Qk! | ‘Cue THE =< BoOrANIGAL GAZCITE EDITORS: JOHN MERLE COULTER AnD CHARLES REID BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF F THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS: J. C. ARTHUR Fritz Nout Purdue University. Boreas? of Bonn. CASIMIR ee OLLE, VoLNEY M. SPA Gen Univ city ‘of Michigan. J. B. DET ROLAND THAXT University of Padua. pity University. ADOLF ENGLER, WILLIAM Tere ‘Henne of Berlin. Missouri Botanical Garden. LEON GUIGNARD, ES. MansHaut Wa L’Ecole de Pharmacie. Uni Hemi ni "Cambridge. Rosert A. HA EUGEN. WARM Un vers a Wisconsin. University of Copenhagen Jinzo MATSUM Veir WITTROCK al Academy of Sciences, hanenas Tiwary. Tokyo. ya Stockholm VOLUME XXX JULY— DECEMBER, 1900 WITH TWENTY-THREE PLATES AND FORTY-ONE FIGURES IN THE TEXT Mo. Bot.Gardei., | 1901. CHICAGO, ILLINOIS PUBLISHED BY THE UNIVERSITY OF CHICAGO 1900 TABLE OF CONTENTS. On the endosperm and embryo of Peferonia pellucida (with plate 1) Sue Duncan S. Johnson I New or unrecorded mosses of North America. I. (with plates 11-v) J. Cardot and 1. Thériot 12 The Ye of the oo sac in some meapantage daar plants (with $s VIand VII) - - Karl M. Wiegand = 25 The developmen and function of the cell pn in pais plants (with plates V1II H. G. Timberlake 73,154 Contributions from the Cryptogamic Laboratory of Harvard University. XLIV. New or little known unicellular eae bag hope Cohnii (with George Thomas Moore 100 Origin of the cones of the multipolar spindle in Gladiolus ith plate x1I} vuther A. Lawson 145 Physiological observations on some perennial herbs (with sisi XIII) A. Rimbach 171 Contributions from the Rocky Mountain Herbarium. I - - Aven Nelson 189 Cell and nuclear division in Fudigo varians (with plate xiv) - &.A. Harper 217 Double fertilization in Compositae. pail S6ite from the Hull Botanical Laboratory. XXI (with ree XV and XV W.J.G. Land 252 A new chromogenic micrococcus (with four aa - - Mary Hefferan 261 On the nature of the stimulus which causes the change of form in polymorphic green alge. Contributions from the Hull Botanical Laboratory. XXII (with plates XVII and XVIII) - Burton Edward Livingston 289 Observations on Lessonia (with — doy - - Conway MacMillan 318 Studies in Crataegus. II - - - - 3: (CD pedald © 335. The achromatic spindle in the spore mother cells of Osmunda regalis. Contri- butions from the Hull Botanical Laboratory. - XXII (with plate xxi) R. Wilson Smith 361 Studies on chromogenic bacteria. I. Notes on the pigment of Bacillus poly- chromogenes (with sixteen figures) - E. M. Chamot and G. Thiry 378 Notes on the division of the cell and nucleus in liverworts (with plate XXII!) ; an Hook 394 BRIEFER ARTICLES — Some observations on apple tree anthracnose (with twelve figures) A. B. Cordiey 48 VOLUME Xxx] v vi CONTENTS [VOLUME XXX New i Naa checN and Cruciferae of the Sierra Madre, Chihuahua, q Mexico” - c B.L. Robinson 58 Note on the mechanics of the seed- oe awns of Stipa avenacea (with five figures) —- - - - - - L. Murbach 113 Some new species of Wyoming plants —- - - - Elias Nelson Some new North American mosses (with plate x1) - John M. Holzinger Notes of travel. III - . - - - DD. G. Fairchild Another note on the flower visits of oligotropic bees - Charles Robertson Photography in botany and in horticulture (with two figures) F, A. Waugh and J. Horace McFarland A new species of Neovossia (with one figure) - - ke. R. Hodson Note on the origin of tannin in galls - - - Henry Kraemer David Fisher Day (with portrait) — - ty - - John F. Cowell Observations on the root system of certain Cactacex - Carleton E. Preston Non-sexual propagation of Opuntia - - - - Carleton E. Preston” Gaurella = Gauropsis - - - - : - I.D.A.Cockerell 351 New species of Trimmatostroma - - - : M. W. Doherty 400 The International Botanical Congress —- - 404 Notes on the flora of the banks and sounds at Beaufort, N. C. Duncan S. Johnson 405 Notes on the validity of Asplenium ebenoides as a species - Wm. R. Maxon 410 — ‘CURRENT LITERATURE — 61, 131, 207, 277, 352, 416. . For titles see index under author’s name and Reviews. Papers noticed in “ Notes for Students” are indexed under cides name and subjects. News — 72, 142, 216, 288, 359, 430. DATES OF PUBLICATION. No. 1, July 19; No. 2, August 15; Sig 3, September 15; No. 4, October 15; No. 5, November 15; No. 6, December ERRATA. P. 9, line 2 from below for other read certain. P. 10, line 13, for five read six. P. 22, line 3, for 14 read 14. P. 22, line 4, after 1887 insert ). P. 182, line 14 from below; p- 183, line 16 — below; p. 185, line 14 fro! below ; p. 186, line 5, for Nothocaleis read Nothocalai _P. 185, last line; p. 187, line 16, for Fost read ‘a P. 213, line 11 from below, for Nawachsin read Nawaschin. XXX JULY, 1900 BOTANICAL GAZETTE JOHN M. COULTER anp CHARLES R. BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS J. €. ARTHUR Purdue University CASIMIR DECANDOLLE Geneva J B. DETONI University of Padua’ ADOLF ENGLER University of Berlin LEON GUIGNARD LEcole dé Pharmacte, Parts ROBERT A. HARPER Oniversily = Wictudie JINZO MATSUMUR imperial reais To aye FRITZ. NOLL fie Bice i Bonn VOLNEY. M. SP Un oe = ee Te ROLAND Sprints ER rvard University WILLIAM oe SE 2SSOuUTTE Paiwuical Garden H. cesar 6 WV A Un er “3 Cie EUGEN. 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No reduction i is sails to dealers or agents. | FOREIGN AGENTS: Great Britain WM. WESLEY & SON, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin _ St., Strand, London. 18 Shillings. SW. 46, Schonebergerstr. 17a, 18 Marks Vol. XXX, No. J Issued July 19, 1900 CONTENTS P i ON THE ENDOSPERM AND EMBRYO OF PEPEROMIA PELLUCIDA = PLATE a Duncan S. Joh - NEW OR UNRECORDED MOSSES OF NORTH AMERICA. I (witH PLATES I-v). /. Cardot and 1. Thériot - - - - . {2 THE DEVELOPMENT OF THE EMBRYO-SAC IN EN 4 NON ee PLANTS (WITH PLATES V1 AND Vil), Karl M. Wiegan 25 BRIEFER ARTICLES. SoME OBSERVATIONS ON APPLE TREE ANTHRACNOSE (WITH TWELVE FIGURES). A. B. Cordley . ¥ i a 5 = 3 i . ‘: NeW CARYOPHYLLACE® AND CRUCIFER@ OF THE SIERRA MADRE, CHIHUAHUA, MEX- 1co. B.L. Robinson - - “ - j - a : 58 CURRENT LITERATURE. BOOK REVIEWS - - - - . ce ‘. . = 61 THE FLORA OF MONTANA. THE CELL. 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THE PRUDENTIAL is INSURANCE ‘LE. fo COMPANY PROD EN vials . OF AMERICA pr he JOHN F. DRYDEN, President Home Office: Newark, N. J. VOLUME XXX NUMBER I BOTANICAL “GAZETTE FULY, 1900 ON THE ENDOSPERM AND EMBRYO OF PEPEROMIA PELLUCIDA. DuUNCAN S. JOHNSON. a (WITH PLATE I) In the spring of 1899 I began the study of the Piperacee, with well-preserved material of the genera Peperomia, Heckeria, and Piper, collected by the late Professor J. E. Humphrey on the unfortunate expedition to Jamaica in 1897. It was soon dis- covered that seeds were needed for sprouting in order to complete the work, and it was therefore laid aside temporarily and my time given to work on the related genus Saururus. The recent paper by Professor Campbell (’99) on Peperomia pellucida announced results differing from those I had obtained, and led me to reexamine my slides, with the result that I was satisfied with the essential correctness of my former observa- tions, and noted several interesting features in addition. As the intended detailed study of the group may be deferred for some time, an outline of the most important observations thus far made on Peperomia pellucida Kunth is given here. The flower consists of two stamens, and a carpel sessile in the axil of a top-shaped bract (dr, fig.1). The ovule is single, basal, and orthotropous (fig. 7), with a single integument and one archesporial cell. The development of the flower as far as fol- lowed agrees with the account given by Schmitz (’72), and the development of the macrospore in the nucellus is as described by Campbell. I 2 BOTANICAL GAZETTE [JULY The primary archesporial cell cuts off a tapetal.cell above, and then immediately forms the definitive macrospore below. The tapetum divides to three or four tiers of cells, which finally form, together with the outer layer of cells of the nucellus, a - persistent plug of yellowish thick-walled cells, directly under the micropyle (é#, figs. z, 2, 9, 73). The first four nuclei formed from the large macrospore nucleus are located at the periphery of the pretty dense proto- plast of the embryo-sac, arranged like the spores of a tetrad and connected by strands of granular cytoplasm (es, fig. 2). Soon after this we find eight ellipsoidal peripheral nuclei, imbedded in the cytoplasm surrounding the enlarged vacuole of the embryo-sac. Up to this stage the sequence of phenomena has not been very different from that in the normal angiosperm embryo-sac, except for the lack of the bipolar grouping usually found. But now each of the eight nuclei divides again, as Campbell has shown, to form an embryo-sac with sixteen similar nuclei, pretty uniformly distributed in the peripheral layer of cytoplasm (jig. 7). A little later than this, before the pollen tube reaches the embryo-sac, the cytoplasm begins to get denser about one of the nuclei at the top of the embryo-sac, and finally a definite limiting membrane surrounds this and forms the oosphere (0, fig. 1). This egg is not directly under the micro pyle, but is pushed aside slightly by the aggregation of a smaller amount of cytoplasm about a second nucleus at the top of the embryo-sac to form the single synergid, as we may call it from its position (sy, fig.z). The position of spindles in certain cases” seems to indicate that this is a sister cell to the oosphere. Some times other-nuclei are found near the egg, but often not, and in no case was there seen a definite massing of cytoplasm about any of these. At first the synergid does not have a definite wall, but later on, as it persists, at and after fertilization, a dis- tinct wall can be seen (sy, figs. 3, 7, 9, r7, f2), At about the time the pollen tube enters the egg certain of the remaining fourteen peripheral nuclei begin to move together to form a compact group, of usually eight nuclei, surrounded by 1900] ENDOSPERM AND EMBRYO OF PEPEROMIA 3 cytoplasm. This group may appear at the lower end of the embryo-sac, in the middle near the wall, or center, or above, and near or even in contact with the egg (espn, fig. 3). The remaining peripheral nuclei retain their position near the wall, at first as naked nuclei in the thin layer of cytoplasm, but later each of these and a small portion of cytoplasm is separated from the great mass of cytoplasm in the embryo-sac by a flat saucer-shaped wall (fn, figs. 3, 7,9, 72, 13). Nowhere was there noticed any tendency of a number of these to collect in a basal position, or anything in their behavior to suggest their homology with the antipodals of the typical angiosperm embryo-sac. The short top-shaped stamens bear two pollen sacs each. Certain pollen grains (three or four in a section showing fifteen fertile ones) remain with unthickened walls, but apparently are not used for the nourishment of the fertile ones. The nucleus of the finely reticulate-walled pollen grain divides to two at a time soon after the formation of the tapetum in the embryo-sac of the same flower. The pollen grains are shed after the embryo- sac has reached the four-nucleate stage. They lodge on the large abaxial lobe of the carpel (st, fig. 7), and grow downward through a conical mass of small-celled conducting or nutritive tissue to the fusion canal of the carpel, and thence to the micro- pyle (pt, fig. 1). Just when the division of the generative nucleus occurs was not made out with certainty, but in cases where the pollen tube had just reached the embryo-sac the pollen-tube nucleus was seen at its very tip, and a single large generative nucleus, with cytoplasm about it, at the level of the nucellus. In other cases of about the same age there were apparently two generative nuclei. No indication of a sterile prothailial cell was discovered. The pollen tube often extends some distance into the egg, and after its entrance two nearly similar nuclei are found within the egg. The exact fate of the pollen-tube nucleus and the second generative nucleus was not determined. _ For a considerable time after its entrance the male nucleus lies in the egg near or even in contact with the female, but without 4 BOTANICAL GAZETTE [JULY fusing with it (figs. 3, zz). Its presence in the egg seems, however, to have an important influence on the other contents of the embryo-sac. At the time of its entrance the group of eight nuclei, each with a single nucleolus, is usually found in the cen- tral or upper part of the embryo-sac near the egg, surrounded by a considerable mass of cytoplasm, but not separated from each other by cell walls (espn, fig. 3). Soon after this the walls of certain of these nuclei are seen to be flattened against each other (espn, figs. 3, 6), and a little later still the group may consist of one or two larger, elongated, constricted nuclei, each with two nucleoli, and six or four normal-sized nuclei each with a single nucleolus. These larger nuclei have been formed in each case by the fusion of two of the original nuclei of the group. The first one or two of these fusion nuclei form centers to which the other nuclei of the group draw up closely and fuse on to add their bulk to that of the larger nuclei. Finally a very large nucleus is formed, with at first several normal sized nucleoli, and later fewer very much larger ones or a single one (espn, figs. 6,7): The wall of this large nucleus shows at first several projecting lumps or knobs, each indicating the portion contributed by one 4 of the fusion nuclei (espn, figs. 4, 5,6, 7, 8—a series of sections of the same group). The line of contact of the walls of the fusing nuclei is at first evident by the darkly stained region where the two peripheral chromatin nets press against each other (espm,_ jigs. 5, 6,7). Later the lumps on the wall gradually smooth out and the chromatin net becomes evenly distributed about the periphery of the usually transversely elongated nucleus, which lies in a pretty dense mass of cytoplasm just below the oospore (espn, figs. 7, 8). From this time on this nucleus behaves like the endosperm nucleus of the typical angiosperm embryo-sac. It sometimes begins ‘its development before any activity is noticed in the ‘fertilized egg, except that the wall of the latter becomes more distinct and the sexual nuclei flatten against each other (fig. 72)+ In other cases the sexual nuclei fuse during the fusion of the endosperm-forming nuclei (fig. 7). The endosperm nucleus 1900 | *-ENDOSPERM AND-EMBRYO OF PEPEROMIA 5 divides by mitosis, the first spindle being approximately trans- verse. The number of chromosomes here is seemingly very large, compared with that in other spindles found in the embryo-sac or in the nucellus, but they were so densely packed in all the cases seen that it was impossible to make accurate counts. , : It is certain, however, that the amount of chromatin in this and its daughter spindles is greater than that found anywhere else in the plant (espn, figs. 9, rz). A cell plate is formed in the typical way at the middle of each spindle, the fibers of the latter being stretched out laterally to a surprising extent (espn, jig. 10). The new cell-wall thus formed stretches from the oospore to the base of the embryo-sac and cuts the latter com- pletely in two, forming thus two endosperm cells. Each of these divides further, forming a cell-wall immediately at each division, till in the oldest seeds seen there are forty or more endosperm cells, each with a large nucleus, several nucleoli, and dense cytoplasm, filling up all of the embryo-sac not occupied by the embryo and synergid and flattening the degenerating peripheral nuclei against the wall of the embryo-sac (esf, figs. 12, 13, 14, pn; figs. 9, 13). The fusion of the sexual nuclei is completed, at the latest, soon after that of the nuclei of the endosperm group, and before many endosperm cells are formed the oospore divides to form the embryo. In the few cases of the early divisions of the embryo seen the first wall seemed to be longitudinal, and the position of the walls in the slightly older embryos, often seen, seemed to confirm this (em, figs. 12, 13). The oldest fruits available, as was evident from their position on the spike were nearly ready to separate from the mother plant, z. €., were nearly ripe. In these the embryo consisted of more than twenty cells, but showed no sign of a definite suspensor and no indication of the organs of the young sporophyte. In fact, the whole structure has much the same shape and but slightly larger size than the one-celled oospore (asp, fig. 9, em, figs. 12,17). 6 BOTANICAL GAZETTE [ JULY ‘The ultimate fate of the long persistent synergid has not been made out with certainty as yet, but so far as it has been that it has at the time of fertilization, except that the wall becomes more distinct (sy, figs. 3,7, 9, 11, 72). In many cases where the embryo consisted of six or eight cells a single large cell could still be seen beside it, which seemed quite distinct — from the endosperm cells that press against the embryo on all — other sides, and this is interpreted as the still persistent synergid — (sy, fig. 12), and possibly the cell at the right of the embryo under the tapetal plug in fg. 73 is another case of the same sort. In several of the very oldest embryos seen there was a group of cells, smaller than any other cells in the embryo-sac, located in in this same position beside the embryo. The evidence obtained work, including probably the study of the sprouting seed, be necessary to make this certain and to determine the ultimate fate of these cells. The absolute size of the embryo-sac in these mature seeds is but little larger than when the egg is differentiated (figs. 2, 3 73—note the magnification of each). The relative size and position with reference to the other parts of the fruit is shown in fig. 14. The oblate spheroidal mass of endosperm is about one eighth the length of the whole seed. It is separated from the integument at the top by three or four layers of cells of the tapetum and nucellus. Below is the great mass of perisperm which lies the flattened and distorted, but still darkly staining nucleus ( psn, fig. 15). In these cytoplasm layers are also found large clear, or finely granular, spherical masses of an undetermined chemical nature, but presumably serving as food ( psp. fig. 15+) The single integument is but two cells in thickness, and both of these take part in the formation of the seed coat oF si) Sh a ee i aaa a a eer Bl ees ee eae bs ees ek 1900] ENDOSPERM AND EMBRYO OF PEPEROMIA 7 testa. All the walls of the cells of the outer layer become thickened till the cell cavity is practically obliterated, and the thickness of this layer is about the same over the whole of the seed, or slightly greater toward the base (int, figs. 14, 15). The cells of the inner layer thicken the outer walls greatly, especially near the upper end of the seed, where large knobs of the thickening substance project into the cell cavity (én, figs. 14,15). The cavity is never entirely filled as in the outer cells, but considerable space remains which is packed with starch like the cells of the perisperm (7, figs. 14, 15). The innermost layer of thickening substance of the outer walls of the cells of this layer is of quite different consistency from the rest of the wall, and shows in sections as a uniform border about all the hollows and projections of the latter. At the base of the seed several layers of cells of the chalaza thicken their walls, like those of the outer integument layer, to complete the protection of the seed (jig. 74). The seed does not escape from the carpel, but the latter apparently remains adhering closely to it when the whole falls from the mother plant. At this time the carpel is four or five layers thick, except at the base and in the stigmatic region (cf, fig. 15). The outer layer is of large, cuboidal, nearly empty cells, interspersed with knob-like hydathodes. Its cells have unthickened walls, except for the fine striae found quite generally on the outer epidermal walls of the whole inflorescence (¢, fig. 75). Next within this layer we find two or three layers of thin- walled flattened cells, with little contents. Closely adherent to the integument is the inner layer of the carpel, made up of large cells of about equal height and meridional length, but elongated equatorially to twice this length. These cells have the basal wall considerably thickened, with comparatively low ridges pro- jecting above this general thickening (fig. 75). The lateral and outer walls of these cells have anastomosing ribs surrounding thin spots or pits, forming cells closely resembling those of the velamen of the roots of many epiphytes in structure, and per- haps in function also. The basal or inner end of these cells is 8 BOTANICAL GAZETTE [JULY occupied by a granular mass, apparently of some firm substance deposited by the protoplast as an addition to the protective layers of the fruit and seed, or possibly connected with the absorption of water by these cells (fig. 75). The subtending bract increases but little in size after the macrospore is formed, and as the fruit ripens the bract withers and is squashed down by the swelling carpel (dr. figs. 1, 4). In comparing the foregoing with Campbell’s results it will be seen that my observations confirm his in regard to the origin. of the macrospore and its development to a sixteen-nucleate ripe embryo-sac. Campbell thinks that one of the upper of these nuclei goes to the egg, and one to each of the two naked syner- gids; while eight others, which he interprets as probably antipodals, temporarily collect at the base of the embryo-sac, but later disperse and become indistinguishable. The other five nuclei play no prominent part, there being according to his observations no nuclear fusion analogous to that of the polar nuclei of the ordinary angiosperm embryo-sac. In my own work I have seen but a single synergid which is long persistent and has a distinct wall. The nuclei of the group which Campbell interprets as possible antipodals I find are ultimately fused together into one endosperm nucleus, there being no special basal (antipodal) group of sterile cells or nuclei. Again, Campbell says that the at first flattened embryo finally : fills the whole embryo-sac and that there is no endosperm what- The meaning of these very striking peculiarities of the embryo-sac of Peperomia pellucida (and other species of the same genus) is not easy to determine. The extra division of the embryo-sac is quite unique, and so also is the lack of a basal group of sterile cells or antipodals, Finally the fusion of s0_ large a number of nuclei into one, in forming the endosperm nucleus, is approached only by the cases of fusion, at quite 2 1900] ENDOSPERM AND EMBRYO OF PEPEROMIA 9 different stage of development, of the several nuclei in the endo- sperm cells of Staphylea pinnata and Corydalis cava. In these forms, according to Strasburger (’80), when walls appear about the endosperm nuclei several of these are enclosed ina single cell and these later fuse to a single nucleus. The fusion of polar nuclei during instead of before fertilization is found also in Allium fistulosum (Strasburger ?79, p. 21), and this case may perhaps be considered as analogous to that of the endosperm- forming nuclei in Peperomia. That these peculiarities of Peperomia are to be considered primitive rather than higher specializations seems to me unwar- ranted by the evidence at present available, especially when we consider the fact, which I have ascertained, that such closely related genera as Piper, Heckeria, and Saururus have essentially typical angiosperm embryo- sacs. These latter forms develop a small amount of endosperm in a manner similar to that found in such distantly related and certainly not very primitive forms as the Nymphaeacee. Again, the lack of any grouping of the extra peripheral nuclei in the embryo-sac of Peperomia fails to give any encouragement from this source to those who look upon the antipodal group in the angiosperins as a second egg-apparatus (Lotsy, ’99, p. 106). So also the fusion of the eight nuclei to form the endosperm nucleus, if we regard it as at all homologous with that of the polar nuclei, seems to indicate that this is a purely vegetative or nutritive process, rather than anything like a sexual fusion as suggested by Mann (’92). Finally, the development of the cell walls in the endosperm directly after nuclear division each time, instead of by the method of free celle formation, as in the pro- thallus of the higher pteridophytes, is not favorable to the view that Peperomia is a transitional form between these forms and the typical angiosperms. I am inclined to believe that the peculiarities of the embryo- sac of Peperomia have been secondarily acquired, and are analo- gous to those found in other angiosperms of peculiar habit, e. g., many aquatic, parasitic, and saprophytic forms, Io BOTANICAL GAZETTE [JULY It is probable that a careful study of the sprouting seed will show the meaning of some of these peculiarities to the plant. I hope soon to be able to determine whether the tissue which I have called endosperm here has the same function as in Sauru- rus, of absorbing the perisperm for the benefit of the embryo during the sprouting of the seed. I trust also that a further study of related forms may discover some intermediate type of embryo-sac, that will indicate more definitely the possible deriva- tion of the peculiar one found in Peperomia. In conclusion and summary: the macrospore nucleus of; Peperomia pellucida forms sixteen free nuclei, of which one goes to the egg, one to the synergid, eight more fuse to form a single endosperm nucleus, while the other five remain sterile and degenerate. The nearly ripe seed contains an embryo of fifteen or more cells surrounded by endosperm cells in which the walls are formed directly from the cell plate of the spindle. JoHNs Hopkins UNIVERSITY, Baltimore. LIST OF WRITINGS REFERRED TO. CAMPBELL, D. H.: Die Entwickelung des Embryosackes von Peperomia pel- ucida Kunth. Berichte d. Deutschen botanischen Gesells. 17 : 452 . 1899: HOFMEISTER, W.: Neue Beitriige z. Kentniss d. Embryobildung d, Phanero- gamen. Abhandl. d. K. sachs. Gesellsch. d. Wissensch. 6: —. 1859- Lotsy, J. P.: Contributions to the life history of the genus Gnetum. Annales du Jard. Bot. de Buitenzorg II. 1: 46. 1899. MANN, G.: Development of the macrosporangium of J/yosurus minimus. Trans. Bot. Soc. Edinburgh. 1892. SCHMITz, F.: Die Bliithenentwicklung der Piperaceen. Hanstein Bot. Abhandl. 2: 1875. STRASBURGER, E.: Die Angiospermen und Die Gymnospermen. Jena. 1879- : Zellbildung und Zelltheilung, Dritte Auflage. Jena. 1880. EXPLANATION OF PLATE I. Abbreviations used: ax, axis of the inflorescence ; 47, subtending bract; cp, carpel; em, embryo; es, embryo-sac ; esf, endosperm; esfz, endosperm nucleus; ¢sv, principal vacuole of embryo-sac ; esw, wall of embryo-sac ; 74, integument; mf, micropyle; zc, nucellus ; 9, oosphere nucleus ; osf, oospore ; BOTANICAL GAZETTE, XXX PLATE I WV AX Ops \ ANG ~) ‘ ) re al RP = \ at NS lanmarce A l\y im epeaes me | YAY ‘ Sayre Ma STAR XOxNEH PILOT EAT CYT LTP i wey 1900 | ENDOSPERM AND EMBRYO OF PEPEROMIA a E gn, peripheral nucleus of the embryo-sac; fs, perisperm: fsfm, nucleus of perisperm cell; #7, pollen tube; sé, stigmatic lobe of carpel; sy, synergid ; tp, tapetal cells. All figures are camera drawings from microtome sections. Fic. 1. Longitudinal section of axis, bract, and carpel containing nearly ripe (sinteen nucleate) embryo-sac. X 150. F Longitudinal section of an ovule with a four-nucleate embryo- sac. X 440. F1G. 3. Longitudinal section of an embryo-sac after -the entrance of the pollen tube and male nucleus into the egg, and showing the group of nuclei that fuse to form the single endosperm nucleus (those with dotted outlines are in the next section to the one from which the rest of the figure is drawn). * 775+ Figs. 4-8. A series of sections of a group of nuclei fusing to form the endosperm nucleus, in an advanced stage of fusion and with large fused nucleoli; in fg. 7 the other contents of the embryo-sac are shown ; the fig- ures are numbered in the order of succession of the sections. X 775. Fig. 9. Longitudinal section of embryo-sac, showing the fusion of male and female nuclei in the egg, and a spindle of the second division of the endosperm nucleus; the upper nucleus of spindle from another section. x 775- Fic. 10. Part of a tangential section of embryo-sac, showing begin- ning of the formation of a cell-wall from the cell plate of the dividing endo- sperm nucleus. X 775. 