinder among the Filicales, their reason being that it seemed to present more of the features of a central cylinder such as is typical for dicotyledons, that is, a medullated Monostele in Van Tieghem’s terminology. In determining the Position of the family, therefore, in any natural system of Classification, it was hopeless to try to reconcile this single dicotyledonous character with the remaining filicinean characters, and so the vascular system in the family was regarded as anomalous. CAMPBELL: On the prothallium and embryo of 0. Claytoniana and O. cinna- ™omea. Ann. Bot. 6: 49. 1892. 416 BOTANICAL GAZETTE [DECEMBER It is fair to note that Zenetti dissented4* from the prevailing view, and evidently for the reason that he attached some value to the nature of the central cylinder from the phylogenetic standpoint. Hence he sought to find the same type amongst the vascular cryptogams. He rejected the ordinary fern type because it is ‘polystelic,” and the lycopod type because there is no pith, obviously overlooking Se/aginella laevigata, Phylloglos- sum, etc. So finding no living form with which comparison could be established he turned to paleophytology. Among the Lepi- dodendraceae he found the prototype sought for, especially in such of these fossils as L. Harcourtii, and the Sigillarians, because in these the wood is broken into bundles between which there are medullary rays. But he evidently did not grasp the signifi- cance of bundles and medullary rays in relation to leaf traces and branch traces. In O. regalis, too, the protoxylem is endarch, while in those ancient lycopods it was exarch. The stele of the Lepidodendraceae, as in all plants bearing palingenetically small leaves, was cladosiphonic, while O. regalis is phyllosiphonic, as are all primitively megaphyllous plants. Hence any attempt to establish a relation between the central cylinder of modern ferns and of those ancient horsetails must fail. Indeed, of the early fossil forms preserved, the one with a central cylinder most closely resembling that of the Osmundaceae, as has been pointed out by Dr. Scott,#" seems to be the cycadofilicinean Lyginoden- dron ( fig. 26). Further, we dissent just as strongly from the view that the family is anomalous in the matter of its vascular system. The typical fern stem possesses an amphiphloic siphonostele, as is especially revealed by a study of development. But degenerated forms of this are to be met with in almost every family, some examples of which have been noted. The Osmundaceae, as has been shown above, all exhibit some degree of degeneration from this type. It is therefore evident that the cauline vascular sys- tem of this family is neither primitive nor anomalous among the Filicales. * ZENETTI: of. cit. 73. 4ESCOTT : of. cit. Igor | ANATOMY OF THE OSMUNDACEAE 417 SUMMARY OF OBSERVATIONS. 1. An internal endodermis has been demonstrated in Osmunda cinnamomea, but in none of the other species examined. This inter- nal endodermis is in textural continuity with the external endo- dermis through branch gaps, and sometimes through foliar gaps. 2. Internal phloem has been found in OQ. cinnamomea in the region of branching. This is continuous with the external phloem through ramular gaps. 3. The external phloem of the Osmundaceae forms a contin- uous cylinder, a fact which De Bary has stated for O. regalis; and is not broken opposite the medullary rays as Strasburger has affirmed of the same species. Isolated sieve-tubes have been found in the medullary rays of O. cinnamomea. 4. The xylem forms a cylinder broken only by foliar and ramular gaps. 5. Brown sclerenchyma has been shown to be usually present in the medulla of QO. cinnamomea, not uncommonly in O. regalts, and not at all in O. Claytoniana. It occurs likewise in Todea barbara, but has not been observed in Z. superba. 6. The medullary and cortical tissues of the Osmundaceae are histologically equivalent. Brown sclerenchyma, which is not an intrastelar tissue in other ferns, occurs in both medulla and cortex; and in O,. cinnamomea the brown sclerenchyma of the medulla is in continuity with that of the cortex. 7. In O. cinnamomea the typical ramular gap is one through which internal and external ‘endodermis, internal and external phloem, cortex, and medulla connect. Every stage of degen- eration has been observed in Q. cinnamomea, however, down to the completely closed steles. O. regalis has a gap in the wood only, and O. Claytoniana usually none. 8. The so-called “quergestreckte Zellen” pointed oo by DeBary in OQ. vegalis, and more fully described by Zenetti, have been found in all the species studied. They are sieve tubes, Possessing all the characteristic features of sieve tubes, even that of callus plugs. Their irregularity of orientation is shared by the other peripheral tissues of the central cylinder, and is 418 BOTANICAL GAZETTE [DECEMBER apparently due to disturbance caused by the exit of the large leaf traces. g. Callus plugs have been demonstrated in the sieve tubes. 10. A study of the growing point has further shown that the ‘‘quergestreckte Zellen” and the typical protophloem are of the same kind; but it has failed to verify Strasburger’s statement that the pericycle and the endodermis arise from a common maternal layer. ‘11. The phloem forms a continuous sheath in the leaf. 12. The root possesses a a acialee diarch, occasionally triarch, vascular axis. SUMMARY OF CONCLUSIONS. 1. The internal endodermis in O. cinnamomea is to be regarded as phloeotermal in nature, a fact denied by Strasburger in homologous cases. 2. The medullary and cortical tissues seem to be morpho- logically equivalent. 3. Observations on the anatomy wt the Osmundaceae have been confined heretofore to the cosmopolitan O. regalis, and the subtropical Todeas. From these observations it was concluded by Van Tieghem that this family possessed a type of central cylinder anomalous among the vascular cryptogams, a type (the medullated monostelic type) peculiar to the phanerogams. The writer dissents from this view. It appears to be the case that the central cylinder of O. cinnamomea is not medullated monostelic, for the medulla is obviously extrastelar. Further, it cannot be regarded as gamodesmic on account of the topo- graphical distribution of the phloem. The most obvious inter- pretation seems to be that it is a degenerate form of the amphi- phloic siphonostelic type of central cylinder (polystelic type of Van Tieghem). O. cinnamomea, O. regalis, O. Claytoniana form a series arranged in order of degeneration of their central cylinders, and the same is true of 7. darbara and T. superba. The present research was carried on in the Biological Depart- ment of Toronto University under the direction of Dr. E. C. -_ 1901] ANATOMY OF THE OSMUNDACEAE 419 Jeffrey, to whom I wish here to express my obligations for his advice throughout. My thanks are due to Professor R. Ramsay Wright for the facilities afforded in the department. For some of the material used I am indebted to Mr. Oakes Ames, Assistant Director of the Botanical Gardens, Harvard University; Sir William Thistleton Dyer, Director of the Royal Gardens, Kew; Dr. Brodie, Toronto; and Mr. R. B. Thomson, B. A. UNIVERSITY OF TORONTO. EXPLANATION OF PLATES XIV-XVII. Abbreviations used, ep, Callus plugs. P, pericycle. ¢, cortex, pA, phloem. é. ¢., external cortex. p. ph., protophloem. z. c., internal cortex. px, protoxylem. é, endodermis. u, “ quergestreckte Zellen.” é. é., external endodermis. ¥7%, root. z. é., internal endodermis. _ 5s. s. s., strands. 