1G. 11. Approximately transverse section of the upper end of an embryo-sac like that shown in fig. 9. X 775. Fic. 12. Longitudinal section of the upper end of an embryo-sac through the synergid and an eight-celled embryo. X 775. Fic. 13. Longitudinal section through the upper end of nucellus of a nearly ripe seed, containing an embryo of twenty or more cells surrounded by endosperm. 75. 1G. 14. Longitudinal section through axis, bract, and a nearly ripe fruit; endosperm and embryo practically as in fig. 13. X 75. Fic. 15. Part of section of carpel, integument, and perisperm from jig. 7g (at the left from the base of the embryo-sac). X 775. NEW OR UNRECORDED MOSSES OF NORTH AMERICA, J, J. CARDOT and I. THERIOT.* (WITH PLATES I1-v) PHASCUM CUSPIDATUM Schreb. var. Americanum Ren. & Card. var. nova.— Costa longe excurrente apice saepius decolora varietati piliferum proximum, sed _foliis brevioribus, me < magis papillosis pedicelloque brevissimo erecto distinctum Varietas mitraeforme Limpr., foliis papillosis similis, differt fol majoribus longioribusque, costa minus longe excurrente é calyptra conico-mitraeformi. ' Wisconsin: Madison, on ground in pastures, clover fields, and fallow ground (L. S. Cheney, 1893. Ren. & Card., Musci Amer. sept. ex Stl no. 267). Missouri: old fields near Emma (C. H. Demetrio, 1891). Illinois: — Microbryum Floerkeanum var. Henrict Ren. & Card. in Bot. GAZ. 14:91 9, from Kansas, leg. Henry, seems to be also a stunted form of the a moss. It has also the calyptra cucullate, a character which separates it from Microbryum Floerkeanum. All the specimens we have received from North America as Phascum cuspidatum belong to this var. 4 mericanum. 189 GyMNostomuM cuRVIROSTRE Hedw. var. COMMUTATUM Card & Thér. (Hymenostylum commutatum Mitt., Musci Ind. Or., p. 34 Weista curvirostris var. commutata Dicks., Handb. Brit. Mosses, 212). Newfoundland (Rev. A. C. Waghorne). © This variety has long, narrow leaves, everywhere long and smooth. In t of irregular cells papillae. and the cells of the areolation are he type, the upper areolation is composed » rectangular, quadrate, and triangular, with scatter GYMNOsTOMUM CURVIROS TRE Hedw. var. scaBprum Lindb. Musci Scand. 22, ‘ * Besides the new species, we shall eral species named by of w describe in this paper and the following se ; Renauld and Cardot in the Revue Bryologigque, 1892-1893, i hich only short, Provisional diagnoses were published. 12 1900 | MOSSES OF NORTH AMERICA 13 Missouri: Benton county, on moist rocks along Indian creek (C. H. Demetrio, 1893). Minnesota: Lewiston cave (J. M. Holzinger, 1889), Bear creek (J. M. Holzinger, 1890). Wisconsin: Madison (L. S. Cheney, 1892. Ren. & Card. Musci Am. seft. exsicc. no. 269). This form shows contrary variations to the preceding: the leaves are smaller, the cells quadrate, papillose; besides, the stem and the nerve are generally covered with high papillae. HyYMENOSTOMUM MICROSTOMUM R. Brown, Trans. Linn. Soc. 12:572 (Gymnostomum microstomum Hedw., Musc. frond. 3: 71, pl. 308). According to Lesquereux and James, Manual 56, this species is not known from North America, and all the specimens that have been communi- cated under the generic name Aymenostomum are to be referred to the Weisia viridula var. gymnostomoides C, Mill. Yet the no. 54 of Sullivant and Lesquereux J/uscz Bor. Am. belongs undoubtedly to the Hymenostomum microstomum R. Br., at least in our set. Quite inseparable from Wezsta viriduda as to the vegetative organs, and differing only by the capsule closed with a membrane finally perforated in the center Weista Wimmeriana BS., Bryol. Eu. 33-36: 4, pl. 7. (Gym- nostomum Wimmerianum Sendtn. in Flora 23':59. 1840. Ayme- nostomum murale Spr. Musc. Pyr. no. 236. Gymnostomum murale Sch. Syn. 37. 1860. [ Ed. 1.]) Minnesota: Taylor's Falls (J. M. Holzinger, 1895). Resembling the slender forms of W. viridu/a in size, habit, shape, and areolation of the leaves; distinct chiefly by the inflorescence, which is paroi- cous, or sometimes, as in our Minnesota specimens, synoicous. The peri- stome is generally rudimentary; in our American specimens, however, the teeth are rather perfect, with 4-5 articulations. DIcHODONTIUM Oxympicum Ren. & Card., Rev. Bryol. 19: 74. 1892.— Dioicum, humile, caespitosum, obscure viride. Caulis gracilis, erectus, 5—7™™ altus. Folia madida patentia subrecurva, sicca laxe erecto-flexuosa, apice incurvata, 1.50-1.75™™ longa, 0.6™™" lata, e basi ovata vel oblonga paulo latiore breviter lingu- lata, obtusa subobtusave, subundulata, marginibus plana, cellulis prominulis minute crenulata, supra basin integram distincte den- ticulata, costa valida percurrente vel subpercurrente; cellulae 14 BOTANICAL GAZETTE [JULY i minutae, obscurae, parietibus incrassatis, irregulariter quadratae, a 5-8 metientes, utraque pagina papillis prominentibus ornatae, — cellulae inferiores majores, elongate rectangulares, laeves. Folia perichaetialia longiora, basi laxius reticulata, superne papillis elongatis subcylindricis obsita. Capsula in pedicello pallido, 6-7" longo, siccitate dextrorsum torto erecta inclinatave, — oblonga, arcuatula, circa 1.5™" longa, 0.5™ crassa, collo dis- tincte strumoso, sicca plicatula et sub ore leniter constricta; operculum ignotum. Peristomium purpureum, elatum, dentibus triangulari-lanceolatis, 15—20-articulatis, usque ad medium in granulosis. Planta mascula ignota. Plate IJ. Washington: Olympic mountains (L. F. Henderson). A true miniature of D. pellucidum Sch., from which it is easily distin- — guished by the much smaller size, the leaves denticulate above the base, the cells of the areolation much smaller and with more prominent papillae and the strumose neck of the capsule. DICRANELLA LAXIRETIS Ren. & Card., Rev. Bryol. 20: 30. 1893.— Dioica, pusilla, gregaria. Caulis simplex, brevissimus, I-2™™" altus. Folia madida mollia, erecto-patentia, sicca flexu- osa subcrispata, ascendendo sensim majora, circa 2™™ longa, 0.25-0.3™™ lata, lineari-lanceolata, apice obtuso minute denticu- lato, marginibus integris planis vel parce reflexis, costa percur- rente vel sub summo apice evanida, rete laxo, cellulis inferioribus. majoribus elongate rectangulis, superioribus breviter rectangulis. subquadratisve, omnibus parietibus angustis. Folia perichaetia~ lia vix diversa. Capsula in pedicello tenui pallido, 5-6™™ longo, erecta, minuta, oblongo-subcylindrica, arcuatula, 0.75—1™™ longa, operculo convexo oblique longe subulato. Annulus e duplici vel triplici serie cellularum compositus. Peristomii dentes purpurei, . valde trabeculati, longitudinaliter striolati, usque ad % inferiora in 2 crura longa subulata bifidi. Planta mascula ignota. Plate TL. ‘, poe in a deep and shaded ravine near Lafayette (A. B. Langlois, 1891). One of the smallest species, resembling D. debilis L. & J., but with shorter stems, softer and more flexuous leaves, which are minutely denticulate Sipe“ oh sli er x aguas = lla le I el 4 ‘ F : : ® 1900 | MOSSES OF NORTH AMERICA 15 at apex, softer areolation of wider, shorter, and thin-walled cells, and narrower rather asymmetrical capsule. DicraANELLA Hower Ren. & Card., Rev. Bryol. 20: 30. 1893, et in Bull. de l’Herb. Boissier 4:15. 1896.— Dioica, caespitosa, subnitens, lutescenti-viridis. Caulis erectus, simplex vel parcis- sime divisus, 6-10" altus. Folia laxiuscula sicca erecto- flexuosa, madida plerumque plus minus subsecunda, I.5—2.2™™ longa, 0.28-0.35™™ basi lata, lanceolato-subulata, acuta subacu- tave, integerrima vel summo apice obsoletissime denticulata, marginibus ubique planis, costa lata, 4—™% basis et totam fere subulam occupante, cellulis angustis, linearibus, inferioribus bre- vioribus. Folia perichaetialia e basi subvaginante longius et tenuius subulata. Capsula in pedicello rubello, siccitate sini- strorsum torto, 5-7™" longo, subhorizontalis, oblonga, arcuata, circa 1™ longa et 0.28-0.35™™ crassa, sicca sub ore valde con- stricta, operculo alte conico. Annulus nullus. Peristomium D. variae.—FPlate Il. California: Mt. Tamalpais, Marin county, on wet banks (Marshall A. Howe, 1892-1893. Ren. & Card., Musci Amer. sept. exsicc. no 203) cies or a regional race; characterized by the more distant and more flexuous leaves, subsecund when moist, generally longer, plane on the borders, the broader nerve, the narrower cells, the somewhat narrower capsule in dry state, and the green-yellowish and brighter tinge of the tufts. DICRANUM VIRIDE BS. var. laeve Ren. & Card., var. nova.— A forma typica habitu multo laxiore ceca minus congestis dorso laevibus distincta. Newfoundland: Bay-of-islands, old stump (Rev. A. C. Waghorne, 1895). Dicranum ancustuM Lindb., Soc. pro Fauna et FI. fenn. 1880, et Rev. Bryol. 9:83. 1882. Lindb. & Arn., Musc. As. Bor. 2:80 (descriptio locupletissima). Northwest shore of Hudson Bay, lat. N. 63°55’, long. O. go0°20’ (G. Comes, 1893-1894). We found some stems of this rare species amongst specimens of Audacomnium turgidum. r moss, known only from some localities of north Finland and from Siberia. It is easily distinguished from D. Bonjeani De Not. (D. 16 BOTANICAL GAZETTE [yuLy ; palustre BS.) by the leaves straight, not undulate, convolute, and entire, the j ! thinner costa, the less porose cells, and the perichaetial leaves long piliferous. FIsSIDENS BRYOIDES Hedw., var GYMNANDRUS Ruthe, Hedwi- — gia 9:178. 1870. Limpr. Laubm. 1: 430. (F. gymnandrus Buse | Musc. neerl. exsicc. fasc. 4, no. 77). Northwestern Montana: in the vicinity of Lake MacDonald, Flathead county (J. M. Holzinger and J. B. Blake, 1898). A peculiar form, easily distinguished from the type by the antheridia naked in the axils of the stem-leaves. FISSIDENS SUBBASILARIS Hedw. var. Bushii Card. & Thér., var. nova.— A forma typica differt foliis latioribus brevioribusque, obtuso rotundatis, nervo fere ad apicem producto et areolatione magis opaca cellulis parietibus crassioribus. . Missouri: Eagle rock, on gravelly ground (B. F. Bush, 1897). Desmatodon systilioides Ren. & Card., sp. nova.— Monoicus, gregarius. Caulis brevis, erectus, 2-3™™ altus. Folia in rosu- lam congesta, patula, oblongo-lanceolata, apice sat subito brevi- terque acuminata, acuta, marginibus planis superne inaequaliter denticulatis, nervo valido rufescente percurrente vel breviter excedente, cellulis inferioribus laxissimis, subrectangulis, inani-_ bus, hyalinis vel lutescentibus, mediis et superioribus minutis, - rotundato-subquadratis vel subhexagonis, papillis numerosis _ obscuratis, marginalibus 2~—4-seriatis, quadratis vel breviter rec- tangulis, vix vel parum papillosis, limbum distinctum lutescen-— tem translucentem efformantibus. Folia perichaetialia vix. diversa, paulo breviora. Capsula in pedicello lutescente vel pallide rubente, siccitate dextrorsum torto, 8—12™ longo erecta, ee ee eR oe EE PRE SOR ee ee, PERRET ee ee See, S 1900 | MOSSES OF NORTH AMERICA 17 shortly excurrent, and by the total lack of peristome; at least, all the cap- sules we have been able to examine do not show the slightest trace of this organ. The pellucid border of the leaves shows some relationship between D. systylioides Ren. & Card. and D. Portert James, but the latter has a much narrower capsule with a highly conic lid not adhering to the columella, and the peristome and annulus are well developed. Barbula eustegia Card. & Thér., sp. nova.— Dioica? gregarie caespitosa. Caulis brevissimus, 1-2™" altus. Folia siccitate erecto-flexuosa, madida recurvo-patula, lineari-lanceolata, acuta, subacuta obtusiusculave, superne plicato-canaliculata, inferiora minima, 0.5™" longa, superne sensim majora, superiora I™ longa, marginibus planis vel parce revolutis, integris, costa sub apice evanida vel eum fere attingente, dorso papillosa, cellulis inferioribus pellucidis, laxiusculis, laevibus, oblongis, subrectan- gularibus, sequentibus quadratis, superioribus parvis, vix 4-5# metientibus, quadrato-rotundatis, obscuris, minute papillosis. Folia perichaetialia caulinis majora, externa e basi semivaginante sat subito in acumen elongatum canaliculatum, patulo-arcuatum constricta, interna oblonga, late breviterque acuminata, rete omnino pellucido. Capsula in pedicello capillari pallido, 12-18™ longo, siccitate dextrorsum torto, erecta vel obliqua, oblonga, I-1.2™ longa, 0.3—0.4™™ crassa, operculo conico-subulato capsu- lam aequante vel superante. Annulus duplex, 0.07™ latus. Peristomium purpureum, membrana basilari 0.08™™ alta, dentibus circa 1™ longis bis convolutis, valde granulosis. Sporae laeves, 84 crassae. Flores masculi ignoti. Verisimiliter dioica—Plate Idaho: Cedar creek, Latah county, on ground (L. F. Henderson, 1897). This moss, received from Mr. Henderson only in very small quantity, seems a miniature of B. favifes BS., from which, besides in its small size, it differs chiefly by the lid as long as the capsule or even longer. By this char- acter, as well as by the form of the leaves, it resembles also 7richostomum dicranoides Sch. (T. macrostegium Sull. Icon. Suppl. 35, £7. 22) from Central and South America and the Antilles, which has been also recorded from Ala- bama ; but this last species has the beak of the lid thinner, the leaves larger, broader, denticulate above, a less opaque areolation of larger and more dis- tinct cells, and the peristome less twisted, with a shorter basilar membrane. 18 BOTANICAL GAZETTE tome ramosus, circa 1™altus. Folia sicca et madida erecta, subimbricata 1.75-3™" longa, 0.75-1 lata, ovato-lanceolata, marginibus planis integerrimis, inferiora mutica, superiora pile breviusculo minute denticulato, basi paulo decurrente instruc costa sat valida, basi 60-80 lata, superne canaliculata, apice versus indistincta, cellulis inferioribus juxta costam linearibt hyalinis, caeteris parvis, obscuris, quadratis vel subrotundatis, bistratosis. Folia perichaetialia subsimilia, basi laxius reticulata, Calyptra cucullata. Capsula in pedicello crassiusculo stricto, 1.5>2.5™™" longo, exserta, erecta, breviter oblonga, sicca subcy- lindrica, exannulata, operculo convexo-rostrato. dentes flammei, circiter o. 35™™ alti, integri vel parcissime perfo- rati, e basi late triangulari longe subulati, superne minute granu- losi, articulis 15—25.—Plate IV. — Idaho: near Moscow, on dry rocks (L. F. Henderson, 1894). Closely allied to G. montana BS., but sufficiently distinct by the larger leaves, with a stouter nerve and a shorter and a thicker hair, and chiefly by the peristomial teeth almost entire, not divided and scarcely perforated, W more numerous articulations. GRIMMIA Montana BS. var. Idahensis Ren. & Card., var. nova —A forma typica differt capsula pro more majore magisq ‘ exserta et pedicello paulo longiore, siccitate plerumque flexuos subgeniculato. Folia pilo saepe destituta. North Idaho: west end of Lake Pend d'Oreille (J. B. Leiberg, 199% Ren. & Card., Musci Amer. sept. exsicc. no. 289). GRIMMIA SUBSULCATA Limpr., Laubm. 1: FE). Idaho (J. B. Leiberg, 1889; J. H. Sandberg, 1892). Northwestern M tana: in the vicinity of Lake MacDonald, Flathead county (J. M. Holz and J. B. Blake, 1898). A long time confused with G. alpestris Schleich. The distinctive acters quoted by Limpricht are: for G. alpestris, pedicel straight, caps™ without stomata, leaves not plicate; and for G. subsulcata, pedicel somew curved, capsule with stomata, leaves with two longitudinal folds in the Up 1900] MOSSES OF NORTH AMERICA 19 part. The last character is the best, for the pedicel of G. sudsulcata is some- times néarly straight and the capsule without stomata, while the folds of the leaves are always distinct, especially on a transverse section. It is to be noticed that Limpricht cites erroneously G. /ame//osa C. Miill. as a synonym for G. a/pfestris: on the contrary, from an original specimen Miiller’s plant is proved identical with G. sudsu/cata. The true G. a/festris Schleich. has been gathered by Messrs. J. M. Holzin- ger and J. B. Blake in the same region of northwestern Montana where they have collected G. subsulcata, Orthotrichum Idahense Card. & Thér., sp. nova.— Monoicum, laxe depresso-pulvinatum, inferne fuscum, superne lutescenti- viride. Caulis basi decumbens, longe denudatus, irregulariter ramosus, 2—3™ longus, ramis ascendentibus. Folia madida erecto-patentia, sicca erecto-appressa, 1.75—2.50™™ longa, 0.50- 0.75 lata, oblongo-lanceolata, subobtusa, integra, marginibus fere e basi usque infra apicem arcte lateque revolutis, costa sub- percurrente; cellulae ubique unistratosae, parietibus valde incrassatis, inferiores rectangulares, juxta costam lineares, mar- gines versus quadratae vel breviter oblongae, caeterae rotundae, utraque pagina papillis grossis bi-trifurcatis obsitae. Folia peri- chaetialia subconformia, basi laxius reticulata. Vaginula nuda. Capsula in pedicello brevissimo vix emersa vel semi-emersa, madida ovata, collo brevi attenuata, sicca subcylindrica, infra os leniter constricta, octostriata, striis e cellulis 4-seriatis, longiori- bus, lutescentibus, parietibus crassioribus compositis, stomatibus emersis. Operculum ignotum. Peristomii dentes 8 bigeminati, vel 16, siccitate reflexi, plus minus pertusi, minute granulosi, superne lineolati; cilia nulla vel fugacia. Calyptra lutescens, apice fusca, ramentis longis, numerosis, denticulatis, papillosis obsita. Sporae papillosae, 20-22 crassae. Flores masculi ses- siles.— Plate V. ae Moscow mountains, on rocks (L. F. Henderson, 1893). the superficial stomata and the peristome reflexed when dry and ea Sepaica: this species belongs to the group of O, arcticum Sch., but is easily distinguished from all the other species of this group by its lax tufts, emergent capsule, and peristomial teeth less opaque, covered with less dense papillae. x i 20 BOTANICAL GAZETTE [JuLy — nova.— A forma typica differt capsula exserta in pedicello eam 3 aequante foliisque usque medium versus revolutis. Ab QO. papil- acumine breviore et minus angusto distincta. Calyptra valde pilosa. Papillae foliorum parum prominentes. Folia nonnullis propagulis saepe instructa. California: region of the upper Sacramento, Sisson, on trunks of Quercus Kelloggii (Marshall A. Howe, 1894. Ren. & Card. Musci Amer. sept. exsttt. no. 291). WEBERA CaRINATA Limpr., Laubm. 2: 261. (Bryum carinatum Boul., Musc. de la France 280. B. naviculare Card., Rev. Bryol. — 13:27. 1886, et B. cymbuliforme Card., loc. cit. 14:22. 188700 Webera cucullata var. carinata Husn., Muscol. Gall. 229). Northwestern Montana: in the vicinity of Lake MacDonald, Flathead county (J. M. Holzinger and J. B. Blake, 1898). By the habit and the dioicous inflorescence, this moss approaches W. com- rows, and the cell walls thinner. Bryum euryloma Card. & Thér., sp. nova.— Dioicum, dense caespitosum, lurido-viride. Caulis erectus, tomentosus, 2-33 altus. Folia conferta, madida erecto-patentia, sicca subappressa, € basi decurrente anguste lanceolata, 3-4™" longa, 0.65-0.75"™ lata, sensim et tenuiter acuminata, integra vel apice obsoletis- sime denticulata, marginibus. anguste revoluta, interdum uno latere subplana, nervo in cuspidem breviusculam, acutissimam, Folia perichaetialia intima sim cuspidata, margine plana. —20™™" longo basi atropurpureo, abrupte lindrica, sicca sub ore constricta, collo ello sensim defluente instructa, 2.50-4 longa, 0.75~1 crassa, operculo convexo apiculato. Peristomium B. pseudotriquetri. Flores masculi capituliformes.— Plate V. 1900] MOSSES OF NORTH AMERICA 21 Puget sound, Orcas island, Mt. Constitution, lake border (L. F. Hender- son, 1892). Distinct from &. pseudotriguetrum Schw. and allied species by the smaller size, the narrower leaves entire or scarcely denticulate at apex with broader margin, and the capsule smaller, narrower, and more abruptly pen- BRYUM CRASSIRAMEUM Ren. & Card. var. Covillei Ren. & Card., var. nova.— A forma typica differt cespitibus densioribus, caulibus ramisque gracilioribus, strictioribus, foliis strictis magis appressis, costa pro folii magnitudine plerumque crassiore, cap- sula fusco-rubra et peristomii interni segmentis dorso latius apertis. Rocky mountains (Death valley Expedition, no. 1358; F. V. Coville and F, Funston, 1891). BryuUM TORQUESCENS BS., Bryol. Eur. 6-9: 49, p/. 20. Washington: Pullman, Whitman county, moist banks (L. F. Henderson, 1892). Nearly allied to B. capil/are L., but distinct by the synoicous inflorescence and the capsule deep red when mature. The American form differs from the European type by the leaves being erecto-patent and not spirally contorted in dry state. PTEROGONIUM GRACILE Sw. var. Californicum Ren. & Card., var. nova.— A forma typica Europaea differt foliis longioribus longiusque acuminatis cellulisque alaribus minoribus. California: ‘‘ad rupes Californiae, perfrequens; Bolander”’ (Sulliv. et Lesq. Musci bor. amer. exsicc., ed. 2, no. 349); Sansalito (Marshall A. Howe, nian specimens of P. gracile that we have examined belong to this variety. PYLAISIA POLYANTHA Sch. var. drepanioides Ren. & Card., var. nova.— Forma peculiaris, habitu et magnitudine Hypno palles- centi similis; folia secunda plerumque ad basin acuminis obsolete denticulata, cellulis alaribus minus numerosis et minus obscuris, rete magis scarioso. Capsula minor. Peristomium normale. Minnesota: without locality or name of collector, mixed with a small form of Hypnum uncinatum Hedw. (Herb. Univ. of Wisconsin). 22 BOTANICAL GAZETTE [JULY PSEUDOLESKEA PATENS Limpr. Laubm. 2:806. (Leskea? patens Lindb. in Soc. pro Fauna et Fl. fenn. 1880. Lesguereumia— patens Lindb. in Meddel. af Soc. pro Fauna et FI. fenn. 14:75 1887. Newfoundland: Deer lake (Rev. A. C. Waghorne). his species differs from P. afrovirens in its more slender stems, the leaves erecto-patent (not secund), symmetric (not falcate), and the papillae being set on the middle of the cells and not on the angles. TRIPTEROCLADIUM LEUCOCLADULUM (C. Miill.) Jaeg. var camptocarpum Card. & Thér., var nova.— A forma typica differt tantum capsula brevi, subhorizontali, arcuata, brachythecioidea. | Idaho: Latah county (L. F, Henderson, 1894). AMBLYSTEGIUM SERPENS Br. Eur. var. subenerve Ren. & Card., var. nova.— A caeteris formis minoribus A. serpentis differt foliis enervibus vel subenervibus. Ab A. sudtili habitu robustiore, foli multo majoribus latioribusque, brevius accuminatis distinctum Newfoundland: Bay-of-islands (Rev. A. C. Waghorne). AMBLYSTEGIUM FLUVIATILE Br. Eur. var. brevifolium Ren. & Card., var. nova.—A forma typica Europaea caule magis regula riter pinnato foliisque minoribus, brevioribus, ovato-acuminatis costa pro folii magnitudine crassiore distinctum. Minnesota: Lanesboro (J. M. Holzinger, 1894. Ren. and Card. M/uset Amer, sept. exsicc. no. 327). AMBLYSTEGIUM RIPARIUM Br. Eur. var. longinerve Card. Thér., var. nova.— A forma typica nervo in acumen longius pro- ducto distinctum. Arkansas: Varner, in water (B. F. Bush, 1898). Resembles A. vacillans Sull. in the long-nerved leaves, but in this sp& cies the branch leaves have a short obtuse acumen, while in our moss they are narrowly and acutely acuminate, like the stem leaves. Hypnum Hatter Linn. fil. apud Swartz Meth. Musc. 34 Labrador: |’Anse-au-Mort (Waghorne, 1894); Cook's brook (Waghorné 1897). Newfoundland: Middle Arm, on rocks (Waghorne, 1896). Let aay ee Pen Pee ae ge ee na . ; : ; 1900] MOSSES OF NORTH AMERICA 23 A very distinct species of the subgenus Campy/ium, at once characterized by the very dense tufts, the stems entirely prostrate and divided into pinnate branches, the leaves much crowded, recurved-squarrose from a more erect base, minutely denticulate all around, and with a much shorter point than in the allied species. HypnuM CUPRESSIFORME L, var. RESUPINATUM Sch. Coroll. 133. (HZ. resupinatum Wils., Bryol. Brit. 398). Newfoundland: Chance cove (Rev. A. C. Waghorne, 1891). This variety, considered by many authors as a distinct species, is charac- terized by the leaves not falcate-secund, imbricate or homomallous and pointing upward, and the capsule erect and symmetrical or very slightly curved or inclined. It is connected with the type by intermediate forms. Hypnum MOLLE Dicks. var. SCHIMPERIANUM Sch., Syn. 775 [ed. 2]. (4. Schimperianum Lorentz, Moost. 123, pi. 5, fig. €). Northwestern Montana: in the vicinity of Lake MacDonald, Flathead county (J. M. Holzinger and J. B. Blake, 1898). Differs from the type by the longer and more slender stems, naked below, and by the leaves smaller and with a shorter acumen. STENAY and Le HAvreE, FRANCE. EXPLANATION OF PLATES II-V. Nachet’s objectives 3 and 6, oculars 1 and 3, with camera lucida, All drawings are reduced ¥ in photo-engraving. PLATE IIl.—1. Dichodontium Olympicum. a, entire plant, nat. size; 6, 6, b, leaves X 32; c, basal areolation X 135; 4, marginal areolation in the upper part X 260; e¢, transverse section of the nerve X 260; f, capsule ripe and deoperculate x 26; g, a tooth of the peristome X 105; h, part of the same in the upper part X 285.—2. Desmatodon systylioides. a, entire plant, nat. size; 4, 4, 6, leaves X 32; c, marginal areolation in the middle X 135; d, areolation of the upper part X 135; é, capsule ripe X 32; J, capsule unripe X 32. PLATE II].—1. Dicranella Howei. a, entire plant, nat. size; b, 6, 6, leaves X 32; c, apex of a leaf X 135; d, basal areolation X 135; 4% marginal areo- lation in the middle of a leaf X 135; 4, capsule in moist state X 32; g, Cap- sule ripe and deoperculate, in dry state X 32.— 2. Dicranella laxiretis. a, entire plant, nat. size; 4, 6, leaves X 32; ¢, basal areolation X 135; d, mar- ginal areolation in the middle X 135; ¢, areolation of the upper part X 135; f, capsule X 32; g, portion of the annulus X 135; 4,a tooth of the peristome * 135 24 BOTANICAL GAZETTE [JULY PLATE IV.—1. Barbula eustegia. a, entire plant, nat. size; 4, the same X 3; ¢,¢, ¢,¢, leaves X 32; d, basal areolation X 135; ¢, areolation of the upper part X 135; /, external perichetial leaf x 32; £, inner perichetial leaf x 32; h, capsule X 32; 7, peristome X 60; 7, portion of a peristomial tooth x 285. —2. Grimmia pseudo-montana. a, entire plant, nat. size; 4, 6, leaves X 32; ¢, basal areolation x 135; d@, areolation of the upper part X 135; e, transverse section of the leaf in the lower part X 135; 4% transverse section of a leaf in the upper part X 135; g, capsule ripe X 26; 4, two teeth of the peristome Xx 135. PLATE V.—1. Orthotrichum Idahense. a, entire plant, nat. size; 4, 4, leaves X 32; c, basal areolation x 135; d, areolation in the upper part X 135; é, transverse section of a leaf X 135; 7 capsule'ripe and deoperculate, in dry State X 26; g, areolation of the capsular membrane in the upper part X 135; h, the same, in the lower part, showing a stoma X 135; z, two teeth of the peristome X 135; 7, young calyptra X 26.—2. Bryum euryloma. a, entire plant, nat. size; 4, leaf x 26; ¢, marginal areolation in the middle of a leaf %* 135; @, areolation of the upper part X 135; ¢, Capsule in dry state x I5. PLATE If —| s 7 ——— 5 SS SS | ESS Sa SS ——— —Sj = — | =—— Gs = = {=— = LoL) | Se on a Ss= Site AQ0000 0a an00% oc CARDOT & THERIOT on MOSSES 000 SSS. QO0 COE ce a0 Clejy O, 990% 4 e BOTANICAL GAZETTE, XXX Jandot deb. ei ine Na aoa ed NT See ee ee EE CT a ee ee eee ee ae eee ee aE Se ee Te ee BOTANICAL GAZETTE, XXX PLATE. tif = — = —— SS = —— aang = ————— tt | —— = aoe ——- a ————— ————— on MOSSES PLATE IV BOTANICAL GAZETTE, XXX CARDOT & THERIOT on MOSSES THE DEVELOPMENT OF THE EMBRYO-SAC IN SOME MONOCOTYLEDONOUS PLANTS. a KARL M. WIEGAND. (WITH PLATES VI AND VII) THE material for the following study was prepared in the ordinary way, by fixing in the chrom-aceto-osmic acid solution, imbedding in paraffin, and staining with the gentian-violet-orange | combination. Although the development in Canna was found to be nearly normal, that in each of the other two plants showed some very interesting and important variations. Whether these throw any light on the problem of the homology of the embryo- sac can be determined only by more extended study of other plants. Convallaria majalis L. THE HYPODERMAL CELL AND ARCHESPORIUM. The embryo-sac of Convallaria is derived from a hypodermal cell situated at the apex of the nucellus. This hypodermal cell is first discernible as an enlarged oblong or more or less distinctly triangular cell at the apex of the nucellus and directly under- neath the epidermis; but can also be distinguished from the adjacent cells by its larger size and more granular contents. Very early in its development a single cell, the so-called “tape- tum,” is cut off on the side adjacent to the epidermis. This immediately divides by an anticlinal wall into two daughter cells which lie side by side at the summit of the embryo-sac (jig. 7). The nucellus is comparatively broad, and the growth of the archesporium taking place subsequent to the separation of the wall-cell is to a large extent ina lateral direction. To accommodate themselves to this the two daughter wall-cells undergo repeated anticlinal division, so that eight or ten cells are formed, all arranged in the same plane, and forming a plate of tissue just beneath the epidermis (fig. 2). The nuclei of 1900] 25 26 BOTANICAL GAZETTE (juLy these cells increase somewhat in size until they are noticeably a larger than those of the surrounding tissue, but unlike the sporogenous nuclei remain very dense. As the embryo-sac 4 grows larger these cells are all pushed aside, with the exception — of the more central ones which persist between the embryo-sac and the epidermis until after fertilization. : The other daughter cell resulting from the division of the primary hypodermal cell constitutes the archesporium. This E immediately expands in all directions, partly at the expense of the ordinary tissue. At the time of the first nuclear division, the cell is oblong in shape and occupies a considerable portion 4 of the nucellus. The nucleus during its period of growth passes through stages almost identical with those described for the — microsporangial archesporium.t The chromatin network changes _ during synapsis (fig. 2) to a spirem ribbon (fig. 3), which later segments into the individual chromosomes. Corresponding — Stages in the nucleus of the pollen-mother cells and embryo-sa¢ archesporium cannot be distinguished structurally; and this similarity is still farther emphasized by the synapsis occurring in each at the same stage in development. af lh Si THE FIRST NUCLEAR DIVISION. Several good preparations of the first nuclear division we obtained, both in the nuclear plate and anaphase stages. In th plate stage the fibers are well marked; indeed the fascicles attached to the chromosomes are especially large and prominent The spindle, like that of the pollen-mother cell, is rarely po at the poles, but is more often truncate (fig. 4 The chromosomes are large oblong bodies arranged on the nuclear plate just as they are in the pollen-mother cell ; that horizontally with one end directed away from the axis. Simul taneously the outer and inner ends commence to split longitu dinally, but in perpendicular planes. In one case the page is ors a wholly without the aid of the spindle fibers; * WIEGAND: The development of the microsporangium and microspores in Com vallaria and kaa Bot. GAZ, 28: 328. 1899. 1900] DEVELOPMENT OF THE EMBRYO-SAC 27 the other the fibers seem to accomplish the separation. The former splitting is only partial, while the latter finally divides the chromosome into two V-shaped parts which move at once to the poles through the influence of the spindle fibers. This is therefore a heterotypic division, and similar to the one occurring in the pollen-mother cell. In fact it seems probable that having the spindle alone one could not distinguish the two cases, even after a careful study of the chromosomes themselves. The num- ber of chromosomes was found to be eighteen, and since the number counted in the somatic cells was greater than thirty, it is evident that reduction takes place at this period. During the anaphase of the first division (fig. 5) a definite cell-wall is deposited, which divides the original cell into two nearly equal parts. THE SECOND NUCLEAR DIVISION. The nucleus in each of the two daughter cells resulting from the first division very quickly divides again, and it so happens that the two spindles are formed simultaneously. No cell-walls are formed after this division, at least not before the embryo-sac is nearly mature. At the stage shown in fig. 8 four nuclei are present. The second division spindle was in this case directed longitudinally; therefore in the same plane as the first; but this is not always the case. Sometimes the axis of the spindle is inclined and in fact almost. transverse. The position of the spindle, however, seems to be of little importance, since one may find the two daughter nuclei during later stages either one above the other or side by side. The resting stage between the first and second divisions, like that in the pollen-mother cell, is very short. The V-shaped chromosomes remain distinct and uninclosed by a definite mem- brane. The spindle quickly forms, apparently from the sur- rounding cytoplasm, and at the same time the chromosomes are crowded toward the equator where they arrange themselves in a nuclear plate. These spindles were in all cases much less dis- tinct than the first one, and appeared in the earlier stages 28 BOTANICAL GAZETTE [JULY shorter and more truncate, while in the anaphase no cell-plate whatever was produced. The way in which the chromosomes divide could not be posi- tively determined. The segments on entering the nuclear plate, 7 and even while in the plate as seen from the poles, are much curved (fig. 6). A side view of the nuclear plate shows a dense mass with projecting arms similar to the corresponding figure in the pollen (fig. 6). When the segments move to the poles they are seen to be quite long and straight, and never V-shaped as in the heterotypic division (fig. 7). After reaching the pole they at length fuse, and a membrane is formed around them, thus bringing about a truly resting condition again. It seems scarcely to be doubted that the process here is identical with the second division in the pollen-mother cell. Indeed, every appearance in the one has its almost exact coun- terpart in the other. GROWTH AND DEVELOPMENT OF THE EMBRYO-SAC. The further history of the embryo-sac. is very interesting, — since it shows some deviations from the ordinary process in both monocotyledons and dicotyledons. The two superimposed a daughter cells should probably be considered as constituting the : so-called ‘axial row” in this case (figs. 6-8). At least, so far : as the nuclei are concerned, they are the equivalents of the two : cells first formed in Canna. There is therefore a two-celled axial row instead of a four-celled one, as in Canna and many — other plants. It would be expected then that during develop- — ment the lower cell alone would become the embryo-sac, while | the upper would undergo dissolution as in nearly all other cases. This, however, is not the case. At the time when the spindles of the second division occuf, the archesporium as described above is two-celled. No walls 4 are produced by the second spindles, as a result of which each . cell now contains two nuclei. This stage is followed by a com : paratively long period of growth in which both cells increas€ — several times in size, as do also their nuclei, A large number of 1900] DEVELOPMENT OF THE EMBRYO-SAC 29 preparations were obtained illustrating this condition and repre- senting all stages up to the next division of the nuclei. As shown in the accompanying figure (jig. 8), the upper cell becomes gradually larger and more vacudlate, while the cyto- plasm of the lower stains more deeply and fills nearly the entire cell cavity. No cases were observed, however, where one could infer that either cell was in the process of disintegration. At a period shortly before the opening of the flower further changes occur, the first of which is the immense increase in size of the upper cell. The wall between the two still seems to remain intact, although becoming very thin and delicate. In some cases the wall is so delicate at this stage as to be almost invisible, and may have broken down entirely, in which case both cells would be merged into the general cavity of the embryo-sac, but in later stages two cells are again seen, All of the four nuclei undergo division simultaneously, resulting in the stage with eight nuclei, just as in Lilium (fig. 9). In several cases shortly after this two large nuclei were seen fusing near the lower end of the upper cell, while the lower cell contained the distorted remains of three other nuclei. The latter cell was thus apparently already in the process of disintegration. It seems probable that one of the four nuclei originally formed in the lower cell must have ruptured the thin cell-wall and fused with the upper polar nucleus after the normal manner. In fig. ro is represented an embryo-sac just prior to fertiliza- tion. At this stage the ‘‘ egg-apparatus ”’ consists of two syner- gids and the egg. The former lie close together, with the long axis more or less transverse to the axis of the embryo-sac, and are elliptical in shape, with the wall at the upper end thickened and densely striate. Just below these is the large egg nucleus, separated from the synergids and from the main cavity by very delicate cell membranes which seem to extend completely across the embryo-sac so as to join the lateral walls on either side. Below the egg-apparatus is the very large main cavity of the embryo-sac lined with a thin layer of cytoplasm. In this, near the base, is the large definitive nucleus still showing signs of the 30 BOTANICAL GAZETTE [ory previous fusion of the two polar nuclei. At the base of the 4 embryo-sac are the distorted remains of three antipodal nuclei 4 still separated from the main cavity by a distinct cell-wall. It was impossible to determine whether this had been formed — anew, or whether the old perforated wall had simply been g repaired. - At the time when the main features of this study of Conval- j laria were read at the Boston meeting of the American Associa- tion for the Advancement of Science, so far as the writer was aware no exactly similar case had been observed. Mann? had discovered a cross wall in the embryo-sac of Myosurus, but here the development seems to have been somewhat different ; and Strasburger3 had figured a structure somewhat resembling 4 cross-wall in the embryo-sac of Allium, although it was men- tioned in the text as a plasma-plate. Since that time, however, McKenney* in making a careful study of the embryo-sac of Scilla has described a process very similar to that in Convallaria. The archesporial nucleus underwent a period of growth before dividing. The first division showed the reduced number of chromosomes and was followed by a cell-wall. The two suc- ceeding divisions in each daughter cell were not followed by @ cell-wall, so that as a result two cells were present, each con- taining four nuclei. Up to this point the process was similar to Convallaria, but it was further found that only the upper cell took part in the formation of the embryo-sac, and not both in Convallaria, while the lower cell disintegrated. The process in Convallaria is strikingly similar to that im Lilium, and is probably to be considered as simply a modifi-. cation of that method or a transition to it from the ordinary type with two or four cells in the axial row. It differs merely in the possession of a cross-wall, which however partially or *MANN: The embryo-sac of Myosurus minimus L. Trans. and Proc. Bot. Soe Edinburgh 29: 351. 1892. *STRASBURGER: Die Angiospermen und die Gymnospermen (1879). 7 be Jigs. 81-86. * MCKENNEY: Observations on the development of some embryo-sacs. Contrib. Bot. Lab. Univ. Pennsylvania 2: [no. 1] 80. 1898. 1900 | DEVELOPMENT OF THE EMBRYO-SAC 31 entirely breaks down before the fusion of the polar nuclei, one of. which comes from each cell. Potamogeton foliosus Raf. THE HYPODERMAL CELL AND ARCHESPORIUM. In the earliest stage obtained the hypodermal cell at the apex of the nucellus had commenced its period of growth. No definite form can be ascribed to it at this period, although it is perhaps more often wedge-shaped. Even from the beginning it usually contains more protoplasm than the other cells, a feature which at a later stage becomes still more noticeable. After a short period of growth this cell divides. The inner daughter cell resulting from the division becomes immediately the archesporial cell (fig. 14). The outer daughter cell divides again by an anticlinal wall. Two cells now lie side by side above the archesporium, as is represented in fig. 12, which shows a cross-section through the apex of the nucellus. fig. 13, also a cross-section of the nucellus taken from the same inflo- rescence as fig. 12, shows that each of two cells now divides again, so that four daughter cells are formed all in the same plane. This last division may take place either before or. after the first periclinal division of the wall-cell, commonly however before. Periclinal divisions in all four cells now begin, so that at length four rows of cells are formed between the arche- sporium and the epidermis. The process may continue until as many as six layers are produced ; and these all persist until the embryo-sac reaches maturity, although often in a more or less compressed condition. In the anther the hypodermal celi divides into two parts, one being destined to produce the archesporium, while the other after two or three periclinal divisions constitutes, together with the epidermis, the wall of the anther. The stages leading up to the production of the embryo-sac in the ovule are in many respects very similar to those occurring in the young anther. The hypodermal cell here also divides by a peri- clinal wall into two daughter cells, the innermost of which 32 BOTANICAL GAZETTE [JULY becomes the archesporium, while the outer, the so-called ‘tapetum,’ forms a part of the sporangial wall. In Rosa’ and Fagus® the epidermis has been found to divide several times by periclinal walls, and thus to form a considerable portion of the tissue between the archesporium and the apex of ~ the nucellus. The unusually large number of cells in this — region in Potamogeton at first suggested that the same phenom- enon might also be found here; but a careful study showed that no divisions of the epidermis beyond an occasional doubling of — individual cells ever take place, and all of the tissue can be ~ easily traced to the primary wall-cell. Moreover, the similarity to the process in the microsporangia is so evident as to require : no other explanation. ae THE FORMATION OF THE EMBRYO-SAC. The lowermost cell formed by the first division of the hypodermal cell begins to enlarge at once, and must henceforth be considered as the archesporium. The whole process during the early stages of development is perfectly normal. The archesporial cell soon undergoes division resultirig in an upper and a lower cell (fig. 75). These probably corfespond to the two resulting from the heterotypic division in Convallaria, but the fate of the two cells we shall find is somewhat different either from that in Convallaria or in Canna. The first division is immediately followed by a second nuclear division in each of the daughter cells, but without the formation of a wall between the two nuclei(fig. 76). The uppermost cell now shows signs of disintegration, as indicated by the cytoplasm, which becomes more dense and also stains more deeply. The nuciei also lose their definite outline, and finally the whole cell becomes much compressed and flattened against the wall-cells above. In 4 very short time, indeed, it can be recognized only as a dark cap at the summit of the embryo-sac. Several preparations were SSTRASBURGER: Die Angiospermen und die Gymnospermen 14. Jena, 1879: °BENSON: Contributions to the embryology of the Amentiferae. Trans. Lint Soc. II. Bot. 3: 410. 1894. 1900] DEVELOPMENT OF THE EMBRYO-SAC 33 obtained showing the various stages of this process with great clearness. The lower cell on the other hand continues to enlarge. The two nuclei are usually located at opposite ends of the cell and are somewhat larger than those of the surrounding tissue (fg. 77). In the next stage observed both of these nuclei had undergone division, so that two were present at each extremity of the embryo-sac (fig. 78). At the antipodal end, from this stage onward,a small pouch begins to appear which contains the two lower nuclei and finally all of the antipodal cells. One of the two upper nuclei is now sometimes found slightly below the other, and nearer the center of the embryo-sac; but many other preparations show that this is merely temporary or abnormal, and that before the next stage is completed they are both located together at the apex of the cell cavity (fig. 79). Very soon after the last nuclear division a cell membrane can be seen to form around the two nuclei at the mycropylar end of the embryo-sac, thus enclosing them in a little pouch (jig. 79). The membrane constantly grows thicker, until at length it is a distinct wall, and the two nuclei are henceforth entirely Separated from the cavity below. This seems to preclude entirely the possibility that a polar nucleus may pass down and fuse with one from the lower group of cells, although the latter remains in the general cavity of the embryo-sac until a much later period. At a still later stage three nuclei instead of two are to be found in the micropylar enclosure (fig. 20). Two of these are small and differ but slightly from those of the surrounding tis- sue, while the third is much larger and located below close to the membrane. It seems probable that the two smaller nuclei are produced by the division of one of the two original nuclei, and are really to be considered as synergids; while the other and larger one is the egg derived directly from the other nucleus without division. The two nuclei at the antipodal end of the embryo-sac after the pouchlike extension has commenced to form usually lie one 34 BOTANICAL GAZETTE [JULY above the other and are both quite large, each containing a large chromatin mass in the center (fig. 79). The lower one now divides into three daughter nuclei, all very much smaller than the parent nucleus, and with the chromatin scattered instead of being aggregated in a ball. These are all to be considered as antipodal cells, and persist in the little pouch until after fertili- zation. The upper nucleus continues to enlarge somewhat until about the time of fertilization, when it undergoes division fol- lowed by a cell-wall, as a result of which one daughter nucleus is inclosed in a cavity at the antipodal end of the embryo-sac (fig. 2r). This then must be considered as a fourth antipodal cell, since it is one of the four parts of the original lower nucleus. The other daughter nucleus is the polar nucleus, but becomes at once the mother cell of the endosperm, and imme- diately undergoes division until a parietal layer of endosperm is formed (figs. 23, 24). The large antipodal nucleus and those of the endosperm, like the original nucleus from which they sprung, are all large and contain very large central chromatin masses. This, in addition to their position at successive stages, points toward a common origin. The large antipodal nucleus continues to grow for some time and is at length very conspicuous. It is by far the largest in the ovule, and the very large deeply stain- ing chromatin mass may be seen even after the embryo has — reached a considerable size. | The nuclei in Potamogeton are all very peculiar, differing © from the ordinary type in having the chromatin mostly aggre- gated in a ball at the center of the cavity, instead of being dis- tributed on the linin network. Although several hundred slides were prepared, none contained spindles in the embryo-sac, and > consequently the sequence of the divisions had to be determined — by other means. The appearance of such a remarkable process of development, and one so different from those already described, although still strongly suggesting certain features of — the process found by recent investigators in related plants, made it very important that every step should be verified as far aS _ possible. For this reason the material was worked over several : 1900] DEVELOPMENT OF THE EMBRYO-SAC 35 times, until the steps became so clear as to leave no doubt in the mind of the writer that the description as given above is correct. To summarize briefly: the mature embryo-sac consists of an egg and two evanescent synergids, each without a cell-wall of its own, but all contained. within a pouchlike cavity separated from the general cavity by a delicate wall. One synergid usu- ally disappears before fertilization. Next below is a large cav- ity containing the endosperm nucleus, and later the endosperm, derived without fusion from the lower polar nucleus. At the base are the four antipodal cells, three of which are very small and chromatic, and are descendants of the same nucleus, while one is very large and together with the polar nucleus is derived from another parent. The antipodals are separated from the main cavity by a membrane formed at the time of separation of the polar nucleus and the antipodal cell. The endosperm even after fertilization never becomes more than a parietal layer (fig. 24). The investigation of plants of the same and nearly related orders has shown that the occurrence of the large antipodal nucleus is not a peculiarity of Potamogeton alone, but is charac- teristic of a whole group of Monocotyledons. Schaffner? found — that in Sagittaria the development was normal up to the forma- tion of the two synergids, egg, and three minute antipodal cells. He then found that the two polar nuclei fused in the ordinary way, after which the definitive nucleus underwent division. This division was always followed by a transverse wall enclosing one of the daughter nuclei in a chamber at the base; and this was the one which became so large at a later period. Sometimes several were found, thus showing that the large nucleus had divided at least once or twice. From the other daughter nucleus Was produced the endosperm. The structure in question was therefore thought to be an endosperm nucleus, rather than an antipodal cell as in Potamogeton. 7 SCHAFFNER : Contribution to the life history of Sagittaria variabilis. Bot. Gaz. 23: 252. 1897. 36 BOTANICAL GAZETTE [JULY In Naias and Zannichellia Campbell® found an axial row of — two or three cells the lower of which alone became the embryo- sac. The development was normal up to the point where the polar nuclei were formed. The free egg-nucleus and synergids — and the minute antipodal cells were very characteristic; but no / fusion of polar nuclei was observed, and he doubts if it ever ‘ occurs. The enlarged basal nucleus is always present from this — stage onward, but is not separated by a wall from the main cavity se as in Potamogeton and Sagittaria. Campbell believes with Schaff- _ ner that this is a product of the first division of the definitive : nucleus, but also admits that it may be the lower polar nucleus — alone, while the upper has gone to form the endosperm. FERTILIZATION, The stages representing the various steps in the process of — fertilization are rarely met with in this plant. Several good | preparations were obtained, however, and these will form the — basis of the following description. 2 The pollen tube enters the embryo-sac through the micro- pyle at a point directly behind the egg. In its course it passes” close to the partially disintegrated synergid, but the tube t quite slender and scarcely inflated after entering the embryo-sa¢, thus differing decidedly from Sagittaria, in which Schaffner described the inflation as very marked. : The two sperm nuclei were last noted in the mature pollen grain where they were both inclosed in the same cell-wall. The: remain united even during their trip down the pollen tube to the” embryo-sac and enter the egg together. Fusion of the egg a! sperm nucleus was not observed. The stage immediately pt ceding this was found however, and is figured in fig. 22. . Her the egg is easily recognized, and lying in close proximity to it, indeed even touching it, are the sperm nuclei. In reality only one actually touches the egg nucleus. The other lies at the sid or even at the back of the first, and is already in the process ° 5 CAMPBELL: A : morphological study of Naias and Zannichellia. Proc. Calih Acad. Sci. I. 1:1, 1897. 1900] DEVELOPMENT OF THE EMBRYO-SAC 37 disintegration. It seems therefore that in this case the two sperm cells never separate. At this stage the wall which separates the egg from the rest of the embryo-sac is prominent, and can easily be seen to inclose the synergids as well. Camp- bell found both sperm nuclei entering the embryo-sac in Naias but only one in Zannichellia. Schaffner found that in Sagittaria one always remained in the pollen tube. THE EMBRYO. The classical investigations of Hanstein? and Famitzin* on the development of the young embryos in both monocotyledons and dicotyledons have made us familiar with the process in a large number of plants. The work of later students has only gone to confirm the results reached by these investigators, at least as regards general features. However, the results obtained from certain monocotyledons by Schaffner and later by Campbell seem to be at variance with those above mentioned ; and as bear- ing upon this point Potamogeton becomes of interest. In Potamogeton the fertilized egg nucleus remains for a very short time in the resting condition, perhaps it divides immediately, but lack of spindles in the young embryos made it impossible to determine this. Immediately after the first division, the basal cell undergoes a change whereby it becomes gradually larger until a size several times the original is reached. At this stage the cell is a very striking object (figs. 24, 26). The very large vacuole, which soon appears, at length forces the nucleus to the bottom of the cell where it henceforth remains, while the nucleus itself undergoes considerable enlargement. Just behind this large nucleus of the basal cell, one can often see a synergid, even as late as the several-celled stage of the embryo. The basal cell never undergoes division, but on the contrary remains for a long time in its enlarged condition attached to the end of the embryo-sac. But sometimes during the later stages 9 HANSTEIN : Die Entwickelung des Keimes der Monokotylen und Dikotylen. _ Bonn, 1870, *°FAMITZIN: Embryologische Studien. Mém. de l’Acad. Imp. Sci. de St. Petersb. VUI. 26:—. 