4t, leaf trace. sc, sclerenchyma. m, medulla. x, xylem. m.r., medullary ray. PLATE XIV. I. Transverse section of the stem of Osmunda cinnamomea, Fic. 2. Transverse section of part of central cylinder of O. cinnamomea. - 3. Tangential section of O. regalis. Fig. 4. “Quergestreckte Zelle” of 7. barbara, showing sieve plates and Callus plugs. . FIG, 5. Transverse section of leaf trace of O. Claytoniana near the grow- ing point. Fig. 6. Transverse section of part of central cylinder of O. cinnamomea. Fig. 7. Sieve tubes of 7. barbara, showing sieve plates, “ globules bril- lants,” and callus plugs. ; Fic. 8. Transverse section of part of central cylinder of O. Claytontana. = PLATE XV. Fic. 9. Tangential section of Todea barbara, showing “ quergestreckte Zellen.” Fig. 10. Transverse section of O. cinnamomea, immediately above point of ramification, showing open branch gaps. 420 BOTANICAL GAZETTE | DECEMBER Fig..11. Transverse section of O. cinnamomea through nearly the same region in another plant. Fig. 12. Transverse section of same plant as in fig. zz, but lower down. FIG. = Transverse section of same plant as in fg. 72, but lower down. 1G. 14. Transverse section of the central cylinder of O. céanamomea, sin intectial endodermis and brown sclerenchyma in the medulla. PLATE XVI, Fig. 15. Transverse section of central cylinder of O. cimnamomea, show- ing internal endodermis and an absence of brown sclerenchyma in the medulla. Fic. 16. Transverse section of the stem of O. cimmamomea in the region of forking, showing absence of ramular gaps. Fic. 17. Transverse section of the stem of O. Clayfoniana. Fic. 18. Transverse section of the stem of O. Claytoniana in the region IG. 19. Transverse section of the stem of O. rega/zs in the region of IG. 20. Transverse section of the central cylinder of O. i i show- ing brown sclerenchyma in the medulla. PLATE XVII, FIG. 21. actin veews section of the central cylinder of O. cénmamomea, showing internal phloem Fie. 22. A part of tlie central cylinder of O. cinnamomea shown in fig. 2z more highly magnified. Fig. 23. A transverse section of the young sporophyte of O. Clayfoniana, showing one foliar gap, und the corresponding leaf trace opposite. IG. 24. Transverse section of the stem of 7. darbara, showing brown eimai in the medulla. G. 25. Transverse section of a part of the stem of 7. barbara nearer the sehte point. G. 26. Transverse section of Lyginodendron Oldhamium, showing 4 leaf gap, a leaf trace opposite, strands of sclerenchyma in the medulla, and strands of primary xylem centrad of the cylinder of secondary xylem. PLATE XIV BOTANICAL GAZETTE, XXXII \" , Be Vii ne a \ Waa N N= AT BRAT Si Dg K psy. SSCS es erry aoe i) *) os Ls re) i ce os xed par (tess _— eo . a li a i, PLATE XV BOTANICAL GAZETTE, XXXII > SS i if 3 fy, ale ee ne FAULL on OSMUNDACEAE. BOTANICAL GAZETTE, XXXII PLATE XVI i Pn. 19 20 FAULL on OSMUNDACEAE. BOTANICAL GAZETTE, XXXII PLATE XVII FAULL on OSMUNDACEAE. :* 4) Be eee er ee ee as Oe dee DRIEFER ARTICLES, SOME ERRONEOUS REFERENCES. In 1794 C. H. Persoon published in Roemer’s ewes Magazin fiir de Botanik an important article that seems to have been almost entirely overlooked. It begins on page 63 under the title, Meuer Versuch einer systematischen Eintheilung der Schwémme, and is con- tinued, pages 81-128, as Dispositio Methodica Fungorum, with four plates. This latter part was reprinted and considerably extended in 1797 as Zentamen Dispositionts Methodicae Fungorum. This work, from pages 1 to 48, is an exact copy of the 1794 article, so that any- One possessing the Zenfamen can give the correct reference to the original publication by simply adding 80 to the page number of any of the first 48 pages. The parts in the 1797 work that are new are the title-page, preface, and pages 49 to 76. The plates are the same. References for original descriptions in the main part of the Zen- famen should therefore be made to the 1794 paper instead. For Puccinia graminis and £. Circaeae, instead of 1797, as always given, the correct date is 1794, which is also the date of the original descrip- tion of the genus Uredo, instead of 1795 as given in Saccardo’s Sy//oge fungorum and in Pfeiffer’s Vomenclator Botanicus. C. H. Persoon also contributed ten species of Aecidium to Gmelin’s edition of Linne, Syst. Wat. 1791, and his name follows the descrip- tions. So we should write, for example, Aecidium Euphorbiae Pers.; Not Aecidium Euphorbiae Gmel. In Usteri’s Annalen der Botanik for 1796, pt. 19, p. 43, Persoon speaks of having contributed to Gmelin’s €dition of this work. The date of these species is sometimes given as 1796, which comes from reference to Gmelin’s edition of Linne, Syst. Veg., a work that is identical page for page, so far as the volume containing the fungi is concerned, with the earlier Sys¢. Vaz., except in the title- -page. In Hist. Fisica y Politica de Chile 8: 43, 1852, Montagne describes Puccinia Malvacearum Bertero, not P. Malvacearum Mont. as always written. That Montagne intended this is shown by his adding: ‘Solo me pertenecen el diagnosis y la descripcion.” Igor] 421 422 BOTANICAL GAZETTE [ DECEMBER In his Flora Fernandesiana 10. 1835, Montagne also writes Uredo Cestri Bertero and Uredo Hydrocotyles Bertero, not Montagne as quoted in references to the work.—E. W. D. Hotway, Decorah, Lowa. PUCCINIA INANIPES. THROUGH some error the description of this species (BOT. GAZ. 31: 332. 1901) is incomplete. It should be: Puccinia inanipes Diet. & Holw., n. sp.—Sori on both sides of the leaf, particularly on the upper, scattered, punctiform; uredosori brown; ° uredospores elliptical, brown, echinulate, 25-30 X 20—25p; teleutosori black ; teleutospores broadly elliptical, rounded at both ends, and when dry with both ends depressed, scarcely constricted, apex with a very slight cucullate thickening, smooth, dark chestnut-brown, 34-42 X 28-31, with long hyaline hollow pedicels which easily break at the base from the host plant. : On Lupatorium brevipes, Oaxaca, Oct. 18, 1899, no. 3677.