1879. a eee athe as lek', 38 BOTANICAL GAZETTE [juLy of development it may, together with the embryo, become — entirely detached, so that the whole mass is then free (jig. 26). : The wall increases slightly in thickness, and is always thicker than any other wall in the young embryo. . The first division of the egg is soon followed by a second and — later by a third. The embryo then consists of a row of four cells ; of which the three upper have nearly the same size. Occasionally | only three cells are present at this stage, but the four-celled — Stage predominates. The next division is vertical, and in the i uppermost cell (fig. 25). This is again followed by one or two ~ more oblique walls, thereby dividing the terminal cell into several — Sectors, each extending from the basal line to the periphery. Periclinal and anticlinal walls form successively in the different sectors, as a result of which the upper cell soon becomes a mass of tissue (fig. 26). a Meanwhile divisions have occurred in the next lower cell. a This first formed an oblique wall, after which several divisions a took place in various directions. The third cell from the apex . has divided once or perhaps twice by vertical walls. The a embryo now is nearly spherical, a fact which makes it very diffi | cult to trace the development farther with the idea of determin- a ing just what portions of the mature embryo are derived from — | the various primary cells. The oldest stage at which one can with certainty distinguish the four original cells is shown im Z. 20. The subbasal cell has here und two vertical divisions. ergone a transverse as well - No further divisions took place in this — embryo w pee together with the basal cell form the true suspensor (fg 29), The fate of the other be inferred from their the upper cell and the a two cells is immediately lost. It caf position that the cotyledon arises from xis from the subapical. We cannot be eR ge a ee EN Te One eee a ee es Fe OP PET NT ee oe ee aT a a a aN a a aaa 1900] DEVELOPMENT OF THE EMBRYO-SAC 39 far wrong in making this interpretation. The half-grown embryo shown in fig. 27 was the oldest stage obtained. At this time only the epidermis was differentiated, while the plerome is scarcely distinguishable. At the upper end of the figure may be seen the cotyledon; and at one side the plumule, arising in a depression at the apex of the hypocotyl. Schaffner and Campbell both find that no division takes place in the enlarged basal cell after its formation. There is certainly none in Potamogeton. Probably the observation of Hanstein that such a division does take place in Alisma was inaccurate. That such a division does occur in some monocotyledons, how- ever, is a well established fact as shown by Coulter ™ in his studies of Lilium. In this plant the basal cell undergoes both longitudinal and transverse divisions. The process in Potamo- geton does not differ essentially from that in Sagittaria and Naias, as given by the two authors cited above. ~The subbasal cell in Naias divides by transverse walls into three instead of two cells, of which the upper forms several cells, while the lower remains undivided. In Sagittaria the subbasal cell divides also once more than in Potamogeton, and the uppermost daughter cells here again form considerable tissue. Canna Indica L. THE HYPODERMAL CELL AND ARCHESPORIUM. Canna represents still a third type of the monocotyledonous embryo- -sac, differing in method of development from Lilium Convallaria, and also from Potamogeton. The hypodermal cell very soon divides into two parts by means of a periclinal wall (fig. 29). The upper cell then by repeated anticlinal division rapidly forms a layer of about nine cells directly above the future embryo-sac. This layer together with the epidermis constitutes the wall of the sporangium, and remains unchanged until finally displaced by the embryo-sac beneath. ** COULTER, J. M.: Contributions to the life-history of Zz/ium Philadeiphicum — The embryo-sac and associated structures. Bot. Gaz. 23 :413- 1897. 40 ' BOTANICAL GAZETIE [yuLy Meanwhile the lower of the two cells formed by the first 4 division of the hypodermal cell has undergone considerable — growth. It rapidly becomes several times its original size, and extends nearly to the base of the nucellus; but the increase in breadth is not so great. Just previous to the first division of its + nucleus the cell is unusually long and narrow, but is completely filled with cytoplasm (fig. 29). 4 The cell gradually becomes longer and longer until at length _ a division takes place whereby two daughter cells are formed (fig. 31). After a short time both of these daughter cells divide _ again simultaneously. As a result we have an axial row of four _ cells reaching from the base to the apex of the nucellus (fg. 32). In the upper three cells a change is noticed almost at once, whereby the cytoplasm becomes denser and darker, the nuclei less chromatic, and the cells in fact show evident signs of — disintegration (fig. 33). The lower cell by its continued growth gradually compresses the other three, which very soon are all crowded into a small disorganized mass in the micropylar region — | ( fig. 34)-0-A large number of sections was obtained showing — all stages of this process in the plainest manner. The lower- — most cell alone finally takes part in the formation of the embryo- E sac. The process in Canna, therefore, is exactly in accord with 4 that in the Iridacez, Rosacee, Polygonacez, and many Ranun- culaceze and Liliacez as described by Strasburger and others. — Owing to the extremely narrow cavity and small nuclei, the changes within the embryo-sac are very difficult to follow. Humphrey” has already given a full discussion of the literature on the embryo-sac of Canna; and also the results of his owl investigations on the same plant. In this case the antipodal cells were not discovered, and he came to the conclusion, as did also Guignard*} that these cells although always formed must disintegrate immediately, Sie todeiahds The development of the seed in the Scitamines. Annals of Bot 10:1, 1896. *3GUIGNARD: Recherches sur le sac embryonnaire des Phanerogames Angio- Spermes. Ann. Sci. Nat, Bot. VI. 13: 136. 1882. ‘ 1900 | DEVELOPMENT OF THE EMBRYO-SAC 41 From the material at hand it appears that a short time after the last division in the axial row, the primary nucleus of the embryo-sac, 7. ¢., of the lower cell, divides, and one of the daughter nuclei passes to each end of the already much elon- gated cavity. The spindles representing the latter division were not obtained, but several cases were found where there were two nuclei at each end of the embryo-sac, and still others where there were four (fig. 34). One or two sections showed the three antipodals, two synergids, and the egg; while near the center of the cavity was a very large nucleus, apparently the definitive nucleus of the embryo-sac. The fusion of the polar nuclei was therefore not observed, but probably took place as indicated by the two nucleoli and cross line in fig. 35. Just previous to fertiliza- tion the egg-apparatus was found to be separated from the main cavity by a delicate membranous wall, and to consist of two very small partially disintegrated synergids located very near the micropyle, and-a much larger egg nucleus suspended some dis- tance below, as in fig. 35. At this stage the antipodals are often in an advanced stage of disintegration, and are more or less clearly separated by a delicate membrane from the cavity above. Canna differs from Convallaria, therefore, principally in the embryo-sac being formed from one cell of the axial row, and one element of the division into four of the mother nucleus. In the latter plant the whole axial row and all four elements of the division of the mother cell go to form the embryo-sac, THE NUCLEAR DEVELOPMENT. At a very early stage the nucleus of the archesporial cell passes into the condition of synapsis, in which as usual the linin is massed together at one side of the nuclear cavity. After a ‘time indications are seen of the gradual loosening of the knot, with a simultaneous migration of the spiral coils to the more dis- tant parts of the nucleus. The nucleolus here, as in Convallaria, is seen to remain intact during the whole process. It stains much deeper with the gentian-violet than does the chromatin, 42 BOTANICAL GAZETTE [JULY and can therefore be readily distinguished. The spirem itself is composed of comparatively few turns of the rather broad — chromatin thread, and is made up of alternate segments of chro- matin and linin (fig. 29). The nucleus lies imbedded in the cytoplasm, which at this stage completely fills the cell cavity, It is now very large, and several times the size of those of the adjacent vegetative cells, and from it numerous radiations — extend into the surrounding cytoplasm. a The nuclei and chromosomes in Canna are so small that lit tle could be done toward working out the segmentation of the | latter. Some features of the nuclear division, however, may be — noted. : : All of the nuclei in this plant possess a true nucleolus, as in Convallaria, and a very meager linin net-work on which the ; small amount of chromatin is unequally distributed. Inthe vege tative nuclei at the time of division the chromatin becomes 1 aggregated into six spherical masses lying just beneath the — membrane, and the nucleolus at the same time disappears. The — spindle now forms, and the ordinary process of division assures — | six daughter chromosomes for each resulting nucleus. This _ | count was made many times with great ease, owing to the small | number of segments, and always with the same result. : segments on their way to the poles ( fig. 30). The four counts here made gave in every case the number as six, instead of three as one would expect after reduction. The spindles of the sections containing these were also obtained (fig. 31). were all in the nuclear plate stage, and here the number wa actually three ; but each chromosome seemed to be compose ' of two parts, one of which was located directly above the other sl aa i ss a aa sa 1900] DEVELOPMENT OF THE EMBRYO-SAC 43 and more or less completely joined to it. The most plausible explanation for this seems to be that the segmentation for both divisions was nearly or quite completed before the formation of the first spindle, and that when they appeared on the nuclear- plate of the second spindle the segments had come together in pairs only to be separated again on going to the poles. No true resting stage seems to intervene between the two divisions. The above results are especially interesting, since three for the reduced number of chromosomes is one of the smallest so far found in plants. The later divisions in the embryo-sac were not observed. The spindle seems to be formed simply by the elongation of the kinoplasmic mass, no multipolar condition being noticed, and the mature spindle is long and slender but usually with obtuse poles. After each division a distinct mem- brane is deposited at the cell-plate, thus forming the axial row of four cells. The resting nuclei often show very distinct radiations from the nuclear membrane; especially is this the case with the lower one which now commences another period of growth before division in the embryo-sac. SUMMARY. Convallaria—The hypodermal cell divides into an upper and a lower cell, of which the inner cell becomes the archesporium and the upper forms part of the wall. This is also the case in Potamogeton and Canna. The stages of the growth and development of the archespo- rial nucleus are identical with those of the nuclei of the micro- sporangial archesporium. The first division of the archesporial nucleus is the hetero- typic division and corresponds to the first pollen-mother cell division in every respect. — The two spindles accompanying the second division are formed simultaneously. In appearance this division is identical with the second pollen-mother cell division and quite different from either the vegetative or heterotypic. A transverse division of the chromosomes could not be demonstrated. 44 BOTANICAL GAZETTE [july Only the heterotypic division is followed by a cell-wall, and | thus an axial row of two cells is formed each containing two nuclei. Shortly after anthesis the transverse wall disappears and the four nuclei all divide simultaneously, producing four daughter nuclei at each end of the embryo-sac. The endosperm nucleus — is formed by the fusion of one nucleus from each group. After — the transverse wall is destroyed, therefore, the process is the — same as in Lilium. i The number of chromosomes in the vegetative nucleus — is about thirty-six. During the heterotypic and so-called — “reducing” divisions eighteen may be counted. The apparent reduction therefore takes place prior to the first division of the — archesporium, + Potamogeton.—The first division of the archesporial nucleus is followed by a cell-wall, but the second is not; so that an axial } row of two cells, each containing two nuclei, is produced as in Convallaria, i The lower cell forms the embryo-sac, while the upper diss integrates. . Four nuclei are formed in the lower cell; the two at the upper end are at once enclosed by a cell membrane, and from | them develop the two evanescent synergids and the egg. 4 The lower remain free. From one of these all three of th small chromatic antipodals are probably formed. The other divides, forming the fourth antipodal cell and the polar nucleus with a cross-wall between the two. The polar nucleus become the endosperm nucleus without fusion. The mature embryo-sac contains two small synergids and large egg nucleus enclosed by a wall near the micropyle; @ very thin parietal layer of endosperm; and four antipodal cells enclosed by a transverse wall at the lower end of the embry sac, of which three are very small and one is very large. The nuclei of Potamogeton are peculiar in having most of the chromatin aggregated in a ball at the center of cavity. 1900] DEVELOPMENT OF THE EMBRYO-SAC 45 At the time of fertilization the two sperm nuclei lie together near the egg nucleus, but only one is really in contact with the latter. The fertilized egg-cell undergoes at first three divisions, form- ing a row of four cells. The terminal cell and the one next below give rise to the greater part of the embryo. In the subbasal cell only one transverse and two vertical divisions ever occur, and the cells thus formed together with the very much enlarged basal cell form the suspensor. Canna.—The heterotypic division is also the first division of the archesporial nucleus, and is followed by a transverse wall. The second divisions occur simultaneously and are also fol- lowed by cell walls. These form an axial row of four cells. The lower cell alone gives rise to the embryo-sac; the other three finally disintegrate and disappear. The further develop- ment is quite normal. The nuclei of Canna are more nearly like those of Convallaria than Potamogeton. They have a true nucleolus and no central chromatin mass. The number of chromosomes in the vegetative divisions is six. When passing to the poles at the heterotypic division there were still six; but later the second division showed only three as the reduced number. Probably the segmentations for both divisions occur during the prophase of the heterotypic division. This number is one of the smallest yet found in vegetable tissue. CORNELL UNIVERSITY. EXPLANATION OF PLATES VI AND VII. Figures 1-11. Convallaria majalis L. Fig. 1. A vertical section through a young ovule, showing the epidermis, archesporial cell, and two of the wall-cells between ; the archesporial nucleus is in the resting stage. Fic. 2. The same at a later stage ; the wall-cells have undergone further oe and the archesporial nucleus is in synapsis. G. 3. The archesporial nucleus in the spirem stage; the ribbon contains doait denser portions of chromatin. 46 BOTANICAL GAZETTE | JuLy Fic. 4. Nuclear-plate stage of the first or heterotypic division of the archesporial nucleus; the chromosomes appear + -shaped, as in the pollen mother-cell. Fic. 5. Anaphase of the same division; the V-shaped segments have . reached the poles, and the cell-plate is forming. Fic. 6. The nuclear plate stage of the second or so-called “reducing” division ; the chromosomes are arranged very differently ; between the two spindles is the wall formed during the previous division. 1G, 7. Anaphase of the same division. Fic. 8, A later stage showing the four nuclei thus formed. FG. 9. Each of these nuclei now divides forming four at each end of the — embryo-sac ; the cross-wall has disappeared. ; Fic. 10. The mature embryo-sac ready for fertilization ; over the apex is the epidermis and the remains of the other wall-cells; the egg-apparatus consists of the large egg nucleus and two striated synergids ; at the base are the disorganized antipodal cells enclosed by a cell wall, and above is the endosperm nucleus. Fic. 11. A spindle from the vegetative tissue, nuclear-plate stage ; the : chromosomes extend in all directions. a Figures 12-27, Potamogeton foliosus Rat. Fig. 12. A cross section of the apex of the nucellus, showing at the cenlet the two wall-cells lying above the archesporium. IG. 13. Same in the four-celled stage. Fie. 14. Vertical section of the nucellus, showing the archesporial cell with resting nucleus, and one wall-cell above. Fig. 15. The axial row of two cells ; the cross-wall was formed after the : first or heterotypic division of the archesporium. Fic. 16. Same, later Stage; the “reducing” division is not followed b! el cell-wall, ie nt See ata Fic. 17. The lower cell developing into the embryo-sac ; the upper © : forms a crushed mass above. 1G. 18. Same, but each nucleus has divided again. : FiG. 19. Same, with a wall forming just below the two upper nucléh Separating them from the Cavity below. 1G. 20. The young embryo-sac with the large egg and two small syne : gids enclosed by a cell-wall, three small antipodal cells at the base and a ve : large nucleus above. : Fic. 21, Same, slightly older; the large nucleus has divided into 4 polar nucleus above and a large antipodal nucleus below separated by a cell-wall the former becomes immediately t Fig. 22. Egg a nuclei lie ne tion, ee he definitive nucleus. 419-428. Ig00. ®Mac the Monotropacee : Bull. N. Y. Bot. Garden I ra i a CS eee eee Ae ee ee ee eee 1900] CURRENT LITERATURE 71 discussion of their anatomical and ecological features. No. 14 of the same series (issued May 30) contains an account (with 1 plate) of the Japanese Mutisiaceze from the collection of Faurie, by A. FRANCHET. No. 15 of the series (issued May 30) contains an account of certain new or little known Chytridinee, by E. DE WiLDEMAN.— Notes on some type specimens of Myxomycetes in the New York State Museum is a useful paper by W. C. STURGIS, published in Zyans. Conn. Acad. Arts and Sct. 10: 463-490, Pls. 60, 61. 1900.— F. E, Ltoyp and L. M. UNDERWoop (Bu//. Torr. Bot. Club. 27 :147-168. fis. 2-4. 1900) have published a review of the North American species of Lycopodium, recognizing 29 species, two of which are described as new.— P. A. RYDBERG (dé/d. 169-189. f/s. 5, 6) has begun a series of papers entitled “Studies on the Rocky Mountain Flora,” the first one being devoted to the Jobatus, aureus, subnudus, and tomentosus groups of Senecio, in which assemblage of forms he recognizes 30 Species, 18 of which are described as new.—W. R. Maxon (tb¢d. 197-199) has described a new Asplenium from southern and Lower California.— S. C. Stuntz (did. 202- 211) has published a revision of the N. Am. species of the genus Eleutera (Neckera), recognizing 6 species, all with new names.—G. N. Best (ddzd. 221-236. és. 7, 8)has published a revision of the N. Am. species of Pseudo- leskea, recognizing 7 species, two of which are new, and 4 varieties, all of which are new.— E. P varieties.—J. K. SMALL (zh¢d. 275-281), in continuing his notes and descrip- tions of N. Am. plants, has described 11 new species.— FLORA W. PATTER- SON (iid. 282-286) has described 17 new species of fungi—Geo. V. NASH eg A Garden, 1: 429-437. 1900) has described 11 new grasses ep the southern states, and a new Trisetum from Michigan.—J. K. SMALL a 437-447) has published a revision of the genus Bumelia in North Bac recognizing 13 species, five of which are described as new.—N. L. Poa (ebid. 447-449) has described 7 new species of Ci ratacgus.—MRs. tex (Zo vid BRANDEGEE has published a second paper in the series on Cac- It lool 5*1~9. 1900), the first having appeared in Evythea (3:1 23. 1895). new nS critical remarks upon numerous species, and descriptions of one Species of Cereus and two of Mamillaria.—J. M. C NEWS. PROFESSOR Dr. H. AMBRONN has been called to the assistant professor- ship of botany in the University of Jena. PROFESSOR J. W. ToumeEy, of the University of Arizona, has been j appointed assistant professor of forestry in the new school of forestry at a Yale University. ’ WITH THE BEGINNING of the present volume of the BOTANICAL GAZETTE, : Dr. J. C. Arthur, on accdunt of ill-health and pressure of work, retires from : the position of an active and responsible editor. For eighteen years he has — been identified with the journal, and has been a large factor in whatever 7“ cess it has achieved. In 1882 he became an associate editor, and in 1886 oF he became an editor, sharing financial as well as editorial responsibilities. Th GAZETTE will not lose his counsel and active cooperation, for his connects will be continued as an associate editor. THE FOLLOWING FELLows in Botany have been appointed by the Un versity of Chicago for 1g00-Igor: S. E. M. Coulter (Hanover College), B. E. Livingston (University of Michigan), and A. A. Lawson (University of ornia). A.C. Moore, a Fellow of last year, has been appointed Assist honorary fellow. WE LEARN from Science (June 15) that the appropriations for the Dep ment of Agriculture for the fiscal $4,023,500, being an increase of more ‘Stations in the forty-eight states and territories, $12,000 for the stati Alaska, $10,000 for a new station in Hawaii, and $5000 for an investig and report to Con Puerto Rico. Division of Agrostology, $25,100, appropriation of $30,300; whi increased from $130 1000 tO $170,000, an increase due in large measure q petition of some 225 members of the House of Representatives. 72 [0LNs Horsford’s Acid Phosphate The most efficient remedy known for the relief of languor and exhaustion, so common in the spring and summer months. Taken after exhaustive illness it acts as a wholesome tonic, giving renewed strength and vigor to the entire system. Taken before retiring, quiets the nerves and induces refreshing sleep. Sold by Druggists. Genuine bears name Hor SFORD’s on the wrapper. oz0don in a new size 25c. of the Liquid The event of the year in dentifrices. Beware of counterfeits and substitutes of this, theworld’s best known dentifrice. Insist upon getting the genuine at the stores. It necessary send 25c. direct to the Proprietors, P. O. Box 7, New York Ci City. eS HALL & RUCKE L e NEW YORK LONDON ENS BORATED TALCUM Delightful After Bathing Uxury After Shaving A POSITIVE RELIEF FOR CHAFING eel Hr —s and SUNBUR Ye nd - ‘ctions of the skin. R ‘ | “Wee Pitation. . : ie S (the origi ( Rinal), 2 “#7, haps, than Worthless On Sor it. , ders, which are t~ ; : s Joa} here, or Mailed for 25 cents ERMA RO MENNEN co,, Newark, N uo we Boston: Philadelphia: Chicago? 504 Fulton 8t, 69 Tremont St. 924 Chestnut St. 74 State St. upon Chlorides, Pour a little Platt’s Chlorides trequently into the traps of the water closets, wash basins, sinks, etc., and all foul gases will be neutralized and disease-breeding matter destroyed. att’s Ch is less, colorless liquid disinf fi nomical; sold in quart bottles only, by druggists, high-class grocers and house furnishing dealers, Prepared only by Henry B, Pratt, Platt Street, New York, No. 179 THIS IS ONE OF OUR LATEST It is one of 60 styles illustrated and . scribed in our catalogue “ B,” for 1900, . ° ifs Rolling and Carrying Chal i idi "t exist for Th of invalidism doesn’t exist i ahicbwe cannot furnish a suitable chair. es We also make the best types as well as the largest™® to be found of ne Coun lining Chairs and Adjustable Co 10 ior Sick Folks, Well Folks, and ue all of which are illustrated and described in our n iti g for informat , par av sae et rize. *C.” In writin ticula’ GEO, F. RGENT cCOMPAY 8 eee oe at a NOMINAL PRICE. Chicago's larg- est music house, ments, at almost nominal prices. Good durable uprights as low as $100, w from $200. Send for complete list. the makersare: Decker Bros., Hardman er Fischer, Hazelton, and others. If yo are interested ina pian write. Any piano not proving exactly as represented may be returned at their expense. Address LYON & HEALY, 100 Adams St., Chicago. Ne gs, 0,do not fail to ee a ae ae a ae ae ae a ee ae ae ae a a a a ee ee a a ee ee a ee $9 =] © # fa8] i=] jor mn a Q ie) S OTT eS ae Te he hh ee Te eS ee ee ee ee eee ee Oe ee ae ae ee SA k 289 N. Fourth Ave., next 23d St. New YE) ee WITHOUT THE USE OF ann peer FeO GEST . ae i Penne INSTITUTION IN TH rs For the exclusive treatment of cancer, tume of mali : ibenig ths, exe in the abdominal cavity by a graduate withheld as ian to m n. nd E. BROWN & SON; North nF A. FIVE OF EM] FINE ‘UNS, EM} Big Four Route |i@eaaes . BETWEEN NEW YORK ann BUFFALO. 33 OF “EM THE ONLY LINE FROM THE Northwest CONNECTING IN THE CENTRAL UNION DEPOT, CINCINNATI, WITH N 5 . Charleston, S.C WRITE FOR MAPS ATES, Etc., TO H. W. SPARKS secr, CHI roe J.C. TUCKER, GN 234 Clark Street, CHICAGO, Po cag pk nN ne Se CHICAGO : % ¢ ALTON ¥ RAILROAD. CHICAGO 0 KANSAS ciTy, CHICAGO «x0 ST. LOUIS, HICAGO ano PEORIA, C T. LOUIS «xo KANSAS CITY, ; ckwiesa | Railroad | -C.RUSSELL,. 7. CHOICE OF MANY ROUTES TO CHARLESTON BY THE SEA Wi? RAINS. Mary, TRAINS BETWEEN IEW YORK anp St.Louis": KANSAS CITY. it Railroad. Vestibule Trains, Comfortable Coaches, Luxurious vg nt Cars, B.D. CALDW TRAPHIC Man Fine Café Ca Roomy PatiorC ars W.LEE : DOWELL E.G.RUSSELL at VIA THE POPULAR N. E. A., July 7-13, 1900 MONON ROUTE HICAGO. [NDIANAPOUNS & fOUISVILLE [AlLWAY and C. H. & D. R’y sesians pps THE BEAUTIFUL ND HISTORIC SOUTHERN ALPINE REGION D. ’ AND RETURNING IF DESIRED VIA WASHINGTON _Throngh Pullmar HOT SPRINGS. A carxAS. acon . DENVER. Colo. ON LIFORNIA AND OREG Parlor and Dining Cars on Day Trains Palace Sleeping Cars on Night Trains ph- City Ticket Office, 232 Clark Street CHICAGO any portion of 1 are con ss reo be aa tenn pla tn su to =A aon rite to the Pundersig y- trip, Chicago & ‘Alton, it will ned for ms aps, pam CAGO, ILLINOIS, ime -ta ables, e GEO. y, CHA ral Passenger and Ti ket A. CHIC icket Agent, S INVESTIGATE ( THOROUGHLY § §& » BEFORE YOU PURCHASE A TYPEWRITER. THE SMITH PREMIER Will merit your approval as It Tully meers requirements af every point "ART CATALOGUE FOR THEASKING. © THE SMITH PREMIER TYPEWRITER Co. x SYRACUSE, NY. U.S.A. )" Because The Mammon : Typewriti Is easy, to Operate and io Schooks aad Cale Its Interchangeable Tppe — kes it the Most Usd’ Teachers and Clergymet inclose 4! Send for new Catalogue, and ine stamp for a correct Map of the "i mine the ae 1 4 i. os XY Spacing Attachment just added to the Hamma THE HAMMOND TYPEWRITER a 537-551 E. 69th St., New ——— The whether shapely or ugly, matters little, so long as it does not break or crum ble, and the quality is smooth and ng. DIXON'S test PENCILS their smooth, tough ; S L SPEE Simp hardness, suitable for all kinds of school work, and are indispensable in the drawing class. Samples worth double the money will be sent on receipt of 16 cents, if you mention this publication, Joseph Dixon Crucible Co — ITER) : ANKLIN TYPEWRITER (| JERSEY CITY, NJ.” | QURAN REIN PEENeW Mi) RELIEF PEN at No. 314. Ease in Writing Unsurpassed o 0 other varieties of stub pens. . [5 ¢) styles fine, medium and blunt points. . . ASK YOUR STATIONER FOR THEM. THE ESTERBROOK STEEL PEN Co, 26 John St., New York. Works, Camden, N. J. A. F bea han apbsenees eee ON. { i mE English Studies OF The University of Chicago LATEST ISSUE, Just Published. LEONARD COX, The Arte or Crafte ‘ | Rhethoryke, 1530: a Reprint, edited CARPENTER, Ph.D. The first English Price, 50 cents, — OTHER NUMBERS 1 ae ot of Gods. 196 sie Ay RiGGs, Ph.D. 8vo, paper. Mi ‘ ‘Se og oe pantie | in the in E a treatm worth 75 eer, emt of Nature tween Pope and Words- ¥ x ‘ «™ Sent Postpaid on receipt of price, ae Address THE UNIVERSr ry OF CHICAGO PRESS CHICAGO, ILL. We recommens “ KOH ILNOOR” PENCILS a I be had o y High genet ~~ a hy iataral: Tealer in Am FAVOR RUHL & CO., 123 W. HOUSTON STREET NE Thirty Years of | Babbitt’s Best S04) i i ' It has stood the only infal- : lible test—the test of time | Made by B. T. BABBITT, New York, —_——_—_ Y SECRETS EXPLAINED E Cause of Impaired Vision able opportunity for treatment. Everyone should know how to care for the eyes, and thus avoid dan- gers sure to follow neglect. An excellent book, “THE EYE, ITS CARE AND TREATMENT,” free for 2c.stamp. THE GLEN SANITARIUM CO., Glens Falls, N. Y oe hee ee The Botanica] Gazette Edited by John M. Coulter, C. R. Barnes and J. C, Arthur, with American and foreign associates, Monthly; at least 80 ima ° =) Pa al eo < or o 4 a i] ° o n = ad students, and news items. $4.00 a year; foreign, $4.50; single copies, 40 cts. All subscriptions and requests for sample copies sho ; uld be addressed to _ PURE! HEALTHFUL! STRENG DY «og: University Presg Division, Chicago, Il. Sold at om eoneeee: an RY WHERE The University of Chicago ; sh "ie waisted boy leaks with nvy on ages BRAND clad p laymate. Tl hes ; 1 st will send, pr Pi id,ohe shirt and collar on receipt of $1.00, bio rae style and a wanted. Addre The Improved BOSTON | GARTER ae ee ee a Oe Ae The Standard | for Gentlemen ALWAYS pate The Name “ BOST = ae shaciped on every loop. YS — flat to the leg—never lips, ty nor Unfastens. aN SOLD EVERYWHERE. > Sample pair,Silk me Cotton 25¢ = GEO. FROST co., Makers, Boston, Mass., "U.S.A, Oe EVERY p PAIR WARRANTED PENANG “] go woolward for penance.’ * wey sips ee him in Rome for want of linen. —Love’s Labor Lost, Act V, Scene 2. The torture which the wearing of wool next to the skin inflicts, and the sanitary qualities of linen under- wear, were apparently well recog- nized by the Master Mind of English literature. Dr. Deimel’s _Linen- Mesh Underwear is too comfortable to appeal to those penance. Its perfect tary qualities entitle it to the con- sideration of all who ev that a —“— body goes best with ~All true Deimel Linen-Mesh gar- ments bear the above Trade-Mark. If you cannot = them from your dealer, write to u Booklet and ete pieces free. We also manufacture 1 finest dress shield in existence. an be washed, are odorless. A guarantee with every pair. THE Deimel Linen-Mesh System Co. 492 Broadway, New York. 1 Montgomery St., San Francisco, Cal. ngton, D. C. a4 15th St., N. W., Was 10-12 Bread St., London, E. C THERE Is ONLY ONE EX | RACT OND érGoNewg ty es THIS Is IT! wih Bult Wrapper, Invaluable for all Aches, Pains, Inflammations, Catarrhal Trouble and Piles. JD DIRECTIONS WH APE REE egy ea POND’S EXTRACT CO., NewYork and London. ponte “IT’S ALL IN THE LB Have you a i ° eautiful Picture == Stereopticon ore in your School ? If not, why not purchase one and make your teaching twice as effi- cient? All admit the eye is the only medium to reach the brain, and pictures impress truths in one-tenth the time of oral demonstration. Illustrated story books prove such to be the case... T, € have thousands of sub- a jects in slides, and make anything desired itha for the purpose. Write for catalogue to— Is Always Made Wi 4 ra RILEY BROS.. Korona Came 16 Beekman St, RAO age NSES- \ TRADE L New York City OUR F AMOUS_LENSES Branches ae aan ae +» tor7 Walnut St, Mass., 36 Brom mfield St, Room 38, beep lis, Minn., 222 Nicollet Ave. m St. a Send for Catalogue GUNDLACH OPTIC Chicago, I., orx TE — 700 Clinton Ave., South, ROCHES! a “ESPECIALLY THE Registered by U.S, Patent Office BUFFALO ITHIA WATER of Virginia.” Springs Nos.1 and 2. For Albuminuria and Bright's Disease CHRONIC AND ACUTE. | Samuel 0. L. Potter, A.M., M.D., M.R.C.P. -; London, hea aps of the Principles and Practi of Medicine eg the College of Physicians and Surgeons, San mA cisco, a recogniz ate hag na cage science is known, in his handbook of Pharm cy, Mater! a Medica and eas er , Under hea of ALBUII TIINURIA, a e 600, 6 of nn > Tinks the citation of remedies, says: BUFFALO. LITHIA WATER i is h ighl ende | the: f reme Ble; fin acon: BUFFAL ALO LITHIA WATER: of albumen, epithelium, ie Sr anular ntirely disappear from the urine under its a distre while in thos: Pease areca y ER ‘s for sale by Grocers and Druggists generally. €stimonials which defy all imputation or questions sent to any address. PROPRIETOR. BUFFALO LITHIA SPRINGS. VIRGINIA. A wie i te oh te oh ae hh me eh oe ae Kh oe he ae oe a ae: SE ate 22 we Se A ee ot oh es 24 oe wth os ck es ce Ao wes ch OH Se ee ea ee a ee a ee Ae eee, tian AP atta ba, Ba, ih th ht, tl, a ll, **Preserbes Health’’ “* Prolongs Life *” BAKER’S BREAKFAST COCOA PIANO “Among ail the instruments o makers, here and abroad, | today prefer the We because of its sympathetic tone quality.” 7 April 5, 1900. EM “Its exquisite tone has heen a soumed@ delight.” ; April 7, 1900. CLEMENTINE DE‘ “ Perfect for accompanying the voice” 7 March 22, 1900. ERNST VAN “Congratulating you upon the mooie superiority of your magnificent pianos.” Feb. 7,.1900. ALY and caterer.” —Dietetic and Hygienic Gazette, Walter Baker & Co, Lid. “The quality and tone of your beautiful ments have been entirely satisfactory tom April 5, 1900. POL PLAM i DORCHESTER, MASS. WAREROOMS: =~ on Every Package Established 1780, Fifth Avenue and 16th Street, New’ Qe eee eee eee eT eT er eT eye ee ‘__ >. 2 >. 4 as tt me tt Des A ae ck te tek De De. De. mee wee 4 Oe.d Det De. de Be te A bl =r TP SEevruwwres= Pe ONelUvrhCUTClhUPCUOrhU DCUMC SECU ell rhlU OCCU eTClCU eC lULehlUCc Te hULrrrhlUCl Ol rele hm LU UL a 268 Wabash Avenue, Chicago. Fury SP SFRkFaesaeeayp 181 Tremont Street, Boston. - iM 2a ‘Fill itup yourself=-we have no list of your possessions. bull whether you own a cycle,asuna| ),. YaCht.a creel.a bait box.or,)) ;- anything from acastle towlf, mse Canteen you should. ep it bright.and. if =d0 itwith 2S 75/ Cor es nn \ “at >t ‘ ye “4 ‘i ie Sa a “ \ i] APES Vateenets ss Gems fe M4 | 5 i 5 Vy i} ii if Vol. XXX 4 J.C. ARTHUR : rdue University CASIMIR OLE Gen bo} B. DETO “ : Havant of Padua ADOLF ENGLER University of Berlin : LEON GUIGNARD 7 L Ecole de Pharmacie, Paris : ROBERT A. HARPER University of Wisconsin -JINZ6 MATSUMURA Imperial University, Tokyé @bhe Anivers AUGUST, THE EDITORS JOHN M. COULTER ann CHARLES R. BARNES, 1900 No. 2 . BOTANICAL GAZETTE WITH OTHER MEMBERS OF THE BOTANICAL. STAFF OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS FRITZ NOLL api a Bonn VOLNEY M: SPAL Uni we > de ROLAND THAXTER Harvard University WILLIAM TRELEASE Missouri — Garden H, MARSHALL WA Oniversity re ele eee EUGEN, WARMING University of Pas VEIT bed he oval ae ee Sciences, Stoc CHICAGO, ILLINOIS _Bublishe by the Gnibversity of Chicago itp of Sotseks press COPYRIGHT 1g00 BY THE UNIVERSITY OF CHICAGO Botanical Gazette | a a@ontbly sournal Embracing all Departments of gr essa Subscription per year, $4.00 umbers, 40 Cents The subscription ree must be paid in advance. No numbers are sent = the expiration of the time paid for. No reduction is made to dealers or agent FOREIGN AGENTS: q Great Britain—-W™M. WeEsLEY & Son, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin _ St., Strand, London. 18 Shillings. SW. 46, Schhnahchete: 17a 18 Marks Vol. XXX, No. 2 Issued August 15, 1900 CONTENTS : THE DEVELOPMENT AND FUNCTION OF THE CELL PLATE IN HIGHER PLANTS (WITH PLATES vit AND Ix). A. G. Timberlake — - 73 _ CONTRIBUTIONS FROM THE shea eee egae LABORATORY OF HARVARD cg 4 VE XLIV. NEW OR TLE KNOWN UNICELLULAR ALG. I. CHLOROCYSTIS COHNII ea PLATE X). George Thomas Moore - 100 | BRIEFER ARTICLES. NOTE ON THE MECHANICS OF THE SEED-BURYING AWNS OF STIPA AVENACEA (WITH FIVE FIGURES). LZ. Murbach 113 Some New Species oF Wyominc PLANts. Elias Nels = ae Sia New Nortru heiageke MossEs te, PLATE ai Jobo 3 M. Holsinger . -. 122 Notes oF TRAVEL. III. D. G. Fairch st -- ANOTHER NoTE ON THE FLOWER and OF OLIGOTROPIC ; make CBeaeel Robertson - 130 CURRENT LITERATURE. _ BOOK REVIEWS . . i - - : - . ‘ Bn: A PRACTICAL GARDEN Book. THREE POPULAR BOOKs. 2 5 MINOR NOTICES - * “ . - . : 3 2; i 133 e NOTES FOR STUDENTS : 3 i 2 ‘ ry - eee, 3: A : . 142 : Separates of any article (not less than 50 co ill d onl n special order in advance ~ pies) will be printed only upon spe Publicati ation. 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Wesley & Son’s New Botanical Catalogu ) wae Being CONTENTS: No. 127 128 of Bibliography § History Biographies and Pt Herbals Early Botanical Science Linnaeus Handbooks Dictionaries NATURAL Microscopy Morphology and Physiology HISTORY and Botanical classification Nomenclature SCIEN TIFI oaeagelhe Botany Encyclopaedic works Cryptog. Phanerogams Natural Pees of Plants (Floras) Fossil lants — of Scientific Societies Periodicals pplied Botany Botanical Agriculture and ad et to the end of the 18th century Gardening Ornamental Plants Food Plaats Medical Botany Tobacco Gums, Caoutchouc, ett headings Fibres Forestry Agriculture 92 pages Diseases of Plants ; William Wesley & Son, Booksellers, 28 Essex Street, Strand, . - London, England BSS 5555555005545 559 Allgemeine Botanische Zeitscl ui Systematik, Floristik, Pllanzengeographie etc. BOTANY PRISER : io) BERS Die “Allgemeine botanische es eg ” bringt vor allem Abhandlungen iiber Pest pf ru nosen kritischer Arten nd Bastarde, Schilderungen Hloristisch Une geographisch interessanter ae ‘strlen ag sar Referate iiber systematische, fl und 1 Pf St ig ga S ten, Berichte iiber die Thiti. gheit Fitoliacker Institute, Vereine 7 vereine etc., una deren Schriften und Katalo ie biog ra hische Notizen etc. | sine besondere Sorgfalt wird auch d Re id “abe Exsiccatenwerke, botanische Te auschhataleg 4 botanische Reisen ewend. m Ziel u. a. die Herausgabe von Lixsicca attenwerken bildet. rege Der komplett vorliegende Jahrga 99 wurde unter Mitwirkung von 43 Botanikern ne Si naa 40 Ori 3 — 31 Referate, — Sangaben von 20 bot. Zeitschriften, ‘erichtet ae c. von K's c., tber 32 Tauschvereine und Exsiccatenwerke, Ube ®. und brin ion gelangende Personalnachrichten von Bolsi” n gt simtliche zur Kennt lies Rrdicile ur Kenntnis der Redakti ia | ie Die “Allgemeine botanische Zeitschrift” erscheint piinktlich am 15. jeden pe ye so ee rsehen in mindestens Bogenstirke, k Mk. Abonnenten portofrei unter arr Bosca a eee | er Herausgeber: A. Knewciers st oe Karlsruhe in aaes (Deutschland), Werderpi®™ " Glumaceae Exsiccatae. | Die “lum den ‘Carices exsiccatae,’”? werden auch Mitarbeiter aus gesucht. ” sliedern S es sich in Gramineae Cyperaceae et Juncaceae exs. Spe reichliche Tuametaie einer Spezies einsendet i ; i jpezies bti Ng diesem Werke, Felt zu ( mace 4 jief. aa : -); die Carices A Lief. 8 Mark f Buchhandel 10 Mark «ae Jede Lieferung enthilt ik Wadia ni Mark (10 fres.), im Buc La inte t Lief.C Cyperaceen und po ataeriay ge Bis jetzt sind ik ee : 7 Lief. Carices, 2 “atten sinf 4: Lief. ist i d die Etique Prick hergestellt eae elte Material eda sate Mp Santee obapta : acli ramineae), Prof, Buchenau (Jun ge formen); f a it Palla rae icp: e), Ww vagrants — pikeathal kowitz. r Material ‘cinsenden will in Kacherah ck ine Gicetsthlend), Werderplatz 48: _ Paid-up Cash Capital, $1,000,000.00 ASSETS, - 27,700,511.50 Life Insurance in force, . 3 $ 00, 334,554-00 Returned to Policy holders, . 395734,920.89 ‘ J. G. BATTERSON, President. wtb DUNHAM, Vice President. H. J. MESSENGER, Actuary. ; JOHN E. MORRIS, Secretary, E. V. PRESTON, Sup’t of Agencies. ie a aN Sek eT ADAPTABLE TO ANY STYLE OF RECORD OR ACCOUNTS WE VISIT EVERY TOWN IN THE UNITED STATES SALES OFFICES IN ALL THE PRINCIPAL CITIES BAKER-VAWTER COMPANY LOOSE LEAF SYSTEMS Z < tf ATCHISON, KAN.-CHICAGO—HOLYOKE, MASS. 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The Lakeside Press PRINTERS AND BINDERS THE p TION, ot OF BOOKS DEMANDING TASTE AND CAREFUL EXECU- SPEC] ATE EDITIONS, COLLEGE CATALOGUES AND ANNUALS A Street, Chicago. TH Be 0D DS Sh AS DD LD 1D dD HG ' AN OPPORTUNITY TO VISIT THE EAST rist tickets on sale va omg Sho eee aictiest Goeth ern bd es pir after June rst. Chautauqua Lake, Niagara Falls, the St. Lawrence River, White Mountains and the Atlantic Coast ae es: Spon of Traias”™ sie amie ong meee kept tuteneetd in arrangaal for your trip. Sent free on application to F. M. BYRON, G. W.A., 144 Van Buren E NEW Ss otp b Shecive tdi BOSTON TRAIN w in servic Catalogue of North American Plants. 2 # 2 SECOND EDITION *” A. A. HELLER » 547 West Walnut Street LANCASTER, Pa., U.S. A. > Re OOD paper and printing G printing on one side 0 page only, larger type genera arranged and numbettl according to the Engler & Prantl sequence, and extél sive use of synonyms, are tee tures of this edition. Mad Price $1.00 per copy Pape covers, or $1.25 bound in cloth ee hal Sample pages may be ™ upon application. The Botanical Gazette Edited by John M. Coulter, C. R. Barnes and EC. Arthur, with American and foreign associates. Monthly; at least 80 pages. science of Botany in all of its departments, con- taining results of research, book reviews, students, and news items. $4.00 a year; foreign $4.50; single copies, 4o cts. All subscriptions and requests for sample copies should be addressed to The University of Chicago University Press Division, Chicago, Ill. “IT MA sen E aersis “MY oon i. tke aoe en hip and ‘6c in aes d Us st cara we ns tab ta! men Ratt Weeguin extab WELL CO., "THE train leaving Chicago at 6.30 p.m. daily was spoken of by the late Geo. M. Pullman as the “Finest Train in the World.” It is electric lighted, a Réclining Chair Car and Dining Wide-vestibuled trains are operated a ioe Pon Chicago and St. Louis City, Omaha and Den- “dh pene ed with modern, roomy, comfortable Pullman Cars and Re- clining Chai rs. opean plan Dining Car service is a special feature of ex nce this line BEST LINE CHICAGO OR ST. LOUIS TO perfect ver a ion and strictly first- class coo S t = if a U : ‘*Colorado Outings” ‘“‘California”’ re the titles of descriptive booklets 7 . A which 3 n n can be had without charge upon application to fae fh yl Bim erg, Passenger Agent, CB: & Q. KR. HICAGO. ea excellence on : Delicate china, roses, spotless linen, ati The School and Society THIRD EDITION ‘‘The [lost Noteworthy Book of the Year” BY JOHN DEWEY Professor of Pedagogy in the University of Chicago am. E problem of element tary education is one that forces itself - only on teachers ag ohe'te pe rds, but is felt with continuously growing anxiety b y the parents, and e educational situation has nowhere been ’so sleet stated nor so s raphically illustrated as in the odd hundred pages of Pugrey JoHN DEWEY’s ne $ e as terest excited there, the public has the guarantee of their interest and compre lity for all who feel the responsibility of bringing the meaning of life home to the child. **A most Pau contribution in the discussion va the educational Problems of the day, by an expert in pedagogics.’’— 7he Outlook. $<. 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HOME OFFICE - Presidens. ferred to, and issue, this attractive a} GOLD BOND palit in amounts of $5,ll and upward. The Protecti afforded is absolute The Investment excelled. Write for full information. VOLUME XXX NUMBER 2 BOTANICAL GAZETTE AUGUST, 1900 THE DEVELOPMENT AND FUNCTION OF THE CELL PLATE IN HIGHER PLANTS. H. G. TIMBERLAKE. (WITH PLATES VIII AND IX) Historical. AstpE from the work of Strasburger there have been very few investigations reported that have treated fully the subject of the cell plate and its history in the vegetable cell. So far as I know, the term cell plate was first used in its present sense by Strasburger in the first edition of Zedlbildung und Zelltheilung. Xt was here that he made the statement that the beginning of the cell plate is to be found in swellings of the connecting spindle fibers. The subject was more fully discussed in the third edition of the above work, and I shall refer to that in greater detail below. In 1878 Treub published his classic researches on the réle of the nucleus in cell division,* in which he describes the process of cell plate and cell wall formation in the living cells of the pro- embryo of Orchis latifolia and the ovules of Epipactis palusins. By keeping the tissues in a 1.25 per cent. solution of KNO, he — was able to make an extended study of the above processes. The first indication of the formation of a cell plate is a collection of 8ranules across the equatorial region of the spindle, which move into place from various directions in the cytoplasm. They * Quelques 7 aan sur le réle du noyau dans la division des cellules végétales. Amsterdam, 187 73 74 oy BOTANICAL GAZETTE [ AUGUST have, therefore, no connection with the spindle fibers. After reaching the equator the granules fuse into a continuous layer across the central spindle. The growth of the cell plate occurs © by the addition of new material either all around, if the spindle is in the middle part of the cell, or on the free sides, if the spindle is near one side of the cell so that the cell plate has reached the mother cell wall on that side where it is first formed, 7. e., the whole nuclear figure moves across the cell, building the cell plate as it goes. By plasmolyzing the cell by the addition of more KNO, Treub was able to demonstrate that the new cell | | these new fibers (of. cit., Anthoceros and the m | : ; : . j q 4 | : : i 4 ‘ p 1900] THE CELL PLATE IN HIGHER PLANTS 75 spore mother cell of Anthoceros and has found that the cell plate is formed upon cytoplasmic strands (not spindle fibers) uniting the four chromatophores. A portion of this cell plate becomes converted into a cell wall. In endosperm formation there are a great number of nuclear divisions that are not followed directly by divisions of the cell. The spindle in each case disappears after each nucleus divides.3 When the above process has continued until there are a great many nuclei lying free in the protoplasm, new fibers are formed connecting each nucleus with all of its immediate neighbors. In these fibers the cell plates are built in the usual manner. In a later work‘ these new fibers were described as growing out of a hyaline plasma mass around each nucleus. On the nature of the cell plate elements, 7. ¢., the granules which form the cell plate, we have an interesting statement. The cell plate is formed of small granules whose chemical nature is hard to determine. That they may be starch is indicated by the fact that they take in some cases a blue stain when treated with iodine. In most cases, however, they are not thus stained. They may be a substance between starch and cellulose. This hypothesis is rendered more probable by the fact that the cell plate elements are apparently used directly to form a cellulose = instead of being converted into a protoplasmic layer which Splits and excretes a cellulose layer between the halves. In the pollen mother cells of the Cycadacex, Juranyi® described a process of cell wall formation by the conjunction of 4 cellulose ring, growing in from the mother cell wall and a new wall formed in the connecting spindle. In 1882 Strasburger modified his previous view as to the °rigin and chemical] nature of the cell plate elements.” In this Paper he holds that with the aid of suitable stains it can be shown *STRASBURGER, Op. cit., p. 345. 4 Ueber Kern- und Zelltheilung im Pflanzenreich. Hist. Beitr. 1. 1888. ‘Zellbildung u ‘ius nd Zelltheilung, 3. Aufl., 342. a thej ve : 19:31, — tiber Structur u, Bildung des Zellkernes. Ref. in Botan. Centralb. 7U eber den Bau u. d. Wachsthum der Zellhiute 172. Jena, 1882. 76 BOTANICAL GAZETTE | AUGUST that the granules of the cell plate react as albumen. As to the origin of these granules it was observed that they are not added from the surrounding protoplasm but are within the fibers them selves. The fibers appear as if made up of small granules. These granules are really contained in the fibers as if in tubes, and they collect at the equator, forming apparent equatorial swellings af the fibers, z. ¢., the cell plate elements. The cell plate elements, — after growing by the addition of new granules, finally fuse intoa continuous plate. In 1887 Went® showed that connecting fibers and other spindle fibers are identical, and that the former apparently increase in number before the cell plate appears. He was not able, however, to account for the origin of the new fibers. Prior to the formation of the cell plate, Went observed a stained substance between the fibers of the connecting spindle next to the daugh- ter nuclei, leaving thus a clear zone in the equator of the spindle. This stained substance was thought to be nucleolar matter dis- solved in the nuclear sap. Inthe subsequent stages these two darker zones move to the equator where the cell plate is built. Its origin Went did not see, but he showed that its growth® peripheral, as had been well established by previous observels: Went, however, observed for the first time that the connecting | fibers disappear in the center of the spindle during the gro ol of the cell plate, so that the peripheral fibers form a ring -_ nected with the growing region of the plate. Opposed to the view of Strasburger as to the cytoplasm origin of the cell plate by fusion of swellings of the connectitt fibers, was that of Zacharias, who contended that the spines is of nuclear and the cell plate of cytoplasmic origin, thus fol lowing Treub’s theory of the origin of the cell plate from oe. granules. The cell plate is formed by the entrance of substal | from the surrounding protoplasm into the equatorial region of the central spindle, which is a portion of the dividing nucl ye ee Beobachtungen iiber Kern u, Zelltheilung. Ber. d. deutschen botan. Ges. - ° Ueber Kern u. Zelltheilung. Botanische Zeitung 46: 56. 1888. 1900] THE CELL PLATE IN HIGHER PLANTS id (Mutterkernrest) having a homogeneous structure and hyaline appearance. In a later contribution he ascribed the filar struc- ture of the central spindle to the effect of reagents, especially absolute alcohol. . In connection with his researches on fertilization Guignard *° observed that the number of connecting fibers is equal to the number of chromosomes, and that these fibers are formed by the fusion of smaller primary fibers. Later Strasburger™ confirmed the views of Went as to the staining properties of the nuclear sap and the relation of the substance in it to cell plate formation. He also described the formation of a temporary plasma membrane about the connect- ing spindle, and suggested that the change in the form of the nuclear figure, 7. ¢., the bulging out of the connecting fibers, is due to the osmotic action of the nuclear sap held within this membrane. In 1893 Wildeman™ described in the rhizoids of mosses a process of division wall formation in which the spindle is first arranged so that its long axis is parallel to the long axis of the cell. Previous to the formation of the cell plate, it is changed to an oblique direction, but the young cell wall is so curved as to become attached perpendicularly to the mother cell wall in accordance with Sachs’s law of “orthogonal trajectories.”’ ’ In 1895 Strasburger*3 claimed new evidence for the identity in character of connecting fibers and other spindle fibers in the fact that their staining qualities were the same. In preparations stained with the triple stain of Flemming he observed that, while pe cell plate is being formed, the connecting fibers go through gradations from violet (the color of spindle fibers) to brown (the cytoplasmic color), thus confirming his earlier opinion as to the cytoplasmic nature of spindle fibers. From the fact that ro Nouvelles études sur la fecondation. Ann. des Sci. Nat. Bot. VII. 14 : 163- Ir 4 - Histologische Beitrige 1: 162 ef seg. "2 Etudes sur |’ d attache des cloisons cellulaires. Mémoires couronnés et mémories + Say, etr, p- p. iy } ey Acad. Roy, d. Sci. de Belgique 53:19. 1893. aryokinetische Probleme. Jahrb. f. wiss. Bot. 28: 193. 1895- 78 BOTANICAL GAZETTE [AUGUST shortly before the cell plate is formed ‘‘extra-nuclear nucleoli” appear in large numbers in the equatorial region of the spindle, and that by the time the cell plate is complete these have dis- appeared, he argues that the nucleolus takes part in building the cell plate. : In the pollen mother cells of Liiium Martagon, Farmer * found that the protoplasm of the connecting spindle on each side of the newly formed cell plate is relatively transparent. | The series of contributions which‘appeared from the Bont laboratory in 1897 presented several important facts in regard to the formation of the cell plate. Mottier*s found in the pol len mother cells of the lily that the cell plate is split into plasma membranes before the cell wall is laid down. In cells plasmolyzed in fixing, a complete plasma membrane was see around each daughter cell, while no cellulose wall could be observed (of. cit., p. 192). In the pollen mother cells of Heme rocallis fulva, Juel** found that after the first nuclear division the cell plate did not reach the mother wall but remained free in the cytoplasm until the second nuclear division occurred and new cell plates were formed at right angles to it. Then some of the fibers radiating from the daughter nuclei become attached to es old plate, and all the cell plates now continue their respectité growths toward the mother cell wall. Debski*7 studied the di sion of the segment cells of Chara and observed that the long axis of the spindle is parallel to the shorter axis of the cell # further that the cel] plate is not formed across the equator : the spindle but nearer one nucleus. In the formation of ascospores in the asci of Erysiphe and Peziza, Harper*® fou 4 Kerntheilung in Lilium Antheren, besonders in Bezug auf die Centrosome frage. Flora 83 : 167. 1895. *S Beitrage zur Kenntniss der per Sia Se in ag Pollenmutterzellen ein Dikotylen u. — Jahrb. f. wiss. Bot. 1897. 5 6 Die Kern heilungen in den Pulls saci. yon Hemera files bei denselben PAN Unregelmassigkeiten. Jahrb. f. wiss. Bot. 30: 2°: HE pte liber Kerntheilung bie Chara fragilis. Jahrb. f. wiss. Be 397. 30: 227. ripe Kehibaig und freie Zellbildung im Ascus. Jahrb. f. wiss- Bot. 30% - age Spa a a ah ea Re i 1900] THE CELL PLATE IN HIGHER PLANTS 79 that the plasma membrane around each ascospore was formed by the growth and lateral fusion of the polar radiations, a proc- ess which is essentially similar to the formation of the cell plate in higher plants. In the division by which the ‘‘beaks”’ are cut off from the gametes in Basidiobolus, a typical cell plate is formed, accord- ing to Fairchild,’ from two rows of granules, whose origin was not discovered, in the equatorial region of the spindle. In some of the Sphacelariacee Swingle* confirmed the previous obser- vations of Strasburger, that the cell plate is not formed in a connecting spindle, but arises in the cytoplasmic network. Swingle suggested, however, that its formation may be under the control of the nuclei by means of the kinoplasmic fibers which radiate from the poles, although there are not enough of such fibers actually to form it. An important point in this con- nection is the fact that if the daughter nuclei are of unequal size the cell plate is formed nearer the smaller nucleus, as Debski described for Chara. In the oogonium of Fucus, Strasburger™ found a cell plate built of granules which he considers extra nuclear nucleoli arranged in the cytoplasm. The granules divide SO as to form two layers out of which are formed continuous membranes, Discussing the subject more generally, Strasburger” Places great emphasis upon the kinoplasmic nature of the cell plate. He points out the fact that the formation of a cell plate by the fusion of swellings of the connecting fibers is analogous to the formation of the plasma membrane around the ascospores as described by Harper. Inthe case of the ascospores there is no doubt of a transformation of the substance of the fibers (polar radiations) into a plasma membrané. In the case of the cell plate, while it is equally evident that there is a transformation of f ~ 6g Kerntheilung und Befruchtung bei Basidiobolus ranarum Eidam. Jahrb. : » Bot. 30: 285. 1897, ine dey Kenntniss der Kern- und Zelltheilung bei den Sphacelariaceen. Jahrb. f. * Bot. 30:297. 1897. * Kerntheilung und Befruchtung bei Fucus. Jahrb. f. wiss. Bot. 30: 351- 1897. cee, 1 30: 375. vTBope oPlasmastructuren im Kern- u. Zelltheilung. Jahrb. f. wiss. Bot. . 80 BOTANICAL GAZETTE [AUGUST fiber substance into a lamella from which is derived the plasma membrane of the daughter cells, Strasburger was not then certain whether the cell plate forms simply the plasma membranes, or whether a part of it may be changed into a division wall (9. aif, 38). In a subsequent contribution, however, he made the statement that the cell plate forms a plasma membrane (Hawt schicht), that it splits to form a plasma membrane for each daughter cell, and that the substance of the cellulose wall excreted by these daughter plasma membranes and laid dowi as a wall between them. Hof* has recently described in the division of vegetative cells what I take to be the same appearance as that described by Went and Strasburger, viz., the deeply stained portions of the spindle moving toward the equator and there taking part in the formation of the cell plate. He, however, describes the sub- stance of these zones as granular rather than as material i solution in the nuclear sap. After the cell plate is complet? the connecting fibers are entirely drawn in (eingezogen) and theif place taken by alveolar cytoplasm. Just what Hof means by the drawing in of the connecting fibers is a question left unsettled in his description. They may be drawn into the nucleus of into the cell plate. His figures do not help at all in understanding his meaning in this instance. is Nemec? has recently described the same phenomena 4 ee the formation of the cell plate in the growing root tips of Allin Cepa. He adds further that new connecting fibers make thet appearance after these zones have reached the equator. The fact that these new fibers often seem to end free betwee? the original fibers, Nemec considers as evidence for their origin ff™ the stained substance now at the equator. : The observations of Strasburger as to the method of cell plate formation in the oogonium of Fucus were not whol 88 z Ueber die karyokinetische Kerntheilung in der Wurzelspitze von A/iu™ aie Jahrb. f. wiss. Bot. 32 : 313. 1899. ae *3Zellhaute. Jahrb. f. wiss. Bot, 31: 514. 1898. *4 Histologische Studien am Vegetationspunkten. Bot. Centralb. 76: 221-3 1900 ] HL CELE PLATE IN HIGHER PLANTS 8: confirmed by Farmer and Williams.” These investigators found that there isan accumulation of material, which they think is some form of a carbohydrate, in the neutral zone between the daughter nuclei of the future oospheres. There is, however, no trace of a preliminary cell plate. While spindle fibers were seen reaching from the poles to the equatorial region of the cell, no evidence was found that they take part formatively in the building of the division wall. In an early paper Strasburger had pointed out the differ- ence between the method of division of the cell body of ani- mal cells and that of the higher plants, by stating that in the former the connecting spindle fibers take no part in the division of the cell body, but that the division is accomplished by the constriction of the plasma membrane; whereas in the latter these fibers are increased in number and take part in the division of the cell by forming a cell plate. This view Carnoy” attempted to overthrow in his work on the testicular cells of some arthro- pods, in which he claimed to have found a cell plate tormed of two parts, a cytoplasmic part formed by thickenings appearing on the junctures of the cytoplasmic network, and a spindle part formed by swellings on the connecting spindle fibers. I have not been able to find that this work has been confirmed by sub- sequent investigators. The spindle portion (plague fusoriale) of Carnoy is undoubtedly the midbody (Zwischenkérper) subse- quently described by Flemming*® and others as arising from granules in the equatorial region of the spindle, but not always as swellings of the spindle fibers. Notwithstanding the fact that it takes no part in the division of the cell body, Flemming was inclined to consider it the homologue of the cell plate in the pa plants. The midbody has received a great deal of atten- tom the zoologists, and various views have been held as to its significance and fate. The literature on the subject has been 26 Pe re Contributions to our knowledge of the Fucacez. Philos. Trans. of the Roy. a 190 : 623. 27 : La cytodiérése chez les arthopodes. La Cellule rx : 375. Neue Beitrage zur Kenntniss der Zelle. Arch. fiir mikr. Anat. 38: 690. 82 BOTANICAL GAZETTE [aucust recently succinctly reviewed by Ballowitz,? so that I need not go over it here. Ballowitz finds that the midbody is formed by the fusion of swellings of the central spindle fibers. _ His obser- vation differs from Flemming’s in that the latter thought the granules were sometimes between the fibers. Ballowitz and Flemming agree that the midbody takes no part in the division of the cell, but may be seen between the two daughter cells after division is complete. Ina very interesting paper by Kos- tanecki,° upon the relation of polar radiations to the division of the cell body, the view is developed that the division is effected by a cell plate built in the cytoplasm by means of two systems of polar radiations which connect the granular cell boundary with the centrosomes. During the metaphases the longer fibers from éach centrosome, which cross each other ia _ the equatorial region of the cell, contract and so change their points of attachments to the plasma membrane until they reach | the equator. They now draw in toward the central spindle, pull ing in with them a substance similar to that of the cell boundary; i oe Descriptive, The present investigation was undertaken to attempt to “ mine in detail the exact sequence of events. during the divis! of the cell body, and to correlate, so far as possible, the fact * Zur Entstehung des Zwischenkorpers. Anat. Anz. 14: 390. | ai Ueber die Bedeutung der Polstrahlung wiahrend der Mitose u. ihr. veri zur Theilung der Zellleibes, Arch. f. mikr, Anat. 49: 651. : : ey, Oe 1900] THE CELL PLATE IN HIGHER PLANTS 83 thus brought out from the point of view of the physiology of cell reproduction. MATERIAL. My observations were made largely upon (1) vegetative cells found in the growing root tips of various phanerogamic plants, among which may be mentioned Alum Cepa, Lilium longiflorum, Fritillaria imperialis, Fyacinthus orientalis, Vicia Faba, Phaseolus vul- garis, Pisum sativum; and (2) pollen mother cells of Lavix Ameri- cana and Larix Europea, and dividing pollen grains of Js versicolor and Hemerocallis fulva. In the first group I found the liliaceous plants were much more favorable for observation, and of these, owing to the ease with which the material could be obtained, Adium Cepa was the one generally used. I have satisfied myself, however, that the phenomena I shall describe occur in all of the plants men- tioned, and my conclusions are of course drawn from a study of all of these plants. In many particulars there is a notable differ- ence between the phenomena that may be seen in such tissues as the root tip and such cells as the pollen mother cells of the larch. These differences | shall attempt to correlate to some extent With the nature of the cells. METHODS. As some of the phenomena herein recorded are at variance with those observed by previous investigators, I have thought it worth while to give a detailed account of the methods used, even though they are those already well known to cytologists. Vari- ous killing and fixing methods were employed. Flemming’s chrom-osmo-acetic acid (both formulas); Hermann’s platinum chlorid-chrom-acetic acid; Vom Rath’s platinum chlorid-picro- esmo-acetic acid; Keiser’s mercuric chlorid-acetic acid; 96 per cent. alcohol, and a mixture of mercuric chlorid, formalin, acetic and formic acids arranged by Professor D. C. Worcester, of the department of zoology of the University of Michigan, and soon ‘© be published by him. Of these methods, the one best adapted to my purpose was the stronger solution of Flemming, 84 BOTANICAL GAZETTE [aucust since the kinoplasmic portions of the cell in division were nearly always well fixed in it. It proved in this respect to be muth superior to the weaker solution for the tissues studied. The material was kept in the killing fluid 30-48 hours, washed in running water, hardened by carrying through the different grades of alcohol and imbedded in paraffin in the usual way. Some | care was necessary to avoid plasmolysis in the early stages of hardening. I found that the objects could not be left in the lower grades of alcohol for a very long time; 30 minutes in I§ per cent., 45 minutes in 30 per cent., and one hour in 50 per cent. seemed to be about the maximum limits. Above 50 pet cent. more latitude as to time could be allowed. into the gentian-violet solution 20-25 minutes. They Wee © again rinsed in water and passed rapidly through a weak orange solution (1 part saturated solution orange G in H,O + 1 ae , H,0), dehydrated, clarified quickly in clove oil, and mounted # balsam. For Staining objects killed in the fluids containili mercuric chlorid, Zimmermann’s fuchsin iodine green® . Heidenhain’s haematoxylin, preceded by Bordeaux red. We ground stain, were found most serviceable. # Morph, u. Phys. d. pfl, Zelikernes 6, 1coo] THE CELL PLATE IN HIGHER PLANTS 85 In order to show clearly certain zonal differentiations in the spindle as well as other general structures in the cells I have used photo-micrographs for many of the illustrations. I have considered this the more satisfactory, inasmuch as my prepara- tions have showed some phenomena varying markedly from the descriptions of other observers. OBSERVATIONS. For purposes of description we may group the phenomena under three heads representing three stages in the development of the cell plate, viz., preparatory ; genetic; growing. 1. . Lhe preparatory stage. At the time when the chromosomes have collected at the equator to form the equatorial plate the spindle is of the form that Hof has recently termed ‘“ monaxial,” 7. ¢., a spindle having a single axis but not necessarily ending in single definite points as poles (figs. r, zo, 23). The poles of the spindle in both the larch and the onion are jn most cases somewhat blunt. In the latter plant my observations agree entirely in this respect with those of Hof (Z.c.) and Nemec (2. c.). In the case of the larch T have not confirmed Strasburger’s 3? observation of a centro- sphere. In my preparations the poles generally appear blunt as they do in the onion, The fibers which appear in connection with this stage may be grouped into three systems, connecting fibers, mantle fibers, and radiating fibers. These systems correspond in general to the system of fibers similarly named by older investigators, especially among the zoologists, except that here, as will appear later, the distinction cannot be so sharply drawn between con- necting fibers and radiating fibers. If we study the above mentioned Systems of fibers more in detail, it will appear that while Many of the connecting fibers are collected into thick | Strands or bundles, and others appear as single fibers, such an arrangement does not indicate a necessary difference of charac- eves frequently a single fiber diverges from a bundle and * Karyokinetische Probleme fig. 25. 86 BOTANICAL GAZETTE [ AUGUST unites with another bundle or continues as a single fiber to the pole. The arrangement into bundles seems to be produced } partly, at. least, by the fibers being crowded into the small spaces between the chromosomes. This explanation is rendered more probable by the fact that, as Strasburger has pointed out, the arrangement into bundles is much more evident at the equator than in the polar regions of the spindle (of. ciz., p. 183), 2.é., the fibers forming a single strand often seem to diverge beyond the equator. Some of my preparations show that this divergence reaches to the ends of the strand (fig. 23), so that such strands are most compact where they are adjacent to the chromosomés. The appearance of single fibers crossing obliquely from one bundle to another, together with the fact that such fibers often cross and recross each other, was taken by Belajeff as evidence that the fibers are drawn out portions of a protoplasmic net work. Guignard’s observation that the number of connecting fibers is equal to the number of chromosomes may be explained by the hypothesis that the fibers are collected into larger strands in the spaces between the chromosomes. Guignard, as previously | | stated, described such fibers as secondary fibers formed by Ls lateral fusion of previously existing smaller primary fibers. He did not, however, distinguish any other than the connecting fibers. Strasburger later showed that there are other fibers extending from the poles to the chromosomes. These are mantle fibers. They have been so clearly demonstrated recent by Osterhout, Mottier, Nemec, and others, that I need give detailed description of them in this connection. Suffice to say’ I find no apparent difference in structure between them and the a connecting fibers. : The third system of fibers to which I have referred may be described briefly as including those which extend from the poles into the cytoplasm. They are much more abundant in the lat than in the onion; hence I shall first describe them as the appear in the former. They seem to have the same structure the connecting or mantle fibers. Strasburger has figured them in the larch as radially arranged granules, while the conne¢ 1900] THE CELL PLATE IN. HIGHER PLANTS 87 fibers are shown as continuous lines. In my preparations the granular trophoplasm through which the fibers extend often has a radial appearance where the fibers are very abundant, but it is always possible in good preparations to distinguish the single threads (fig. 23). It is only by means of their distribution and arrangement that I would distinguish the radiating fibers from the other systems. They may be said to be centered in a gen- eral way on each pole, and to radiate through the cytoplasm in all directions toward the cell boundary, but at this stage very few appear to reach the plasma membrane. Their arrangement around the poles is by no means regular. Some of them lie across the poles in such a way as to have both ends free in the cytoplasm. With these it is of course often difficult to deter- mine whether a single fiber is seen, or whether the appearance is given by two fibers extending from the same point in opposite directions into the cytoplasm. In many cases, however, I have been able to satisfy myself that a single fiber extended through the pole and that its two ends lay free in the cytoplasm (fg. 23). This was most evident in cases where the fibers lay in such posi- tions as to form appreciable angles with the axis of the spindle. Where the apparent radiating fibers are continuous or nearly so with the long axis of the spindle it is often difficult to deter- mine whether they extend through the poles into the region of the spindle itself or whether they end at the poles. In some cases it appears as if such fibers are merely prolongations of the connecting or mantle fibers. I could not be certain on this point on account of the abundance of fibers in the polar regions of the spindle. Some, however, may be traced through the poles into the cytoplasm closely adjacent to the spindle. Still it can be readily observed that many of the radiating fibers which lie Closely adjacent to the spindle do not extend beyond the poles These latter fibers bear an interesting relation to the connect- Ing fibers, They often extend from the pole parallel to the connecting fibers, even uniting with the strands of the latter, nearly to the equator, where they curve outward and end blindly in the cytoplasm, Many of the fibers from each pole cross one 88 BOTANICAL GAZETTE | AUGUST another at the equator. These generally appear to be longerand more abundant than any of the other radiating fibers, but heir identity in appearance with the connecting fibers makes it impossible in many cases to determine whether they extend to the poles or not. From the above observations it seems to me to be improbable that there is any fundamental difference between the fibers of the three systems. While for convenience of discussion it is per haps best to retain the classification heretofore used, it is worth while to understand clearly just what significance is to be attached to the different terms. By connecting fibers I mean those which lie entirely in the spindle and extend across the equator. Mantle fibers are those which lie entirely within the spindle and aft attached to the chromosomes. ‘The spindle is understood, thet, to be made up of the mantle fibers and the connecting fibers Radiating fibers are those which, connected in small part with © the spindle, generally if not always at the poles have at least one end lying free in the cytoplasm. That this definition of radiating fibers may have to be slightly extended will appea later. s Whether there is any difference in origin of the various fibers : I have -not attempted to discover. The recent researches of Belajeff,33 Osterhout, Mottier, and others, have indicated that all firmed by later investigators.35 So far as my observations §% the onion radiating fibers seldom appear at this stage. Nemes however, has described them as first appearing in the very ™ : prophases and persisting through the metaphases. Theit tory, as Nemec describes it, is of much interest. The * Zur Kenntniss der Karyokinese be den Pflanzen. Flora 79: 439 1894 ‘“ i" ; ; Puts Kerne u. Kernkorperchen in meristematischen u. sporogenen Gewebem eltrage zur Biologie der Pflanzen 72225. 3 Hor, Joc. cit., and NEMEC, (oc. cif. 1900] THE CELL PLATE IN HIGHER PLANTS 89 indication of a spindle is an accumulation of hyaline substance around the nucleus, but in greater amount on the sides cor- responding to the poles of the future spindle. The whole body thus forms an ellipsoid spindle fundament (Az/age). This soon shows a filar structure and in many cases these fibers extend outward from the sides and cross in the equatorial region of the cell. The radiating fibers thus appear prior to the other fibers, or it would perhaps be more accurate to say that they grow more rapidly, as rudiments of the other fibers appear at the same stage in the polar regions. It should be noted here that no radiating fibers are described for these early stages, except those extending toward the equator. Such fibers, however, reach to the plasma membrane and are attached to it. The attachment is shown by a small accumulation of violet-staining material at the point of juncture. While Nemec states that the above mentioned fibers persist throughout the process of nuclear division to the begin- ning of the anaphases, his figures of the equatorial plate stage do not show them, but in connection with the late metaphases they are again figured. It seems to me probable that the two : Sets of fibers are not identical. It is not unlikely that those first described changed their position and became connecting or mantle fibers, while the later radiating fibers are a new growth. Nemec himself states that he believes that some new radiating fibers are formed during the late metaphases. I have been able to see some radiating fibers in the onion in connection with the €quatorial plate stage, but not in abundance nor of such length as Nemec described. They seldom reached beyond the “quator. When they were present they showed the same rela- tion to the other spindle fibers as has been described for the larch. MOS Lee dene of the cytoplasm in this stage requires a an ea, In the onion, it surrounds the spindle closely tt ie oe arabs be seen between the fibers in the polar region. it in Sieur slightly denser in the vicinity of the spindle than berry . cell-membrane. In older cells there are rrequent OUTER: the outer portion. In the larch an interesting dif- n frequently occurs. About midway between the cell go BOTANICAL GAZETTE [aucusT periphery and the spindle there is a thin layer of kinoplasmic ~ material, looking as if it were made up of small portions of fibers in a tangled mass (fig. 23). A similar appearance was described | by Mottier (of. cit., p. 180) in an earlier stage of spindle forma tion in the pollen mother cell of the lily. He observed, how ever, that this layer forms a part of the spindle, z.¢., it is the same as the felted layer described by Strasburger and Belajeff. That it forms no part of the existing spindle in the larch, howevel, — would be indicated by its late appearance after the spindle 8 ~ formed. It may be that there is in the larch more kinoplasm — formed than is needed for this stage in the development of the | spindle, and that it becomes absorbed into the cell protoplasm or is used directly for the later stages in the growth of the fibers (p. 47.) Between this kinoplasmic layer and the spindle there often appears a thick layer of finely granular trophoplasm which stains readily with the orange. A similar layer in the pollet | mother cells of Hermerocallis was described by Juel, but he | he did not mention any strong affinity for the orange stain, such as I have observed in the larch. In some of my own prepare | tions of Hermerocallis, I have observed the orange-stained layet F very frequently. ie Throughout the cytoplasm, and in some cases in the spindles also, there often appeared, in this as well as in later stages large blue-stained granules. Their distribution was irregulat: | They were more apparent usually in the larch than in the oni” ort mentioned and not as secondary structures formed simp between the groups of chromosomes. They show in the re other hand, the portions between the receding groups of daug res OE oe ONCE ERS OTD cE ERR ST rset eet PENT ee NED EAM SRST Set eS SRY Oe Ee eT eT TS ON a Te NERO Me ee ee 1900] THE CELL PLATE IN HIGHER PLANTS gl chromosomes often appear as single coarse fibers. This appear- ance is probably due to the closer crowding of the small fibers by the chromosomes. It is possible that in some cases this crowding together has gone to such an extent as to result in an actual lateral fusion of the fibers, such as Guignard suggested. The above-described condition of connecting fibers is doubtless what led Nemec to conclude that during the metaphases there were new coarse fibers formed reaching merely between the daughter chromosomes. I have been able to determine that in most cases the fibers can be traced through to the poles, and that they often appear to separate into smaller fibers beyond the chromo- somes in the way described by Strasburger for the larch. In the larch, in which the fibers show their compound character much more clearly, the true relation is still more distinctly seen (fig. 2). While I have not been able to confirm the observations of Nemec (op. cit., p- 329) as to the presence of secondary fibers during the metaphases which are formed merely between the groups of daughter chromosomes, I have often observed that the above-mentioned blue-stained granules appear to be very numerous in the region of the spindle. They often appear in rows and sometimes seem to be sticking to the connecting fibers. But such a regular arrangement is not at all character- istic. They are as frequently scattered in the ground substance of md protoplasm (fig. 25). That they are formed from dis- integrating secondary fibers seems impossible to believe when We consider the fact that they may be scattered throughout the whole cell as described above. As to the function or ultimate ~ of these granules I could learn nothing definite. They seem 0 si distinct from the ordinary granules of the trophoplasm in Staining qualities alone. The fact that they showed most plainly on the region of the spindle may simply mean that they are more pee distinguished by form from the spindle fibers, which in ag€ are not abundant, than they are from the other gran- ules appearing in the cell. eS eke 8 above mentioned processes have been taking place necting fibers, the radiating fibers and the cytoplasm g2 BOTANICAL GAZETTE [aucusi have undergone important changes. In the larch the radiating — fibers apparently grow longer, so that by the time the chromo somes have reached the poles a great many of them may be traced to the plasma membrane. Any evidence of such a fusion of the fibers and the plasma membrane as Nemec described in the onion I was unable to find. In the equatorial plate stage the fibers certainly did not reach to the plasma membrane. Whether the difference in appearance of the radiating fibers is really due to the growth of the existing fibers or to the forma tion of new ones, I could not determine. In the later stages, | shorter fibers may be seen, which are possibly the radiating fibers of the earlier stages, while the longer fibers are newly formed. The increase in number of fibers which such an explanation _ would demand was, however, not always evident in my prepata tions. In connection with changes in the radiating fibers there _ is a disappearance of the kinoplasmic layer heretofore described as lying between the spindle and the cell periphery. It may be, as previously suggested, that the substance is used in the growth | of the radiating fibers, though J have no positive evidence that such a relation exists. The fine granular zone also disappe@® | during the metaphases. There is now a tendency for the cyi® _ plasm around the poles of the spindle to assume a finely gram: lar appearance. The significance of such an appearance I have discuss in a subsequent paragraph. Following the formation of the diaster there sets in a5 of activities which are concerned immediately with the forain” of the cell plate. In the larch, the spindle soon appears to! 1900 ] THE CELL PLATE IN HIGHER PLANTS 93 differentiated into three zones (fig. 2), as Went first pointed out in other forms. But I cannot confirm the statement that the darker subnuclear zones take their characteristic appearance from a stainable ground substance between the spindle fibers. A care- ful study of the preparation from which fig. 2 was taken has convinced me that the darker portions are due to the structure and arrangement of the spindle fibers themselves. Some of the - fibers are undoubtedly thicker in this region than in the equator (fig. 27,a). With this differentiation in structure is combined the fact that the bundles have here begun to separate into single fibers (fig. 2). The two processes taken together account for the extra density observed. The above facts seem to me to indicate that the kinoplasmic activity preparatory to the forma- tion of the cel] plate begins in the region of the nuclei. The thickened appearance of the fibers soon extends throughout their length (figs. 3 and 27, 6). Concurrent with such thickening the Separation of the bundles into single fibers continues until the central spindle has the same appearance throughout. By com- paring figs. 2 and 3 it can be seen that nearly all of the connect- ing fibers have apparently shortened slightly, leaving clear Spaces just under the two daughter nuclei. In many cells there were often observed single connecting fibers in which the above described changes did not seem to be taking place, but they were never numerous in any one cell. Their distribution was not regular, They were as often seen in the central as in the more peripheral parts of the spindle. I shall have occasion to refer to them in the description of later stages. Granules of ‘rophoplasm may often be detected among the ends of the con- necting fibers at this Stage. These have probably flowed in as the fibers contracted. The process of separation of the fibers described above Sives the appearance of an increase in the number of spindle hp Whether such an increase actually occurs is doubtful. € larch the evidence, so far as I have observed it, seems to roti the apparent increase is due entirely to the above men- Processes. In this connection the hypothesis lately sug- 8ested by Strasburger, that the spindle fibers are increased in 94 BUTANICAL GAZETTE [ AUGUST numbers by splitting, is of interest. His evidence for the hypothe sis may be statedas follows: (1) the apparent rapid increase in the number of fibers; (2) the fibers produce a rapid outgrowth of the cell plate; (3) they are often found lying