-— E. W. D. Hotway, Decorah, Jowa. THE POSITION OF PLEUROCOCCUS AND MOSSES ON TREES. HavinG observed during the past winter that certain chlorophyll- containing plants do not grow most abundantly on the north side of trees, as is commonly supposed by the laity at least, and as stated in at least one of the more recent books on botany, the author herewith presents some of the results of his observations, which are still im progress. The trees on which the observations were made were located in 4 piece of woodland and were principally black oaks with a few white oaks, chestnuts, and beeches. The chlorophyli-containing plants found growing upon these trees were principally pleurococcus and some members of the Bryaceae. These were growing upon all sides on the bark of the trees except the southwest side, and approximately in the following ratio: In ro per cent. of the trees upon the west side; in ro per cent. upon the northwest side; in 10 per cent. upon the north side; in 20 per cent. upon the northeast side; in 35 per cent. upon the east side; and in 15 per cent. upon the southeast side. 1901 | BRIEFER ARTICLES 423 A further examination showed that they grew in the greatest pro- fusion on the shelving side of the trunks of trees with a slant of 10° to 20°; furthermore trunks which were nearly vertical were not inhabited by these minute plant forms. They were, however, nearly always found upon the slanting surfaces near the ground. In some cases the growth extended approximately to a height of 20 or 30 feet, and the position varied as the shelving varied, so that the eae might extend on the same tree at different heights from go° to 270 Rather careful observations thus far obtained tend to show that it is the shelving portions of the trunks of trees which receive and hold the greatest amount of moisture, and as the latter is apparently one of the most important requisites for the development of these green plants, we can readily understand why we find them distributed on nearly all sides of the trees, and not limited, as popularly supposed, to the north side of trees only.— Henry Kraemer, Philadelphia, Pa. CONTRIBUTIONS TO THE KNOWLEDGE OF THE PHYSI- OLOGY OF KARYOKINESIS.' (WITH ONE FIGURE) THE investigations, concerning which this is a preliminary report, were undertaken to throw more light upon the subject of the physiol- ogy of karyokinesis, a subject which has not received attention com- mensurate with its importance. This paper presents only the results regarding the relationship of light of various wave-lengths to the rapid- ity of mitotic nuclear division. The investigations were undertaken at the suggestion of Dr. E. Mead Wilcox and were completed under his direction. I take this opportunity to offer him my sincere thanks for constant and helpful suggestions throughout the course of the investigations. The roots of Aliium Cepa were selected as affording the most con- venient and suitable objects for the experiments. Bulbs of uniform medium size were selected with great care in the local markets. These bulbs were placed over suitable vessels in such a position that the base of the bulb barely touched the pure water with which the vessels were filled. The usual adventitious roots soon formed, and these were * Contribution from the Botanical Laboratory of the Oklahoma Agricultural and Mechanical College. I. Abstract of thesis. 424 BOTANICAL GAZETTE | DECEMBER allowed to attain a length of about one centimeter before the bulb was transferred to the light cages as hereinafter explained. For the purpose of securing light of various wave-lengths, use was made of the usual double walled bell glasses prepared as follows: The first bell glass was filled with pure water and the second one was painted with a very thick coat of lampblack. Two other bell glasses were filled with solutions 4 and & respectively. Solution 4 was made by adding toa 0.06 » solution of copper sulfate ammonia until the precipitate ceased to form. Solution B& consisted of 0.05 7 solution of potassium bichromate. For each of the experiments there were selected bulbs having roots at least one centimeter in length, and as many of these as possible were placed under each of the bell glasses prepared as above. The bulbs were then left under these glasses, so arranged as to allow of the normal respiration taking place, for between two and three days before the beginning of the experiment. Roots were then collected from the bulbs under each of the glasses at intervals of four hours during the twenty-four hours of the day. The tips of the roots thus secured were killed in the usual chrom-acetic-acid fixing mixture and imbedded in paraffin according to the usual method. The sections were cut 13.34 thick and stained according to Heidenhain’s iron-alum-haematoxylin method.’ By means of a very simple method, devised during the investiga- tion, it was then possible to count accurately the number of nuclei in the process of division and the number resting. A uniform combina- tion of ocular, tube length, and objective were used throughout the work. In each case the slide was so adjusted as to include in the field nearly all the portion of the root tip lying back of the apical cells, the field being so placed as to include these apical cells. Within this field thus selected it was a relatively simple matter to make very accurate counts of the dividing and resting nuclei. Nuclei were classed as dividing from the first indication of segregation of the chromatic mat- ter in the prophase to the formation of the daughter nuclei at the close of the telophase. The following tabulation shows the results thus far secured by the us¢ of the above described methods applied to this investigation. Each per- etc.) ——t 1901 | OPEN LETTERS 441 Again, it is claimed that the second publication of Schizonotus (Lindl. Introd. Nat. Syst. 81. 1830) is as a synonym. Again Mr. Rehder quotes correctly, but, as it appears to me, draws wrong conclusions. Lindley says: “ Spiraea sorbifolia (Schizonotus m.).”’ This is not synonymy, but annotation. Of course, “Schizonotus m.” is an abbreviation of “ Schizonotus mihi,’’ and the case is precisely the same as if Lindley had written in full: “Spiraea sorbifolia (which I have distinguished as a separate genus under the name Schtzonotus).” Surely this is not synonymy. If so, why does Lindley, two pages farther on, in naming the focus genera of aaa tic, enumerate “Spiraea, Gillenia, Schizonotus”’ Mr. Rehder admits that “in mer Lindley enumerates (p. 145) Schizonotus as a genus, and characterizes it (p. 441) by mentioning Sfiraea sorbifolia as the type.” What Lindley actually says (Juévod. Nat. Syst. ed. 2, 441) is this: “Schizonotus, Lind?. in Wall. Cat.