closely side by side. The first and third points seem to me to be readily explained by the facts that I have already described for the larch, viz., the rearrangement of the fibers by the bundles separating into single fibers, and the shortening and thickening of the fibers, resulting in a spindle of denser appearance, with apparently more numerous fibers. Strasburger himself points out that the spindle fibers seem to separate from the daughte! nuclei and to become thicker and more densely stained prior to the formation of a cell plate. It should be noted, however, that Strasburger had previously accepted Guignard’s doctrine thé! the connecting fibers are secondary structures formed by tht fusion of smaller primary fibers. The splitting of the fibers, it Strasburger’s sense, would be essentially the same as the sepat# tion of fibers, as described above, with this one exception: # the former case the process is unlimited, for new fibers may ©” tinuously arise by the splitting of original fibers, but in the latter the process is limited by the number of fibers making the bundles. If the splitting hypothesis were true, it W a explain, as Strasburger suggested, the appearance of 14 & peripheral connecting fibers, and thereby the growth of the cell plate; but below I shall describe phenomena which seem vo indicate that the appearance of new peripheral fibers depends not upon such a multiplication of the original connecting fibers, but upon changes in some of the radiating fibers. - Concurrent with the above described changes in the commer | the previously described radiating fibers is evident from 4 care , ful study of the preparations. In fig. 261 have drawn accuratel) : two such fibers extending one from each nucleus. Heré | j ; | q 1900] THE CELL PLATE IN HIGHER PLANTS 95 relation is such as to indicate that the two fibers have fused laterally throughout a part of their length. Whether such a fusion is real or only apparent, I could not determine A care- ful study of the preparation from which the photograph for jig. 2 was made shows all stages in the arrangement of radiating fibers, from that shown in fig. 26 to that in which the fibers extend out into the cytoplasm and cross at the equator in such a way as to form sharp angles (fig. 2g). The significance of these facts I shall discuss in a later connection. The radiating fibers that do not cross at the equator, so far as I could determine, have suffered no appreciable change. It is important to keep in mind that, with the exception of the presence of the trophoplas- mic granules, the changes in the appearance of the spindle which have so far taken place in the larch are due mainly to changes in the existing spindle fibers themselves, and not to the addition of new fibers or of other material. If we turn now to the onion, we note that there is an appar- ent increase in the number of connecting fibers (figs. 12, 73, 14). fig. 13 represents the same stage in the onion as that repre- sented by fig. 2 in the larch. The slight differentiation of the spindle into zones may be seen. While the smaller number of Spindle fibers renders such a differentiation less conspicuous, it is evident that it is caused in the same fashion as in the larch. A fact of importance to note here is that there are now visible more radiating fibers than could be seen in earlier stages. The photograph has not brought these out very clearly, but they may be seen by close inspection. These fibers radiate in all directions from the nuclei, but those are more abundant which “xtend toward the equator. In this connection it may be worth while to point out that these new fibers do not bear exactly the Same relation to the spindle as the radiating fibers that exist in the “quatorial plate stage. Those were centered not on the nuclei or chromosomes but upon the poles of the spindle. It is Possible, however, that they were originally centered upon the mother nucleus, as previously suggested. From a careful com- Parison of this with earlier stages, I am convinced that there 96 BOTANICAL GAZETTE | AUGUST has been a growth of new fibers. It is not impossible that some of these extend into the central spindle, thus increasing the appearance of zonal differentiation, but I could not establish this by actual observation. In an earlier paper Guignard figured such a relation but did not describe it. It seems tobe not improbable that such a condition may exist, and that such fibers may grow in length and form new connecting fibers, either by fibers from the opposite nuclei fusing, or by a com tinuous growth from one nucleus to the other. This process would account for the apparent increase in number of connect ing fibers. Still the evidence is too meager to lead to aly definite conclusion. There is no convincing evidence that there is any real increase in the number of connecting fibers. Its appearance may be due entirely to the changes which take place in the original fibers, as is the case in the larch. On the other hand, the relatively small number of the connecting fibers _ in earlier stages seems to show that some new ones have beet formed. The question needs further investigation. If there are new connecting fibers formed they seem to act simultaneously — with the original fibers in the process of forming the cell plate The point that seems of most importance to me here is the pre ! viously described relation of the radiating fibers to the daughter nuclei, 2. ¢., that they center not upon the poles of the spindle but upon the nuclei themselves. This relation, combined with the fact that the fibers are new formations, may indicate that | the nuclei are the metabolic centers for the formation of spi : f fibers. Such an hypothesis is further strengthened by the viously described changes in the connecting fibers in the laf where the increased thickness is first evident near the ends the fibers, 7. ¢., in those portions nearest the nuclei. The * that the appearance of the new fibers comes prior to the recom struction of the daughter nuclei does not invalidate the ab0Y hypothesis, for, as Juel has shown in the formation of abort pollen grains in Hemerocallis fulva, single chromosomes | have spindles formed between them and the normal nucle” Here would seem to be a case in which a chromosome 4S su : 4 4 4 ; : : : : ; j 1900] THE CELL PLATE IN HIGHER PLANTS 97 acts as a center for spindle formation. While the recent con- tributions on the subject of spindle formation in higher plants vary greatly in detail they all agree in one particular, viz., that the first indications of a spindle are to be found in connection with the nucleus and that the subsequent prophases take place while the fibers remain in this connection. These facts may be explained by the hypothesis that the nucleus is the center for the formation and activity of the kinoplasm. The further history of the equatorial zone in the onion pre- sents a striking difference from its history inthe larch. Whereas in the latter it disappears with the rearrangement and change in the connecting fibers prior to the formation of the cell plate, in the former it soon becomes filled with a substance that stains strongly with the orange of the triple stain. The spindle fibers retain their violet color in the same regions, showing that the orange stain is taken by a substance foreign to the fibers (figs. ‘4,15, 28). In appearance this substance is entirely homo- geneous. I could detect no appearance of granules or any other definite structural elements in it. It would seem to be a sub- Stance in solution in the living cell. It is a significant fact that the cell walls showed the orange stain in those preparations in which this orange stained substance is most evident in the spindle. In many Preparations in which there was a slight excess of Violet, the cell walls were stained blue and the substance in the €quatorial zone could scarcely be distinguished from the sur- rounding blue stained protoplasm. By the use of other stains it was often difficult to color or differentiate this substance. In one preparation, stained with eosin and methyl green, in which the methyl green was in excess, all parts of the cell were stained Steen except the equatorial zone and the cell walls, both of which showed a slight tinge of eosin. In all of the stains used when both the cell wall and the interfilar substance were stained, nets color was the same. There were some cases, however, in Which the walls Were stained while the equatorial zone remained c . : ; . ; Olorless, ¢. §., slides stained in ruthenium red or iron haema- toxylin, 98 BOTANICAL GAZETTE [AUGUST The similarity in staining of this substance to that of the cell walls, together with its presence in the region of the spindle in which the cell wall appears later, I have taken to signify the presence of a carbohydrate substance destined for the formation of the new cell wall. Whether it is at all analogous to the previously described layer around the spindle in the cytoplasm of the larch, I am not able to say. The fact may be of some significance in this connection, that in a late stage of division i the larch a similar orange layer appears in the cytoplasm arount each daughter nucleus. Possibly it shows the presence of material destined for the formation of the walls of the pollen grains. This fact may be correlated with the fact that the © division of the pollen mother cell often does not follow the fist ~ nuclear division. But in the stages showing the young perma nent cell plate, such a substance was generally invisible, thougi in some cells a slight indication of it was seen. There woul seem to be in these cells very slight or no aggregation of reserve cell-wall material. It is interesting to note that in the dividing pollen grains of Iris and Hemerocallis no orange stained inter filar substance was seen. Here, of course, there is no cell wal formed between the two cells. The relation of the spindle fibers to this orange substance i worthy of notice. As above mentioned, they retain their chat j | acteristic color, but they often appear greatly attenuated in bes region. It would seem that the substance had crowded thea F into such a condition (figs. 28, 29), and that it retarded tht previously described processes of thickening and separating © the fibers. | oospheres of Fucus. In the Saprolegniaceze Trow has sugs® that the so-called cellulin granules are a form of reserve cellul / which may be used to close the opening made by the breaking. 4 ofahypha. I have seen preparations of Saprolegnia in whit the young cell wall cutting off the sporange stains strongly ¥ BOTANICAL GAZETTE, XXX PLATE VIIJl BOTANICAL GAZETTE, XXX PLATE 1X TIMBERLAKE on CELL PLATE 1900 | THE CELL PLATE. IN HIGHER PLANTS 99 orange, and strongly orange stained granules are often apparently imbedded in the adjacent thick plasma membranes. The above facts seem to indicate that there may be in the protoplasm some form of reserve carbohydrate in readiness for the formation of a cell wall. I shall hereafter use the term carbohydrate material in speaking of the interfilar substance. The further history of this material is connected with the subsequent stages in the development of the cell plate. The essential difference thus far between the larch and the onion is that in the former the pro- cesses preparatory for cell division have been mainly carried on in the already existing fibers, while in the latter there has been a formation of new fibers and an aggregation of carbohydrate material in the equatorial region of the central spindle. UNIVERSITY OF WISCONSIN. (Zo be concluded.) CONTRIBUTIONS FROM THE CRYPTOGAMIC LABO- RATORY OF HARVARD UNIVERSITY. XLIV. NEW OR LITTLE KNOWN UNICELLULAR ALGZ&. I. CHLOROCYSTIS COHNII. GEORGE THOMAS Moore. (WITH PLATE x) EVER since the discovery by Cohn, in 1872, of the chlomw phyllous endophyte, Chlorochytrium Lemne, there has been consié: erable interest in algae having such a habit, and much speculatiot has been indulged in, both as to their affinities and the methol of Petrocelis cruenta J. Ag. He at first regarded these 100 [4 1900 ] CHLOROCYSTIS COHNIT Iof normal reproductive cells of the Petrocelis, but later came to consider them as something quite separate from this plant. These green cells growing with Petrocelis have been found in this country by Dr. Farlow (6), and recently Kuckuck (11) has decided the plant to be a Codiolum, having no distant connec- tion with its supposed host. After Chlorochytrium was described on Lemna a number of endophytic forms were discovered, some of which showed such marked resemblances to certain fungi that, had it not been for their green color, they would undoubtedly have been placed within that group. One of these “green parasites,” as they were popularly termed, was found by Wright (17) in 1876 grow- ing on various alge off the coast of Ireland, and called by him Chlorochytrium Cohnii. The discoverer of this form was so impressed with its fungus-like appearance and habit that he devoted considerable space to the discussion of how the plant Was in reality a fungus which had but recently acquired the Property of manufacturing chlorophyll. He was even able to observe the stages in this process as the plant developed. It is hot my intention to go into a discussion of how fungi and endo- Phytic algae are related to each other; I merely wish to describe One of the algal forms, and any comparisons to be made with the fungi must be left to another time. While collecting along the beach at Lynn, Mass., in Feb- mary 1897, my attention was attracted to the peculiar granular eesenee of some Enteromorpha which was growing attached to piles. When brought into the laboratory and examined under = See the alga was seen to be covered with a green wipe coe Organism which at first did not seem to have been pe cusly. described. Upon more careful examination and an ei orien of the literature upon the subject, it was Wri i nega plant must be the endophytic alga found by Sh : Chlorochytrium Cohnit. The material collected by me, this s ine ee fectly agree with any published account of Engle, "amd according to the keys in both De Toni (3) and oe Prantl (5) could not find a place within that genus. 102 BOTANICAL GAZETTE [ AUGUST It seemed necessary, therefore, to study the structure and devel: opment of the plant more carefully before it could be decided whether or not it really was a new genus. Since the plant was discovered by Wright, there have beet but three published accounts of this form, for, although tt occurs in widely separated regions and upon a number of dif ferent hosts, it is but rarely collected. Lagerheim (12) in 1884 found it off the coast of Sweden, and the next year Reinhardt (14) came upon it near Sebastopol, while investigating the flora of the Black Sea. The latter observer considered that the variations of the host plant seemed to bea more favorable resting pl” for the unicellular alga, and it was only occasionally th : : individual was found upon the larger more exposed see The only reason discernible for this was the fact that the Bp - omorpha was exposed to the air except at high tide, ant smaller fronds, growing in tufts and more closely adhering the piles, retained the moisture longer and were consequel more favorable for growth. ‘ When viewed with the ordinary low powers of a ™ scope, Chlorocystis appears a bright green color, usually exactly the same shade as the Enteromorpha cells, but ® easily mistaken for them. If the cells of the host pit dividing to form branch-like outgrowths and project above surrounding tissue, there is a slight resemblance to they" 1900] CHLOROCYSTIS COHNIT 103 Chlorocystis, but a careful examination will at once reveal the difference. In shape Chlorocystis Cohnti is usually spherical, although it may be slightly elliptical. It measures from 16-26 in the mature condition. Even though thé plants are frequently crowded together in irregular masses, they never lose their characteristic outline. The question as to the degree to which Chlorocystis may infest the host plant is one upon which my observations do not agree with those made previously. Both Lagerheim (12) and de Wildeman (4) describe the plant as being completely surrounded by the cells of the host, except for a small colorless portion which projects beyond the surface, through which the zoospores escape. Wright in his original description conveys the same idea, although he Says, ‘‘sometimes zoospores attach themselves in such quantities to Schizonema that there is no room to force themselves into the frond,” and at such times they are said to show but little evidence of penetrating the host. Chlorocystis Sarcophyci (16) is described as being completely embedded within the tissue of Sarcophycus. As may be seen from fig. 2, the plants as I found them were not always included within the host, but were quite as often merely attached to the surface of the Enteromorpha. While, as will be described later, the zoospores upon germination may send out processes which penetrate between the Enteromorpha cells and during further develop- ment may be more or less surrounded by these cells, the fact remains that many of the plants pass their entire existence with- Hive having at any time been within the tissues of the host. The crowded condition of which Wright speaks is not necessary to Pring this about, for the epiphytic habit is just as apt to occur ‘Mong single individuals entirely separated from one another, as ae they are grouped together. Even when the lower half of a rts cell is below the Enteromorpha (fig. 5, 9), a Siac — pressure will usually free it, leaving a round ees ers ee it has crowded the host cells apart. At no time ells observed completely covered by the Enteromorpha, 104 BOTANICAL GAZETTE [ AUGUST and the impossibility of such an occurrence is easily understool when we remember that the single layer of Enteromorpha cell is frequently less than half the size of the Chlorocystis, and that they could never any more than surround the alga im a vely superficial way. When the host is a plant made up of a massol tissue, it may be that Chlorocystis assumes a true endophyti habit, but in Enteromorpha it certainly does not seem possible. Occasionally zoospores get between the tubular frond of the hos loosening of the tissue, and finally complete disintegration of wise be due to this alga, ce Chlorocy stis contains a single large chromatophore, whi according to Reinhardt (14), and Wille in Engler and Prantl 1900] CHLOROCYSTIS COHNII 105 is said to lie always upon but one side of the cell. This is not true, however, and it was only after a considerable number of specimens were examined that the ‘one-sided’? chromatophore was observed. It so happened that all the specimens which were first found showed a chromatophore completely lining the cell wall, as indicated in jigs. 7, 2, and 4g. Later, however, exam- ples were found more nearly resembling Reinhardt’s figures (figs. 3, §a); but while this condition frequently occurs, it can no more be considered characteristic than when the whole cell is lined. A large and easily discernible pyrenoid lies near the surface of the cell and can be followed through all the subsequent divisions of the chromatophore (figs. 6, 7). Material killed in picric acid and stained for some time in 2 per cent. acid fuchsin brought. out the pyrenoids well, although Flemming’s fluid with. iron- alum-haematoxylin gave perhaps more satisfactory results. The chromatophore usually forms a definite dome-like thickening where it surrounds the pyrenoid, and this may extend into the cell in the way shown in fig. 5a. When the chromatophore does not entirely line the wall, it radiates from the pyrenoid in irregu- lar bands or ribbons, and these frequently do not pass more than half way round the cell (fig. 5a). This is the condition which Reinhardt figures and which he considered a generic character- istic. It was thought for a time that cells in which the chroma- tophore formed a complete lining might represent a condition subsequent to the formation of zoospores and not really be the adult Chlorocystis. Cultures in a Van Tieghem cell did not give much information on this subject, for although zoospores would be formed and escape, they did not develop to anywhere near Maturity. This was probably due to an insufficient supply of ©xygen, for when cultures were made in Ward cells, or simply see ae cover glasses kept in a moist chamber, the zoospores Hse snag from the time they escaped until they attained in. evelopment. By this means it was settled beyond a ilies at in the great majority of cases the chromatophore rely lined the cell from the beginning, and that it was a per- f . ectly normal arrangement throughout all the vegetative stages of the plant, 106 BOTANICAL GAZETTE [avcust Near the center of the cell is a well-defined nucleus, from which the protoplasm radiates in fine strands. There are usually several small non-contractile vacuoles present and the whole cell contents is often quite granular. Two sizes of zoospores are formed. The larger ones, which are spherical, measure 6-7 w in diameter (figs. zo, 72), while the smaller are only 2.6-3.5 « and are somewhat pyriform in outline ( figs. 11, 13). The method of the formation of the zoospores is identical in both cases, except that there are more successive divisions in the formation of the smaller spores, thus producing a greater number. In the original description of the genus by Wright, it is stated that the zoospores are formed in a very fet hours by free cell formation. By Reinhardt (14) this simulta neous formation is considered one of the points of distinction between Chlorocystis and Chlorocytrium. De Toni also use this distinction to separate the two genera. That the zoospores in Chlorocystis are formed by free cell formation is undoubtedly wrong, and all of my observations go to substantiate those of Lagerheim (12) and de Wildeman (4), who both state that the Spores are formed by successive divisions. All stages im this process may be observed (figs. 6, 7, 9), and nothing comparable to the description by Wright has ever been seen. Also the statement made by the discoverer of the genus that the eal 4 spores are at first colorless and that the protoplasm seemed project itself to one pole and there form a single cilium is o borne out by my observations. Both kinds of zoospores ha each four cilia, with a single chromatophore lining the base ® the cell. In the large zoospores the pyrenoid is easily madt out, and in the hyaline end of both the large and small spores ® | found a lenticular or spherical red spot. ; When the zoospores are fully formed and ready to escapes® | circular piece about 10 # in diameter is cut out from the top Zz outer side of the cell. This may be entirely loosened, or fre: ‘ quently it simply turns back, remaining attached at one - (Jig. IT) very much as in some of the Chytridinee. It see probable that the zoospores do not always escape in this manne : 1900 | CHLOROCYSTIS COHNII 107 for all the other observers of this plant have spoken of a color- less neck which projects beyond the surface of the cell and through which the zoospores make their way. Wright, however, has said to Miss Whitting (16) that when he found Chlorocystis Cohnii developed in the interior of tissue, the cells were some- times quite globular. Certainly the figures of Lagerheim (12) and Reinhardt (14) do not correspond to the description of a plant possessing such a protuberance. If the Chlorochytrium inclusum of Kjellmann (Q) is finally to find a place within the genus Chlorocystis, as has been suggested, we have still another example of a form without the neck-like protuberance. After the examination of dried specimens of Chlorochytrium inclusum Kjell. I am not inclined to think that it is a Chlorocystis ; although the published figures are strongly suggestive of that genus. It is certain that at no time, among the hundreds of specimens of Chlorocystis which were examined, was there anything that resembled a colorless protuberance. It may be that the varying habitat has something to do with the difference in aspect which this alga often presents; at any rate it seems probable that the presence or absence of a colorless tubular portion through which the zoospores may escape is not of much importance. hen the zoospores are liberated they swim about for a length of time varying from a few minutes to two hours. No difference was discernible in the rate or length of activity of the two kinds of spores. In almost every case the spores escaped perfectly free and independently of each other, but in a very few instances it appeared as though they might have been enclosed a delicate membrane as in Chlorochytrium. If there was such ag it must have been very frail and was suggested ofa es the arrangement of the zoospores than by any actual nat ee It always seemed to break up before any reagent a. € added to demonstrate it, and it is quite possible that bie = of the kind exists. Such a membrane enclosing the cae "es could not be of any significance from a systematic a Point, for even in forms where it occurs frequently, there Conditions which bring about its total disappearance. 108 _ BOTANICAL GAZETTE [ AUGUST Material in the laboratory showed the time for the escape of the zoospores to be usually from seven to ten o’clock in the mort- ing. This probably varies with the changing conditions at the seashore, and since the alga was submerged for only a few hours twice a day, it seems likely that the time of zoosport discharge varies with the tides. Efforts to establish this fact were unavailing. Observations made during the night wer _ likewise without result. Perfect aeration was found to be col ducive to the formation and discharge of large numbers of zoospores. The existence of two kinds of zoospores and the fact that conjugation takes place in certain closely related genera would naturally lead to the supposition that something of the sam kind occurs in Chlorocystis. De Wildeman (4) quotes Lager heim as having observed copulation, but I am unable 1 find such a statement in any of Lagerheim’s papers. He does mention having seen two kinds of zoospores, and considers | probable that the larger spore is formed by conjugation, but! think does not claim to have seen the process. From my ide observations I can say that it is certain the larger zoospores ae not formed by conjugation, and that it is possible for both sizes of zoospores to develop into new plants without any fusio® This point was carefully investigated by means of Van Tieght® cell and other cultures, and the zoospores were observed durilf their escape and final coming to rest. There was at no time ay appearance of conjugation, and the development of the spores whether of the large or small variety, was always the same, cells produced being similar in every particular to the charact istic adult plants. It may be that under different physiologi® conditions conjugation might occur, but at the present time ™ light can be thrown upon that point. When a zoospore comes to rest upon the surface of the hos plant, its cilia disappear and a thin gelatinous wall is formé around it. The red spot is lost to view and the Py™ becomes more prominent. If the zoospore is to develop ¥* the host instead of merely attaching itself to the surface, *’ 1900] CHLOROCYSTIS COHNIT 109 colorless neck is pushed out, and this penetrates the Entero- morpha frond between its cells and pushes them apart. When an entrance has been gained in this manner, the neck widens until the whole cell appears funnel-shaped, and this, after further growth, assumes its mature spherical condition. Ina few instances zoospores were observed which had germinated without having come in contact with the host-plant, and it is an interesting fact that some of these sent out colorless tubes of a considerable length (fig. 8). These were all found in cultures of various kinds and may have been due to some unknown abnormal condition. In the first published account of this plant, the zoospores were described as escaping from the mother cell without pos- sessing any color. These colorless zoospores developed into colorless plants which remained so until they had nearly reached adult size when the protoplasm commenced to develop ‘green cromules.” ‘These cromules,” says Wright (16), ‘‘arise as minute points along the inner surface of the cell wall from whence they radiate to the nucleus giving the appearance of a number of necklaces hung in loops.” Although I looked care- fully for some such condition in my material I was unable to observe anything abnormal or unusual. The green zoospores gradually developed into mature green plants with definite chromatophores as described. Resting spores were observed in material that had been kept in the laboratory for some time and had been allowed to dry "P partially. They are formed by the thickening of the wall of tie mature plant and the contents rolling itself into a solid mass of irregular outline, The spore thus becomes of a darker green shade, and the pyrenoid is lost in the increased density of the ceil contents. Be ei - seen from the foregoing that a number of points rae es to the structure and development of Chore cea, a have been considered by former investigators as ra deiaey can no longer remain as such. The habit of the si api and it certainly cannot be regarded as a uni- ophyte. The chromatophore is quite as apt to line 110 BOTANICAL GAZETTE [aveust | the cell wall as to be confined to one side and the radiate arrangement of the coloring matter may or may not occur. The method of zoospore formation is certainly the same as described by Klebs (10) for Chlorochytrium Lemne, namely by successive division, Even the manner in which the zoospores escape seems to vary, and the presence or absence of the colorless tube is of but little consequence. It may be questioned whether or not the material found a Lynn really was. Chlorocystis, since it fails to agree with any published account. Certainly much the easier way would be to - regard it as a new genus. But while the plants found do not agree with the keys in de Toni, and Engler and Prantl, or with any other published account, the points of resemblance are very marked when all the literature is considered as a whole and the various generic characteristics correlated. The few papers of the subject are strangely at variance, and the figures in at eee one case do not agree at all with the accompanying description nor with the specimens distributed by the author. Consequently it seems a case where we are justified in disregarding certall published accounts and in considering that the form above described is really what Wright and Reinhardt meant for the plant Chlorocystis Cohnii. DARTMOUTH COLLEGE, Hanover, N. H. BIBLIOGRAPHY. : eo t. COHN, F.: Ueber parasitische Algen. Beitrige zur Biologie 4. Pia 2:87-108. 