—Spiraea sorbifolia, etc.” Am I pardon- able if I fail to see why the name is published any more satisfactorily here than in either of the two preceding cases? And as far as the enumeration of Schizonotus as a genus on page 145 is concerned, I have already called atten- tion to the fact that it was enumerated in the same way on page 83 of the first edition (1830). Of course, Basilima Raf. 1815 being a nomen nudum, Seringe’s sectional name Sordaria was the first under which this group of plants was distinguished, but it was not used ina generic sense until 1864. It is true that the replac- ing of properly published generic names by earlier sectional ones has been Suggested, but I am not aware that it has found any powerful advocate, even among extremists in nomenciatural reform, and I know of no botanist who has consistently carried out this principle. Yet such a principle seems to be the only excuse for the use of the name Sorbaria. Botanists who accept the oldest generic name must take up Schizonotus Lind]. 1828, while those (few of them in America, I believe) who adopt the generic name with which a spe- cific name is first combined must use Basi/ima Raf. 1836, if they are to be con- sistent with their principles. It is scarcely necessary to refer to Mr. Rehder’s remark that “it would be very unfortunate to revive the name Schizonotus, since it was applied afterwards and has been in use for two other genera,” as this is an argument which will carry little weight with most people, and one which I believe Mr. Rehder himself would hardly have advanced as the only reason for discarding Schiézonotus. “‘Sorbaria of course will not enter into the American flora if Chamae- batiaria is considered as constituting a distinct genus” is another statement to which I must take exception. On the contrary, Schizonotus sorbifolius (L.) Lindl. (Steud. Momencé. 531. 1841), the type of the genus, is peculiarly adapted to the conditions prevailing in the northeastern United States, and is winning its right to a place in our manuals as an introduced plant. It was reported 442 BOTANICAL GAZETTE [DECEMBER from central New York ten years ago (Peck, Rep. N. Y. State Mus. 44:[15]. 1891), and I found it myself in the northern part of this county (Westchester Co., N. Y.) in July 1895. It was at that time that I became interested in the synonymy of this genus, and Mr. Rehder’s article brought vividly to mind the researches which I then conducted upon this subject. Since writing the above I notice that the plant under discussion has been admitted to Dr. Britton’s recently published Manuva/—Joun HENDLEY BARNHART, Zarrytown, New York. (See ey NEWS. Dr. W. R. SHAw has been appointed botanist of the Oklahoma Experi- ment Station. AT THE UNIVERSITY OF WISCONSIN, Charles E, Allen has been appointed instructor in botany, and Mrs. George J. Ruger and H. A. Winkenwerder assistants in botany. HouGuton, MIFFLIN AND COMPANY announce that Sargent’s Sz/va of North America is to be supplemented by two additional volumes, containing 115 plates. The volumes will be published in 1902. A BIOGRAPHICAL SKETCH of Dr. Charles Mohr, prepared by Eugene i Smith, and accompanied by an excellent portrait, is published in the Novem- ber number of the Budletin of the Torrey Botanical Club. IN THE November number of the /ournad of the N. Y. Botanical Garden the report of Dr. N. L. Britton on his recent trip to the West Indies is pub- lished, and also the report of Professor L. M. Underwood on a trip to Porto Rico, AN EXCELLENT PORTRAIT of M. Maxime Cornu is published in Bud. Soc. Bot. de France of September last, in connection with an account of his obse- quies. The address at the tomb was given in behalf cf the society by M. Ed, Bureau. HERMAN B. DorNneER has been appointed assistant in botany in the Indiana Agricultural Experiment Station at Purdue University, vce Wm. Stuart, who has been transferred to the staff of the Horticultural department of the same station. Dr. HANS SOLEREDER, of Munich, has been appointed professor of botany and director of the Botanical Institute at the University at Erlangen. Dr. V. Schiffner, associate professor of systematic botany in the German Univer- sity at Prague, has been called to a similar position in the University of Munich.— Science. R. H. DENNISTON and H. G. Timberlake spent about two weeks in Sep- tember collecting for the State University in the lake region of Vilas county, Wisconsin. Special attention was given to the lower cryptogams, of which about 450 species were collected. These included about 350 fungi and lichens, 50 algae, and 50 mosses. Dr. THomAsS MEEHAN, the well-known horticulturist and botanist, died at his home in Germantown, Philadelphia, on November 1g, at the age T90r] 443 444 BOTANICAL GAZETTE | DECEMBER I9gOI of seventy-five. He was botanist of the State Board of Agriculture of Pennsylvania, in charge of the herbarium of the Binladelphin Academy of Sciences, and editor of Meehan’s Monthly. STEPHEN C. StunTz and Charles E. Allen, of the University of Wiscon- sin, spent the month of August and the early part of September in making collections and ecological notes of the flora of Isle Royale, Lake Superior, They worked chiefly about Rock Harbor, on the south shore of the island. Their collections include 480 numbers of spermatophytes, about go of pteri- dophytes, 340 of mosses and liverworts, 80 of lichens, and 650 of fungi and myxomycetes, Bulletin du Jardin Impérial Botanique de St.-Petersbourg is the French form of the title of a new Russian botanical journal, three numbers of which have appeared, under the editorship of A. Fischer de Waldheim. It is to publish original papers, critical reviews, and reports from the Garden. At the end of each paper a brief résumé in French or German is given. Prom- inent among the contributors to the first three numbers are A. Elenkin, the lichenologist, and A. Jaczewski, the mycologist. A GENERAL MEETING of botanists will be held at the University of Chi- cago on December 31, 1901, and January 1, 1902, in connection with the meeting of the American Society of Naturalists and affiliated societies. The local committee of arrangements extends an invitation to all botanists, whether members of any society or not, to attend the meetings (availing them- selves of the guaranteed railroad rate of a fare and one-third on the certifi- cate plan), to attend the reception and the annual dinner and to share in all provision made for the convenience and pleasure of visiting naturalists. Detailed announcements will be sent on request. NEARLY TEN years ago the late Professor Thomas A. Williams and Mr. David Griffiths, now of Takoma Park, D. C., agreed to issue sets of fungi of South Dakota. Later this plan was changed to include the west in general. The death of Professor Williams caused an abandonment of the undertaking entirely until a recent invoice showed that an unusual amount of very valu- able material had been accumulated. It is estimated that there are on hand about three centuries of specimens, containing many species recently described from South Dakota, Wyoming, Montana, and Arizona. The col- lection is rich in species of fungi that have never been distributed in any set of exsiccati. Mr. Griffiths, therefore, has determined to put the specimens up in packets with printed labels bearing the usual data, and to offer them for sale under the title West American Fungi. The original plan contemplated issuing fifty-four sets, but a much smaller number will now be put up. One century or three may be ordered, to be paid for when issued. The first cen- tury will be ready about the middle of December. GENERAL INDEX. The most genes classified entries will be found under Contributors, Personals, and Reviews. New names and names of new genera, species, and varieties, are printed in ates re synonyms in étalics. A A. A. S., Denver meeting 75 ies, needles of 356 cea 203, formation lela i pen formation 273; Far- formation 272; 3 nie ve. Ruigiolie ye viridiflora 326 ch ead ve, heteracantha sacdaaeg 283; Tou- on 71, 428; vegetation 281 Agriculture, Sheahan es Bureau of t Industry of the U. S., Depart- 55 x of 1 Alae, abnormal Saunders on new nthe cies of 374 Albugo, candida 91; gametogenesis and fertilization } in 77,1 m8 238; Portulacae 793 rin, Perel 5 a Amarantaceae, Lopriore on 71 Amblyolepis setigera are ___ Ainbrosia artemisiaefol nas America, n new é plas of 71 ) Anderson xander P, iB parser 154,379 ceca t furcatus 205 aroliniana 106 Angiosperm s, Hallier on classification of Annuals, formation bed 2 respi 201, 208 Antennaria, N Nilo 434 [DECEMBER, — Apospory, in Anthoceros, Lang on 376 Apples in cold stor orage, Corbett on 151 Aquilegia, pai Seng in 304 Arceuthobium pusillum "Wheeler on 64 Archegonia of Selgin nella apus 130 Argyrostachys, Lopriore on 71, 37 Aristida Se formation 201 we i 153 nold’s * The sea pa at ebb tide’ 222 Amol, work of 1 25% Artemisia frigida 263 Arthrostyliium, Pilger on 4 Arthur, J. C. 307; personal on 228; work of 3 ee a of 327 Asclepias, Cor Poe Strasburger on me in pee mbens 198; incarnata 326; phyt igiacdsides 326; speciosa 326; an 326; tuberosa as6: venient 26 Ashe, W. W., k of 68 Asplenium, Gilbert on 377 Auld and Gibson’s * Codium” 63 Autran, Eugéne, personal 227 B gage of leguminous plants, Burrage Bacterial disease of — Stuart on 429 Baldwinia, Harper wy Balfour, ‘Bagley, aes, 1 380 Ball, C. R., personal 156; work of 434 se er, H. work Ba ne . I5f, 432, aare x ohn H acne cy S., eel? of 430 Basilim: of 1 369, 373, 375, 431, _Basilin oe Bast Shere, middle lamella of 18 Beal, W. J., personal 228; work of 153 Beattie and Piper’s “ Flora of the Palouse Belajeff, w Bennett, Arthur, 58 445 446 Bergen, J. Y. en. 307 Bernard, C ie i ae Bessey, CE 9 "per. 228; “Struc- ture and Siacueh of diatoms” 150 Bessey, E. A., 64, 66, 67,151; personal 156 n ork of 4 iC wate on 434 ogue, E. E, work of 68 Bolley, H. L., work of 68 Bosqueiopsis, Wildeman and Durand on Botanical Society of America 380 Botany, lowa Summer School Bot nae of 76 ; chium ternatum Oneidense, Gilbert ke Rees r, F. O., personal 227; work of 178 Brackett, Gustavus B., personal 155 ay 262 -L., ee 1 75; of 71, 72, 153, 379, » Magee it tis flora of the tea ee: and Canada’”’ 426 k of 68 3ryonia dioica 36 3ulbilis each nace 119 Bulletin Jard. Imp. Bot., St. Petersbourg x I Bruchmann, work of 170 Bryobrittonia, Williams on 72 } ] I 44 Bumelia lycioides 268 mi grass 28 eta u, Ed., personal 443 ae Seviicuen, work of 429 ret Eloise, personal 155 Butten, F. K., work of 374 Cactaceae, structural studies on 35 Cactus Sica Calcium ceeles, tanetion of 142 Calyx, abnormal 351 Campanula rotundifolia 263 se DOA. personal 228; work of 4 Rea eae Peder Srey, ne ee Smee snomalin, ores 7 Carex, Komarov on 434 Garicacee: Urban on Carlton, Mark Alfred, 1 pais: oo on distribution of 431 Rete of Canullieies iydbes ja a8 Celery blight, Towne on ly Cell plate, theory of 3 BOTANICAL GAZETTE | DECEMBER Cell hoe : Cellulos Caccheus myosurcides 205 d on plants of a 153; Robinson on plants o ae Centrosomes, in Helosis and Lilium, Bernard on 152; Yamanouchi on 375 Ceram, a tropical forest in 21 Ceramothamnion, cana on 71 Cerastium, Green on 377 Cercospora apii, Toe on 64 giganteus, anatomy of 46, apadeatns of 3 Chaetonella, Car aes on 378 Chamaebati Chama ioe. “Small on 377 Chamberlain, C. J. 64, 66, 69, 150, 153,223 cemeb tay. es " Methods n plant t histology” Chapaeral als ale 2775 Paik 203; forma- tion sel 27 pare iss ” personal 156 Ches sa 15 NS ‘work of 151; papien Nees 70 Chla oa pene. bl on 150 “Manual of Chodat 70; 71 Chondroxioen, Schmid on 378 Christ, oe wo 153 “ Phyllotaxis” St! Clark, Anna M., wo of 68 ES Peo by a Billings on 433 Clei mous Howers, Du Sablon on 67 bnor Collins Color, spies NO of ao 332 Comber. 2 93, 2 vied dass middle “Sascell of 20 Guy N., pers 4 ompositae, ps boat on a a78 Conidia, homology of 168 Convolvulus a arvensis Pammel on 68 Contributo Charles. E. Arthur wy C. 369, 373: 375, aus 432, 435; John 440; Benn sey, A A. 64, 66, 67, 151; Bray, 1901 | - 195, 262; Chamberlain, C. J. 439; , 63, 64, 2225 Fairchild, ‘David es ke Faull, J. H. 381; Fernald, M..4:. ead Oo H — paar cea 4233 Land, W. fy G. 69; eo son i, 2 egg a be 4233 Bape 147, 14 ; Lyon, Pye 376,'4333 Maca m, A. fe 9; Oliv : By, 92; cree i 306; jase e B.4 Hg Maio » Carle- = E. 353 Rehder, ‘itred 56; Robert- 3 8 era "Albert 142; Shull, H 343; Smith, F. Grace 332; Stevens, ob 1: 77, 257s 2385 Thomas, M. B. Wilcox, E. Cook, O. e., aia 154, 156; work of 434 Copeland, E. B. 356; personal 74 Corbett, L. C., personal 156; work of 151 Cornu, Masons, death of 154; personal 4 Cornus, Small on 71 Corydalis aurea 2 106 Coulte Lae M. 6 153; 371, 373: 374, 2,79, 375, 377, 426, 427, 428, 429, 430, 431, 432, 434, 43 Coulter, Stanley, work of 429 aga Frederick V., personal 155; work 434 Cowies, H.C. 374, 376, 429, 430, 431; personal Crataegus, Ashe on 68; Britton on 71; Sargent on 71; Small > n 377 “oS pa bres on 377 ae on, species ey the U. S., Ferguson on ; tes s 289 “hand Toten of rine oxalate 142 Cucurbita Pepo 367; L n 63 Crypto ogams of Indiana "Thostias on 428 Cc a, Cook on 43 Cuscuta, oe aia he on wept Cylicomorpha, 71 mo. Urb Cynan choideae, pollen of 325 Cynanchum vincetoxicum, Strasburger on pollen in 423 Cypera raceae, Meinshausen on 434 Cytology, of Albugo 77, so 262; of Cyanophyceae 72 INDEX TO VOLUME XXXII D Dale, Elizabeth, work of 375 Dalla Torre Harms’s “ Genera Sipho- a maru Bo no, 1B 29, 439; personal 307 Linens Poa 288, formation 283 Dasystoma a Small on 378 a B. M. 61, 63, 64, 222; work of 152 ny Mara, work of 375 DeCandolle, Diospyros n 428; tomato, Be personal 443 B. M., personal 156 k of 377; and Wildeman’s Reliquiae Deweveanae” 373 Dusén, work of 375 E Earle, Frank S., personal 379; work of 70, Eastwood, Alice, personal 228; work of Echinocactus Wislizeni, anatomy of 44; assimilative tissue of 38 Echium vulgare, Pane of 344 cology of Texas Ehretia elliptica 2 368. cheamarnn septs on Chodat and Bernard on 70; of Nelumbo, Lyon on 3745 pir sala spinulose 171; pro- toplasmic continuity In Embryo-sac of Lilium simdideon, Bernard Rendodernia of a onegsamen 6 Endosperm, of cc Lapa 223; protoplasmic continuity Engler, A., work of 71; “ Palio - 27 Equisetum, Suksdorf on 377 remosphaera viridis 309 448 BOTANICAL GAZETTE oe Canadense, fasciation of 343 3 Ey work of 434 Eupatorium, more ale, Greene ~on ~ 71; brevipes 422; Robinson on 153 Experiment station in Porto "Rico 227 F Fairchild, David ae a personal 156 Fanning, W. G., of 374 Farlow, W. G., ne 227 : asciation 343 aull, J. Mas 381 re dde work of 378 Fendlera upicola 263 Fergus hak of 428 Ferguson, Mange t-C., 435 Sali ald, e500 work of 1 53 s, Christ on 153 “ber teealg duable in Zea Mays 69; lege 77, snare in Ginkgo biloba, Ikeno in Selaginella apus 13 Sraburge $ theory of 240 Fitting, work of 174 Fistulin otha, gre Sele on 70 | Flora, of Indiana, Coulter eee +E orto Ri on 443; of pot ‘lie Valley, peracid ifs 3745 ; of Sou kota, Saunders of S. E. ipinenibens Coulter on pale ees t Indies, Britton of Yukon, Britton and Ryd- = Q oO n 72 wers, abnormal 346; and insects 367 Aisi 28 m 218; oes 11, 262, 266; etpliyile a8 Forti, Achille, baler of 71 Frankenia Jamesii 287 Frasera specios ago ieee Nie a cajorsada 268 an, E. M., work of 374 rede work of 4 430 +» 325, 433 Fong from Porto Rico, — on 70, 377; Henning on on types of 378; of ae Sint o1 oa 430; of west America 4 Fungicides, aed on foliage, Sturgis on [DECEMBER G Galloway, Beverly T. 2 eens 155, 228 Gametogenesis in Albugo 77, 157, 238 a eg s of Selagine ella 124, 170 tory course in plant 18. Garden, New York Botanical 75 Garden en Irish on 428 ree rg ersonal 155 an, "i irae re 7 Gatling “Flora of Tennessee ’ eric nomenclature of cedar pate D eattn 42 xeneva eid 307 =| fay ofp i=] =) xesneriaceae, Urban on 431 oot and Auld’s ges iil 63 riesenhagen, work o ferutine exasperata, Humphrey on 374 Gilbert, Benjamin D., AQAPOOOQOgqda o < AS) : ° 5 ma = fe) 3 - of 6 Gow’s “F lowering sani of Adair county, low Gould, “HOP P., personal a Gramineae, Pilge er on pico formations 118, ue 198, 204, 207. 2 peace of Iowa, Pammel and Weems on 42 9% ms Kansasy —igiay Fi on 67; of Kentucky, G 7 Gray, ioe [Pelec ars ‘bles to 75 Greasewood formation 275 Greene, E. L., work of 71, 153, 377 Grits, David, personal 156, 444; seal Oe North American Sordar 427 Giese Ro bert F., personal 154 Growth, Naboki ch on anaerobic 435; periodicity et in eet 42,5 Gru d, w of 66 Giihertesia esta Rete ih H Halsted, B. D., personal 228 Hapalophragmium, Solow w on 378 Harkness, H. W., death of 154 = _ - a 1901 | Harms, H., work of 71; and eat Torre’s n “Genera Siphonogamarum alae R. M., work of 378 rshberger, J. W., work of 429 [SOS a i ale al 154 Havardia, Sm n 7I Hedlu nd’s UMahouaniile der Gattung Sorbus” 373 edoghyilix, Setchell on 70, 153 ieee pany 370 Heisen, work of 172 Helosis inenes: Chodat and Ber- nard on 70 oc agit pitas on 68 Henderson, L. F., k of 65 Henn nings, P., w my en Herbarium, committee on national 380; Hi n, Britt 3 Hilaria Jamesii formation 201 , work Histology of Osmundaceae 385 Hitchcock, A. S., personal ord work of 6 7 Hollick, A., personal 228 Holm, Theo., work of 377 Holodiseus He Holway, E. W. D. 421, 422; work of 374 Picke. Si Je personal 380 Horkelia, Rydberg Hottes, Chas. F. gtier 154 ae Mad all. A., se enanst 154; work 0, 71 Fitalsire H. B., work of 374 Hucnine. Geo orge C., personal 156» Hyams, C. W., work o Hybridity i in palms, Waugh on 68 Hydnu re er on 153 Bos ae of Cactaceae 37 Pesgindupting. Urban on 431 I Tanthe, Williams on 434 Ikeno, work o Ilex opaca, middle lamella of 23 Indiana, Coulte ron flora of 429 “ate “Sia Cook on 435 and flowers 367 Intercell substance 2 Ipomoea pan a 367 Irish, H. C., na gh INDEX TO VOLUME XXXII J Jatropha, oe 122; spathulata 122 Jepson, W. L., personal 1 Johnson, D = personal 154 Johnson, T.C. 3 Juglans, nigra III; rupestris [11 Juncoides, Small on 377 Juncus, Suksdorf on 37 Juniperus, monosperma 216; occidentalis 216; pachyphloea at, 262; sabinoides III, 212; formation 213; Virginiana ELE, 213 K Kahlenberg, Louis 437 penn physiol gy of 42 ney, Thomas H., personal Sh Kennel Urban on 431 Kofoi work of 150 Kolkwit, cae Komaroy, V. L., Kraemer, Henry 423 Krameria secundiflora 198 Krigia ‘eniphestealis 367 of 431 work of 434 L pease beeenice’s of Glasgow ag ; and gardens of Tokyo o Im 3 Lactariopsis, Hennings on 70 Lamella, origin and nature of the mid- dle I aR te Setchel es! n-Scribner, F., hee ai paps work I American n grass es, II” Gand. W. I. Lang, W. H., wen of 376 Lange, D., work of 374 Larrea Mexicana 287, formation 275 odoratus, abnormal flowers 346 Lauraceae, re on La 2: 152; gee FA, 229 jee dy e of “Ossian dace Leaves, Chas 345 heeriia srpscides, DuSablon on 67 Harms 0 personal 228 Lewis, A arr 42 Liatris cochaulactiys 16 450 BOTANICAL GAZETTE Lichens of Africa, Wainio on 430 Linaria, Canadensis 106; spuria, Du Sa- ., work of 71 - Lister, Arthur, work of 430 Liverworts, Howe on 71; of Africa, Ste- hani on 430, 431 ~ edge apa - Zs a Pyik 148, 292 Longo, Lopri as ein he 71, 377 Lycopus, Steele on a . M. 124, 170, 376, 433; personal Von: Harold, personal 15 $5 work of 374 Lysimachia terrestris, MacDougal on 65 Lupinus subcarnosus 210, os M Macallum, A. i i MacDougal, , personal 227, 228; work of 6s si Srebttook, of plant physi- ology” Macm Sie oo phe gst 155 Magnolia grandiflor 343 Mamillaria Grahami, anatomy of 48 arine algae of Africa, daciea on 430 Matsumura, J., of 377 Medulla of Deas 39; of Osmunda- 397 Medullary rays, middle lamella of 18 Meehan, Thomas, death of 443 Megasporangium, 8 Selaginella apus 131; of S. rupestris 13 ee ‘of Selaginella apus 126; of elampodium, Robinson on 153 liaceae, De Candolle on 71 “ee Suksdorf on 377 ne ee oe et, to Asa Gray 75; to T. < _SEEEEE. 2 P 7 EP cere Elmer D., tng ; work of 71 ate ote od sa ies y; Longe on a 6 Mena Fernald oe plants of 153; Robin- son on plants Mez, Carl, work of 71, 431 Microspor: , of Selaginella apus 435; of Situ 139 rupestris hater Selaginella apus 132; of 139 [DECEMBER , H., work of 377 Tafa work of Sea 137 Sena mall on 71 Mimoseae 268 Minne sot Botanical Studies 374; Sea- side Sta Mistletoe, Wh heeler on dwarf 68 Mitosis, simultaneou us 241 Miyake, Kiichi, work of 379 Mohr, Charles Aeath of 227; bibliogra- phy of 379; sketch of 443; “ Plant life of Alabama” 371 Molisch’s “ Milchsaft und Schleimsaft” reaps " pgrni n 378 Moore, G. T. 309; personal 74, 156 iinaces. of ‘Africa, Gepp on 430; of Greenland, Dusén on 375; of yuoe, Mushrooms, Henderson on 65; Hyams n 65 eae a of Africa, Lister on 43 Mycological notes, Williamson on 429 ta orhiza of arctic plants, Hesselman n3 hiveeackis Gilg on 71; Mez on 431 Myxoderma, Schinidle on 378 N Nabokich, work ; of 435 oS meeting of 444 eedles, Meisner on 356 Nala ubow, a work of 151 Nelson, Aven, personal 228; work of 68, Nelson, ae work of 434 Nelson -, personal 307 + Die scmeenceeey | ““Wahrnehmung des See trices I Nemophila, Eastwood on 70 Nerium oleander, middle lamella of 17 oe Dawson on a, Robinso 153 = aa poi cake ‘instability of Roches- toptera, Urban Sct as of Pellia, Davis on 152 Nutation of peas, Neljubow on 15! O geod — of live 265; formation of ices sane cual on 64 f a p fe Benet Ne wd oe Sa ae et RN Ne SAO | SR eg a eee CUE pe PY se Le eS oe Se Sg = ae = © reer. jor Spas Tee ET eR “4 ; ca iethl : Z s ree = | eal SS ne BS ate gaene is Aj 3 z ' : “ 7 Ee me z4 ars . a ' ‘ & i Igor} Oenothera biennis 367 Oliver, G. W. 306 Oogenesis in Albugo 159 Opuntia, arborescens 279; anatomy of 50; frutescens 279; ul- ae wri “ 49; leptocaulis, ‘anat- 1 Sho milative tissue of 38; Lindheimer aia 274, 278; phaea- antha, anatomy of 49 Orehis mascula Orton, Wm. A., personal 156 Osmotic pressure, effect on form 292 W. Jj. V., per arbuscula, > ssa ersonal 155 undaceae, anatom of 434; personal 74 Oxalis acetosella, Du Sablon on 67 2 Palisade tissue of Cactaceae 38 Palmetto, Trelease on a Pacific 428 Palms, Cook on Pammel, L. H., personal 228; work of 68, 430 omg rg on 153 Panic n 68; Lamson-Scribner odat on new Ros 71 ; Texana 74 a W., personal 156 sd Seige Chester on 151 Pein rce, “6. 1 Eee, 74 Pe elargonium m, abnormalities in 345 Pellaea, Trelease on a cristate 428; atro- purpurea, Treleas Pellia, Davis on nucleus of 152 Pentstemon, Rydberg 78 Persea Borbonia Personals : Allen, C. E. 443, 444; Ander- son 7 228; A Bessey, 7 Brackett, i: B. I 555 Britton, E. G. 228; Britton, N. L. 75; Bureau, E. 443; Butler, ‘Bloiee’ 155s Campbell, D. H. 228; Carleton es Chamber- INDEX TO VOLUME XXXII 451 0. 154, 156; evo E.. B. 74; Ess L. Dorner, H. B.4 443; Duggar, B. M. 156 6; Earle, F. S. 379; Eastwood, Alice 228; Fairchild, D. G riggs, R. i 154; Hall, H. M. 15 Hallowell, Susan M. 379; Halsted, Harkness 154; dp). 8 ; H. W. Hasselbring, H. 154; Hitchcock, A. 3. 156; Hollick, A. 228; eat _ - 80; Hottes, C. F. Fp: ; How 156; jepson, 7 rill, E. tk. Mele c. 227 4433 ee V.443; S culls ; Scho field, oe S. 156; Schrenk, R. . Pimnberlake, H. G. 4433 ie Pay Trelease, se ; Underw ~ Pfeffer, W., Philibert, Henri, death Phloem, of pa. es he wees on 452 BOTANICAL GAZETTE [ DECEMBER a aaeenibi eg Fe Dennis on 428 Fr oe 67; paylogeny of 165 Phyllocomos, Maste 153 Phylogeny ot Albuginaceae 157; of Mu- co of O aceae ae of pent honap oa Be: ‘of ro te dacs 165; of Saprolegniaceae I a ee SPNYs Flahault on nomencla- ture In 37 Pres: ms of 356 Pierce, Newton B., personal 156 Pieters, A. J., personal 156 Pilger, ; a work of 71, 431 palustris 111; ponderosa 106, 262; sil- vestris, middle lamella of 12; proto- plasmic continuity in 224 saps and Beattie’s “ F amy of the Palouse ion” 62 Plants of Isle Royale, Wheeler on 3743 of North Diskots. Bolley and Waldron . — . f South Dako f southeastern Tennessee, oe on 420; Rostowzew on drying 306 Pollen, wane of 325; : se on formation of 433; tube, Longo on Populus Fremont ui 2 Polyembry ry in Toline Gesneriana, Ernst o Port rter, Thoma Conrad, siete phy tablet . 74; biograp y of 37 ben diand, 's8, pence of 50; ets dorf on tote hiss ras ¢ n 70 ound an saa s “ Phytogeography ne a, Rydberg on 378 Prizes, Walker 380 is juliflora 105, 111, 198, 270; for- mation 272 Protarum. arum, Engler on 71 roteidy Jost on synthesis of 429 Pseudomonas, Smith on 430 Pica e roc 106, 262 Ptelea, Small Peridophytess Crna on African 430; orth American, Gilbert on 429; of rt ed bed ed et bed Iowa, Shimek on 153; of Yukon, Underwood on 72 Pteris aquilina 3913 cig lamella of Io bers nia, Circa 421; graminis 421; ee — eo, 421 eae 167 Q Quercus, Leaegee formation 214; Emoryi 216, 26 Gambe _ 217; — 216; earies es a 264; minor 264; un dulata 21 vie Wirsiniane 266; ‘Formeion 21 R Ramaley, ok personal 155, 228 Rehder, Alfred Respiration, intramolecular 303; of bar- ley, Entel 431 Restionaceae, Manes rs on 153 Revegetation of Trestle island, Lange 4 Se Arnold’s ‘The sea beach at ebb-tide”’ 2: 's “Structure rnc states and Canada” 420; Re erlain’ s “Methods in plant his- 372; Chester’s ‘ Manual of a ares: 370; Church’s “ “i Llenr “Codium” 63; Gow’s “Flowering plants of Adair county, Iowa” 373 sriffith’s “‘ North American Sordaria ceae” 427; Hedlund’s “ Monographie der Gattung Sorbus” 373 mson S er’s “A an grasses II” 62; MacDougal’s ‘“Text-boo plant k= Of physiology” 148; Mohr’s “Plant life of Alabama” 371; Molisch’s “ Milchsaft 6 “Phytogeography of Nebraska "374; Robinson’s “ Problems and possibilities Thon avon Et ropa”’ ee Ur bins s Sy icke Antil- lanae”’' 431; von Scheer k’s “ Factors FO eka gk are bre eee oe i Ne a i te = oy eee Te ee a ee aera Ba Y Oe Cale ewerees gis y -ano- f CHOCOL LATES: —) ASSORT) FANCY BOXES == BASKETS etc: ~\GNDIES SENT EVERYWHERE BY By MAIL EXPRES ame Orders by mail carefully executed. \ = = LS 863 snout WAY. = ny rte oat We Z, “te —G Pure! |: » COCOA~” HEALTHFUL TT 2 =CHOCOLATE SPLD AT OUR STORES *°* GROCERS ee t RIDE ACOCKHORSE To BanBurY CROSS. 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Fast Eastern Express eaves CHICAGO 3.00 Arrives NEw YorK 6.00 p. m., BOSTON 9.00 p. m. New York State sting eaves CHICAGO 5.30 Pp. Arrives NEw “York 8.45 p.m., eae 11.30 p. m. Atlantic Express Leaves CHICAGO I1.30 Pp. Arrives NEw YORK 7.00 a.m Detroit Night ee Through Pullman Sleeping Cars Leaves CHICAGO 9.35 p : Dining Car service Arrives DETROIT 7.15 a. m. is noted for its excellence. m. Boston 8.46 a. m. City Ticket Office, 119 Adams Street, Chicago. R. H. L’Hommeprev . : O. RUGGLE General Superintendent. General UN and Tick Agent. Big Four Route FROM CHICAGQ TO Indianapolis, Cincinnati, Louisville, the South and Southeast. THE SCENIC LINE TO Virginia Hot Springs and Washington, D.C., via the Picturesque sare tmger & OHIO R’Y, the short line to Asheville, “{ apt and Florida, W. J. LYNCH, G.P.& TA. =W. P, DEPPE, Ass’* G.P & TA CINCINNATI, O. J. C. TUCKER, G.N.A. 234 Clark St., CHICAGO MONON TRAINS FOR | Indianapolis Dayton MONON TRAINS FORT |i] Kexexexcxexexes_|ill La fayett W.BadenSpr Fr.Lick Spr. Louisville Cincinnati Asheville ee ee PARLOR AND DINING RS wy. Sakon tig lpia ager Sa ENT CARS BY W.H.MSDOEL REED.cem: pass.act. CHASH ROCKWELL PRES.& GEN'L MOA. re CHICAGO. LARGEST CAMERA IN THE WORLD c NSTRUCTED ESPECIALLY BY ORDER OF THE CHICAGO & ALTON RAILWA OTOGRAPH THE ALTON 1 L LIMITED, SEND A 20, STAMP TO GEO, J. CHARLTON, G. P. A., OC. & A, ee CHICAGO, by Py yar aND Boga EIVE ILLUSTRATED PAM WITH Perot ACCOUNT OF cist EXPOSURE pa fit WITH THE moe TRAORDINARY MACHINE, THE CONNECTING LINK between the EAST «=» WEST BUFFALO CHICAGO DETROIT |,.45\s| ST. LOUIS TOLEDO KANSAS CITY Free Reclining Chair Cars on all Wabash Trains Write for Information about any trip you may have in agen sg it “ sae business to assist those who trave F. A. ee. Ass’t Gen. Pass. Ag’t 97 ADAMS ST., CHICAGO C. S. CRANE, Gen. Pass, & Tk’t Ag’t ST. LOUIS, Mo. Field of ot EO Binocular. Field of Ordinary Binocular. Bausch 6 Lomb-Zeiss STEREO Binoculars Are used by the armies and navies of the great Bausch 6 Lomb Plastigmat £-6.8 The Perfect Photo Lens, as the above picture made Bausch é Lomb Prism Binoculars Stand next in el t les Buy Bausch & Lomb Optical Co. New York optically and mechanically, have immense field of view, power, and in addition, has the and give, as no other glass does, a STEREOSCOPIC image. r Booklet with five difficult pictures mailed fr excellen your camera with it. Buy it for your camera, Descriptive booklet matte on fete. D BY ALL DEALERS. Bausch & Lomb Optical Co. New York ROCHESTER,N.Y. Chicago ROCHESTER, N. Y. Chicago (| JUST OUT |; A PLATE ATTACHMENT au For No. 3 Folding Pocket Kodak 3 HAVE ONE ame TO YOURS & — the owner to use either plates or films to focus picture on he! ground glass PRACTICAL REASONABLE We also fit our celebrated Double Anastigmat Lens to these cameras No. 3 FOLDING POCKET KODAK, with GOERZ DOUBLE-AN NASTIGMAT, New Automatic TIB Shutter and a Attachment, with six holders, complete, $62.8 net c Plate Attachment and Six Holders, $7.G4, net. Write for further information to your dealer or to C. P. Goerz Optical Works 52 EAST UNION SQUARE, NEW YORK paWied 3) ee os pans t « SW) A DOCTOR’S REASON. tor, why do you tell nurse to use Ivory Soap?” Two or three years ago, the students at a college in which I am interested, bought some of each kind of soap for sale in the city and made analyses of them. The result was that purity, price and uniformity of quality, indicated Ivory as the soap to be recommended. Since then I direct my patients to use it exclusively.” Ivory Soap — 9942460 Per Cent. Pure. Be oad IN URIC ACID DIA- BUFFALO LITHIA: WATER "esha ATISM, ET C. THIS WATER DISSOLVES URIC ACID AND PHOSPHATIC SEDIMENTS, ETC., ETC. John 56; ssid ti aker, M.D., LL.D., Professor of Matera Medica and Therapeutics in "he sco Chiruretal College of Philadelphia, dics, in the New York Medical Journal, fune, 22, 1899: _ is doubly efficient in Rheumatism and Gout, It + BUFFALO LITHIA WATER dissolv f Uric — ae Posphatic sediments, as well as other products difficult of elimination, while at the same time it exerts a moderately stimu- lant effect upon the renal cells, and thereb facilitates the swift rehibval of insoluable imagerinis from the body. Without such action insoluable substances will precipitate in = paivion nae eg The intense suffering ation b — tozethet = a consecutive pyel and c egy are a heed stil ‘more importa nt. n it ¢ ae This service is performed by the BUFFALO L 1A WATER & Sicolina Seilaces which are ressandate for ‘the production of FALO L ITHIA } The late Hunter McGuire, M.D., LL. D., Formerly Presiient and Professor of Clinical Surgery, University College of Medicine, ” Richmond, Va., and Ex-President of the American Medicai Association, says? og as an alkaline diuretic is invaluable. In Uric Acid Gravel, and indeed in diseases generally dependent apon a Uric Acid Diathesis, it is a remedy of extraordinary potency. | have prescribed itin cases of Rheumatic Gout Ged has sg tie | ecenery remedies, — se good results. I have used it also in my own case, be a great sufferer r fro is malady, an derived more benefit from it than foar. hae Sree i medy.”’ Dr. P. B. Barringer, Professor of Physiology and Surgery, University of Virginia: ‘In more than twen years of practice | have =. rer as + ota -uric acid agent many times, have tried it in a great variety of forms, both in NATURA ATERS and in TABLETS. As ve result to this experience, I have no hesitation in stating oe for bboow § — tsi ee mune uric acid deposits in the ‘ALO ITHIA a og ons with it as a solvent ees porig aa has ‘egg ied limited, acl ? e to compare it one rae er for 0 their disadvantage; but for first class of conditions above set forth | feel that BUFFALO LITHIA WATER Sone” Dr. Thomas es port grad: of Paris (Formerly of Baltimore), Suggestor g Lithia as a Sole vent for Uric Acid, “Nothing | could say ase ‘add - have frequently to the well-known reputation of the BUFFALO LITHIA WATER. sed it with e results in URIC ACID DIATHESIS, RHEUMATISM, and GOUT, and with this obit I have ordered ie ~ Ase Lithia is in no form so valuable as where it exists in the carbonate, the form nature’s mode of solution and division in it is post 2h water which has passed through Lepidolite and Snneetonaie Mineral formations.” of Galea discharged Ad > Lernin Winco Spring No. 2. **It seems on the whole pe that the action of the water is PRIMARILY and MAINLY AERTED upon U wile URAT — vos _— these constituents occur along with and as cementing matter to Phosphatic or Oxalic us materials, the latter m Big ew So detached and broken down as to disintegrate the aterm pe whole in these cases, also admitting of Urethral discharge.” James ‘.. merase M. es A.M., Lo. —— Forme erly Professor of Physiology and Surpery in tha Medical Department q the University of Virginia, and President of the National Board of Health, says: LITHIA - in Ue Ac Sonica ih shighl ke cocigemannee aah peu esource. Profession as an article of Materia Medica.” BUFFALO LITHIA WATER 's ‘or sale by Grocers and Droggists generally. TESTIMONIALS WHICH DEFY ALL IMPUTATION OR QUESTIONS SENT TO ANY ADPRESS PROPRIETOR. BUFFALO LITHIA SPRINGS. VIRGINIA LOOK AT THE LABELS! THE GENUINE ~ BAKER'S CHOCOLATE UP IN PACKAGES LINE ThIESE MOST AND BEST MONEY ALTER BAKE o.,LTp. s ESTABLISHED i780. R & Co. DORCHESTER, MASS, WEBER IANOS All that Art has conceived or sKill devised inthe production ef a piano, is shown in the WEBER Case designs that are strikingly artistic and effective in the simple and ornate alike. Mahogany, Satinwood, White and Gold. Flemish Oak, and English Burl Wainut. Special cases that are correct replicas of the Louis XIV., Louis XV., Renaissance, and Colonial seit. WEBER WAREROOMS Fifth pees cor. 16th Street, = York. Wabas h Avenue, Chicago