1872. Bull. de la Sociéte Bel i : 144. 1G D. L : Die natiirlichen Pflanzenfamilien, Alge® a 1:65. 1897, : 6. FaRLow, W.G.: The marine algee of New England 115. Washingt 1881, . 7. HARVey, W. H.: Phycologia Brittanica 4: f/. 254. 1846-1851- 1900] CHLOROCYSTIS COHNII III 8. JESSEN, F. G.: Prasiole generis algarum monographia #/. 2, fig. 21. 1848 9. KJELLMAN, F. R.: The alge of the Arctic sea. Kongl. Svenska Vetensk. Akad. Handb. 20: 230-231. Stockholm, 1884. 10. KLeBs, G,: Bot. Zeit. 39:—. 11, KuCKUCK, P.: Wissenschaftliche Meeresuntersuchungen, N. S. 1:259. 1896. 12, LAGERHEIM, G. von: Om Chlorochytrium Cohnii Wright och dess fér- hallande till narstande arter. Svet. Vet. Akad. Ofvers. 7: 91-97. 1884, 13. METTENIUs, G.: Beitrage zur Botanik 1: 39. 1850. 14. REINHARDT, L.: Algologizieskya islidowanya. Contributiones ad mor- phologiam et systematicam algarum maris nigri. Odessa, 1885. 15. ROSENVINGE, L. K.: Les alges marines du Groenland. Ann. des Scien. Nat. Bot. VII. 19: 161. 1894. 16. WHITTING, F. C.: On Chlorocystis sarcophyci, a new endophytic alga. Phycological Memoirs. Ed. Geo. Murray, 7:—. 1893. 17. Wricut, P. E.: On a new species of parasitic alga belonging to the genus Chlorochytrium of Cohn. Transact. of the Royal Irish Acad. 26:355. 1877. EXPLANATION OF PLATE X. All the figures are from ink drawings sketched in with an Abbé camera. In the reproduction they are reduced one fourth. Figs. and 2 are drawn with SPOS toc. 5. All the others with a Leitz vy (oil), oc. 3. The mag- eet: given are the original ones before reduction and allow for projec- ion, Chlorocystis Cohnit (Wright) Reinhardt. Fi6. 1. eneral habit showing appearance of cells in Enteromorpha frond. x 280, FIG. 2. Section through Enteromorpha showing relation of Chlorocystis to its host. x 285. ae = ace view. of single cell. ‘“One-sided” chromatophore with biPe. and radiate arrangement. X 830. x rosy 4 Surface view of single cell with chromatophore lining entire wall. Fy . ; py : ’ Pies: 5. Side view ; @, showing arrangement of radiating “ one-sided’ tophore; 4; 200spore which has come to rest directly over Enteromorpha 112 a BOTANICAL GAZETTE cell; ¢, developing zoospore with projecting neck penetrating: Enteromorpha cells. x 830 Figs, 6, 7. Surface views eal first two stages in the formation of X 830. Fig. 8. Germinating zoospores in cultures not in contact with ho FIGs. 9, 10. Sporangia of large and small zoospores respective! F1G. 11. - Sporangium of large zoospores showing method of X 830. Figs. 12, 13. Large and small zoospores, X 830. Fig. 14. Resting spore. PLATE X MOORE on CHLOROCYSTIS a a EE A an eee eee Le TRS PT nT ae oN eR ee te TEN rm 1 BRIEFER ARTIGLES: NOTE ON THE MECHANICS OF THE SEED-BURYING AWNS OF STIPA AVENACEA. (WITH FIVE FIGURES) In the parts of many plants where hygroscopic movements take place in dead tissues, the cause is found in thick-walled mechanical cells of peculiar structure or varying chemical composition. The ripe and dry awn of Stipa avenacea, holding the seed at the lower end, is strongly twisted over half its length next to the seed ina direction Opposite to the movement of watch hands. The remainder of the awn, not having the spiral structure of the former, is not twisted, but is bent at an angle to the body of the awn, thus fur- nishing a brace or support when the seed begins its boring motion, driven by the alternate twisting and untwisting of the dry or wet awn. Little barbules on the upper part of the awn directed away from the seed, assist its progress forward while preventing any backward move- ment. The seed is tipped with a short, sharp point, slightly curved, the > ter to lead the way into the ground. Stiff hair-like barbs on the Wer portion of the seed hold it in the ground when once started. The onward motion is still farther assisted by the increased length of the awn when wet, which amounts, by actual measurement, to 20 per cent. of the whole length. On drying a corresponding withdrawal of the seed is prevented by the barbs. So the alternate wetting and dry- ing of the awn serves the twofold purpose of moving Se er (although this is, no doubt, more commonly accomplished by being fastened with its appendages to some moving object) and placing it in a favorable Position in t Th the lat Aveng barbata bet lo The moist awn straightens out completely. , potash or-other macerating fluid it twists with watch 113 Placed in caustic 1900] 114 BOTANICAL GAZETTE | aueus hands almost as strongly as it does in the opposite direction when dried. The minute structure of the awn furnishes an explanation of the phenomena here described. A thorough investigation of this subject was made by A. Zimmerman* whose purpose was to gain a more acti: rate insight into the torsion mechanism of the awns of wild grass He gives Hildebrand credit for first attempting to explain mechan ically the hygroscopic torsion. His explanation was considered incorrect, and was not in accordance with the views of Nageli and Schwendener, who held that the seat of the mechanism is in the ind: vidual cell. Francis Darwin afterward confirmed this view. Zimmer mann found that in the awn of Avena sterilis (with which, he sajs ” . a * fi ed the twisting power is connt ment. Both the arrangement of the pits in the walls of these cells and The present observations confirm those of others in locating es cause of the twisting of the awn in the individual cells and show ’ : not only a layer of cells but all of the mechanical cells are active im ~ ing about this result. . As is well known, the twisted portion of the awn is composed a cipally of sclerenchyma cells with a fibro-vascular bundle in the el and a band of chlorophyll bearing tissue on each side (/ig- 1). be latter, however, has nothing to do with the torsion. A_ striking pe larity of the mechanical cells is the narrowness and eccentricity of - lumina ; furthermore, this eccentricity in all the cells is alike, 8° = the lumina lie nearest the center of the awn (see figs. 7 and 3): a id Strong Schultz’s solution shows that the material imm around each lumen is stil] very much like cellulose; that it swells contracts more than the outer and denser layers of the cell wall 8 dent, since the surface of a cross section of a single cell . potash is convex, but when washed and dried is concave. * Jahrbiicher fiir wissenschaftl. Bot. 7+ 542. 1900] BRIEFER ARTICLES 115 As before stated, Zimmermann inferred the spiral arrangement of the material in the walls of the outer cells from the direction of the pits, and from the appearance in polarized light; but according to him no one had succeeded in bringing out clearly any striation. After treatment with strong caustic potash and then with dilute glycerine, I found such striations quite wel] marked, passing obliquely ARAB O AGN Alo lox oleisre Yor ao fo eS & gs Tosh ee 28 rae iy FIG. 1. Cross Section of half an awn showing the disposition of the mechanical - 2. An isolated cell (dry) showing the spiral arrangement of the material n.— Fic. 3. An optical longitudinal section of one of the cells (x) 2.— Fic. 4. One of the cells after soaking in a macerating agent.— deni ae am showing the resultant of forces that may cause torsion in the mechan- “~STGS. t, 3, and 4 are enlarged 115 times. on me cell (ig. 2). When seen through the cell in the opposite side. they pass in opposite direction to those in the wall on the nearer (more ae makes patent the spiral structure of the cell wall, denser dense, By refractive) layers alternating with layers which are less 116 BOTANICAL GAZETTE [ AUGUSI That the outer layer of cells is not, in this case at least, exclusively instrumental in effecting torsion, was proved by scraping off the outtr two thirds of a wet awn, when the remaining central portion, on dry: ing, was seen to twist as perfectly as the intact awn. Another evident of this fact is, that after maceration, the largest cells, which belong 10 the middle portion of the awn, are found twisted quite ‘as much as tht smaller outside cells (fig. g). This also agrees with Darwin’s obser tions, quoted by Zimmermann (/. ¢., p. 551). | There can be no doubt, then, that the mechanism is in the ind: vidual cells, and in the inner as well as the outer; and we have as # of cellulose-like material, the molecular structure of which is spiral. There are two causes present, either of which may under favorable © circumstances produce torsion: First, the mechanical cell may be considered as a hollow cylindet whose walls are made up of material in layers of alternating densi)” In the diagram (fg. 5) let fe and hg represent dense layers of matt rial while the less dense are the layers between. When dry, the cells twisted with watch hands. Water enters first into the less dense /#} ers forcing the micelle in all directions. ‘Two of these forces principally concerned here. The one, aé, acting at right angles © the spiral plane of more dense material, may be resolved into its es components ax and xd. One of these acts tangentially and wth simply to increase the diameter of the cell; the other moves the ee plane in such a way as to increase the angle it makes with the a8” the cell, producing, in the wall of the cell toward us, motion ai right to left. The other force, ac, may be resolved into its ee nents, ay and cy, the first of which again merely tends to increase i diameter of the cell, and the second, acting nearly parallel to the & - ponent ax, will strengthen it. In the opposite wall of the cell ® same forces will be found to produce the same result, but when o through the cell, the direction of motion will be just opposite + : in the wall on this side. There can be but one result from the : of such forces — the two forces on opposite sides of a cell, acting ” : Opposite directions about its center, will produce torsion. spot ) Second, the eccentric position of the cellulose-like material | < the lumen of the cell throws the center of the more dense materi one side of its axis, so that the dry cell on imbibing watet will 1900 ] BRIEFER ARTICLES 117 with the denser material on the concave side; at least this would ordi- narily take place. But such a bend in one plane is changed to a twist whenever the proper forces are present. In the case before us, we have not only the proper forces to cause a twist but also to give the motion constancy of direction, 7. ¢., with watch hands. ‘These forces are found in the spiral arrangement of the material. In this case we should expect a waving or serpentine bend rather than a close twist. The fact that many cells are found, after applying reagents, in all stages from a beginning bend in one plane to a wavy twist, leads to the con- clusion that this is, perhaps, the principal force of torsion when the lumen is very eccentric. It is probable that both these causes act in conjunction to pro- duce the generally resulting perfect torsion. — L. MUuRBACH, Centrat High Schooi, Detroit, Mich. SOME NEW SPECIES OF WYOMING PLANTS. Silene Tetonensis.— Stems several, somewhat cespitose from a mul- tcipital caudex, 10-25 high, 1—7-flowered: minutely pubescent throughout, glandular above and often throughout, the leaves often glabrous except on the margins: leaves connate at base and sheathing by somewhat scarious membranes, the petioles often sparsely ciliate ; the radical long-petioled, linear or narrowly oblanceolate, 2-8 long, 2~6™ wide; the cauline linear or the lowest pair narrowly oblanceo- late: calyx obovoid, 7-10" long, with ro purplish nerves, these anas- ‘omosing somewhat near the summit, 5-toothed, the teeth rotund or thomboid-triangular, obtuse with very broad membranous margins: nes Sedat long, greenish-white or rose-color, more or less exserted ; aien so broad, spatulate, with the margins entire or bluntly toothed neas the summit, not at all auricled, 3-nerved, the nerves branched and “nastomosing in the limb; this 3™ long, as broad as the claw or gen- aes a little narrower, with no lateral lobes, emarginate or cleft to the middle, the lobes entire and rounded, the appendages much broader than long and bluntly toothed : stamens nearly as long as the claws of the — the filaments glabrous; styles 3, 1™ long: carpophore very rt. Related to the western S. Watsoni, but is readily distinguished from that by j Y Its broader radical leaves and very different petals. 118 BOTANICAL GAZETTE [avcust The type is no. 6521 from the high grassy slopes of the Teton mountains, August 16, 1899; also no. 6684 from Dunraven peak in the Yellowstone park, August 27. Heuchera saxicola.—Scape, petioles, and inflorescence villous and viscid- glandular, the leaves sparsely so: leaves oval to rotund, truncaté or subcordate at base, doubly and incisely crenate with obtusish teeth, 2-4™ long, on petioles 4-9 long : scapes slender, 2-4" high, with two diminutive bract-like leaves near the summit : spike simple, often inter rupted below, 3-6™ long ; bracts ovate in outline, acuminate, 3-cleftt0 nearly entire and rhomboid, very ciliate on the margin, frequently tinged with violet, 6-3™ long: calyx campanulate, about 7” long, very pubescent and glandular, villous at base, divided to about tht middle ; the tube adnate to the ovaries for slightly over half its length; the lobes unequal in size, ovate or oval, obtuse, white and petaloid: petals wanting: filaments from triangular to subulate, 1™ or less in length: capsule ovoid : seeds oval, hispid, brown. | #1, saxicola is a near relative of H. ovalifolia Nutt., to which it is apparel) referred in the recent published Catalogue of the Flora of Montana, to? specimen, collected by Rydberg and Bessey in the Spanish basin, belongs ® this species. Referring to the original description, we find that H. ovalifola is “wholly destitute of villous hairs,” a negative character which at st Northwest America. As here characterized, it has lanceolate and Jaciniaté 2 bracts, while in ours they are ovate in outline and either rhomboid or 3-clelt The calyx is also described as tubular and with lanceolate lobes. In H, sat cola, on the other hand, the calyx is rather campanulate and the ee broader, on Type, no. 5687, Undine falls in Yellowstone park, July 6, 1899; also, 2822, Little goose creek cafion, July 17, 1896. | Saxifraga cognata.—Cespitose, the numerous short, erect oe (1-3™ long) forming dense cushions, the lower part of the stems ile ered with old leaves: leaves imbricated, oblong-linear to lanceol . cuspidate, 5~r1o™ long, hispid ciliate on the margins, otherwise glab rous: flowering stems slender, about 1° high, sparsely covered ¥ : short purple-tipped hairs, as also the pedicels of the flowers, beat several linear oblong leaves, these 3-5™" long and naked on the * gins, or nearly so: inflorescence cymose, 5~7-flowered, seldom 9° divided two thirds to three fourths; the lobes ovate, obtuse acutish, indistinctly 3-nerved, somewhat less than 2™” long: pe 1900] BRIEFER ARTICLES 119g elliptic, 4-5"" long, white, tinged with yellow toward the base, the upper half with purple dots, 3-nerved, the lateral ones arising near the base and extending to near the apex : ovaries united their whole length, the slender, conical styles more or less divergent. This includes all of the so-called S. dronchialis of the Rocky mountains, and perhaps also of the mountains of the Pacific states. 5S. bronchia/lis, whic is originally from Siberia, is everywhere described as having lanceolate sepals and orange-dotted petals, while our plant has ovate sepals, and petals quite prominently dotted with purple. No, 5551, from Golden gate in the Yellowstone park, June 28, 1899, may be taken as the type of this species. Ribes saximontanum.—A low shrub; spreading stems about 6% long, more or less bristly, with straw-colored and shreddy bark; infra-axillary spines three together, stout, 8—-12™" long: leaves orbicu- lar or broader, truncate at base, 3-lobed half way to the base, the two lateral lobes again somewhat 2-lobed, the lobes with obtuse or acutish teeth, very finely pubescent on both sides or glabrate, 6-20" broad : flowers 1-3, axillary, about 1™ long: calyx cylindraceous, glabrous, white tinged with violet: the tube 2™ broad or Jess, 4™™ long, villous within ; the lobes oblong, minutely toothed at the rounded summit, slightly shorter than the tube : petals cuneate-spatulate, toothed at the Toad summit, about 2™™ long: stamens included, the anthers oblong and obtuse : style divided half way to the base, villous: berry globose, smooth, reddish-purple, 6—10™" in diameter. z iba excellent species, quite different from its southern relative 7. /epfan- ”% trom which it is readily distinguished by its bristly stems, glabrous cane calyx, and divided villous style. It is an inhabitant of high, open ; ae ho. §542. Golden gate in Yellowstone park, June 28, 1899; also no. or Soteld peak, July 20, 1894, by Aven Nelson. Rosa grosse-serrata,— Much branched, 6-18 high, with occasional short prickles (less than 5™ long) or wholly unarmed, the branches eaves Somewhat glaucous ; stipules usually broad, 12-20%" long, acute, or a ie glandular-toothed, the free ends triangular ovate, cent Y Sean usually minutely resinous dotted and finely pubes- lalate sy 5-7-foliolate ; the leaflets nearly sessile, the terminal ase serrate See = aemended and cuneately obvate, sharply and coarsely as T about two thirds their length, 2-4 long, glabrous above, oy finely pubescent and minutely resinous dotted (under 120 BOTANICAL GAZETTE [AUGUST a strong lens) beneath as also the rachis: flowers in clusters of twott four at the ends of the branches or solitary: sepals entire, linear lanceolate, attenuated, the tips only slightly dilated, sparsely pubescett on the back and occasionally hispid-glandular along the margins, petals unknown ; fruit globose, perfectly smooth, akout 12™ in dia eter. From the latter one cannot so easily find characters by which to distingust it, yet it is so different in appearance that its separation as a distinct oie seems to be justifiable. The so-called R. Woodsii of the Rocky mountaits® a much lower plant, and has leaflets only 2°™ long. The following collections of this species are at hand: no 1191, Boulde! creek, August 27, 1894; no. 6787 (type), Madison river in the Yellowsia# park, August 30, 1899; both by Aven Nelson. Lupinus ramosus.— Stems several from a woody caudex, 2-4" hig branched, with divaricate branches, these simple and terminating " short few-flowered racemes: stems and petioles with two kinds pubescence, finely canescent and sparsely villous with spreading | leaves 5-8-foliate, the lower half the length of the petioles, the upp equaling them; leaflets narrowly oblanceolate, obtuse to acute, HE" mucronate, densely soft-silky on both sides, 2-4 long, 5-9” ™ racemes short-peduncled, 3-5™ in length, in fruit a little longer: bat ovate or lanceolate, about 2™™ long: flowers somewhat verticilatt " Scattered, about 1™ long; pedicels 2—3™" long, in fruit i 7 bracteolate, densely silky, as also the pedicels, the lower lip — longer than the upper, which is slightly notched: vexillum very” on the back, the central portion yellowish-white, otherwise pale ® oe lilac; wings pale blue, slightly longer than the vexillum; | light-colored, ciliate on the margin except at the very tip: P?™ ”— 3-5-seeded. : : ; Characterized by its branching habit, short and few-flowered race | the two kinds of pubescence. The type, no. 6576, is from dry banks and benches on Snake Bibi: lowstone park, August 20, 1899. LUPINUS HUMICOLA Tetonensis.—Stems one or more from a root, simple 3—6%™ high, including the racemes, 3-5-leaveds e acute or obtuse and mucronate, glabrous on the uppet me 1900] BRIEFER ARTICLES 121 sparsely strigose, 5-9" long, 8—18™" wide: raceme 4—-12™ long, mostly only 4-5°". No. 6341, Teton mountains, August 16, 1899. Antennaria fusca.— Loosely cespitose : stolons about 5 long: stems slender, 12™ high or less: leaves canescent or tomentulose on both sides, the older ones becoming glabrous ; the radical and those of the stolons spatulate, indistinctly mucronate, 15-22™ long; the cauline linear, 2~4"" wide, the lower somewhat broadened upwards and acute, the upper acuminate : heads 3-13, on short pedicels in close clusters or loosely corymbose, and the head or heads of the lowest pedicel, which is often 2-3 long, usually overtopping the rest: involucres about 6™™ high: bracts (pistillate) in about two series, the lower half bright green and sparsely woolly, the upper portion brown or greenish-brown, oblong, obtuse, more or less serrate. In habit and general appearance the species here described would sug- 8est A. aprica Greene, yet it is not even a very near ally of that. Its dark- colored bracts, slender stems, and the dull and light indument of the leaves easily separate it specifically from Professor Greene's species. The type is no, 6356, growing on dry bottoms and in open woods on Lewis river, Yellowstone park, August 8, 1899. tennaria oblancifolia.— Cespitose : stolons very short: stems slen- der, 15 high or less : radical leaves oblanceolate, those of the stolons Narrowly so, acute, mucronate, about 2™ long, sparsely canescent to glabrous above, canescently tomentose below: cauline leaves linear or oblong-linear, the lower acute, the upper acuminate: heads 4-13, in Close racemose or paniculate clusters : involucres (staminate) 4™ high, the herbaceous portion of the bracts sparsely woolly, the scarious por- tion oval, obtuse, brownish, or white. in ths near to A. racemosa, but it is so strikingly different from sr rs © and outline of its leaves that it must stand as a distinct It is represented Nate, s he Yel by a single collection in which all the plants are stami- Ls 08 AN open, once wooded slope, near Mammoth hot springs in lowstone park, July 3, 1899, no. 5640. ermnale,— Low, 1-2 high, branched from the base, it: ste ore or less branched throughout or simple to near the sum- i ase ce eaves grayish woolly: radical leaves oblanceolate, ae ai the cauline narrowly oblanceolate to linear, 1-3™ S* heads sessile in small glomerules terminating the branches: mi long, 122 BOTANICAL GAZETTE [avaus involucres 4-5"" high: bracts dull white, from ovate in the outer i linear in the inner, obtuse or acutish, nearly all apiculate. A northern ally of the Texan G. Wrightit. Collected on the geyser formations of Norris geyser basin in the Yellor stone park, July 25, 1899, no. 6139. Exias NELSON. UNIVERSITY OF Wyominc, Laramie SOME NEW NORTH AMERICAN MOSSES. (WITH PLATE XI) ald, some twenty miles north, where we pitched our permanent am ited only one of these, Sperry glacier, at the base of which we ® 4 several European mosses heretofore not reported for the United ae as well as some new species closely related to certain alpine species the old world. So far as determined the material collected 140 species of mosses and 20 species of hepatics. The publica a full report is delayed for various reasons, and it is deemed is to publish here only the most important part of the report as © ; Prepared. A more detailed account of this region, botanically # as geologically very interesting, may be found in the Septem” ber of Bulletin of the American Bureau of Geography. ae Dicranoweisia subcompacta Card. et Thér., sp. nova vinato-caespitosa. Caulis simplex vel parcissime ramosus, ete dense foliosus. Folia madida suberecta, sicca crispatula, I-15 oblongo-lanceolata, acuminata, subacuta vel obtusiuscula, “— canaliculata, nervo basi attenuato usque ad apicem product vel pa” lum infra evanido, marginibus inferne planis, superne inflexi rimis, cellulis irregulariter quadratis vel subrectangularibus, laxioribus, juxta costam linearibus, alaribus distinctis, subinflatis: . Caetera ignota.— Plate XT, 1900] BRIEFER ARTICLES 123 Very nearly allied to the European D. compacta Sch., from which it ae by the leaves being more narrowly acuminate and generally subacute, the cells of the areolation larger and with thinner walls, and chiefly by the costa narrower, attenuate below (16 to 25u broad; it is 554 in D. compacta).— Along the trail from Holzinger’s basin to the Rim. . Barbula rufipila Card. et Thér., sp. nova.— B. aciphyllae habity et foliorum forma omnino similis, differt tantum cellulis duplo majorbus et magis distinctis (superioribus 20-30m in B. rufipila, 12-15p in B. aciphylla) piloque saepius minus denticulata interdum integro. Spec- imina sterilia.— Plate XZ. Avalanche basin ; Holzinger’s basin. FISSIDENS BRYOIDES GYMNANDRUS (Buse) R. Ruthe.—New to North America. Cardot det.— Shores of lake McDonald ; Avalanche trail. Grimmia Holzingeri Card. et Thér., sp. nova.— Minima, tenella, pul- vinatula, obscure viridis, inferne fusca. Caulis erectus, 4-6" altus, parce ramosus, ramis interdum attenuatis, subflagellaceis. Folia ~ ferta, minima, o, 50-0.70™™ longa, 0.20—0.35 lata, madida erecta, sicca appressa, breviter ovato-oblonga, concava, omnia mutica obtuse ene: hata, marginibus planis integris, costa canaliculata, usque ad apicem Producta, basi 28 lata, cellulis superioribus _bistratosis, quadrato- subrotundatis, inferioribus unistratosis majoribus, lutescentibus, infimis. oblongis vel sublinearibus, omnibus incrassatis. Caetera ignota.— Plate XJ. This very minute species, resembling in habit the small forms of Andreea Petrophila, is quite distinct from all the European and North American species of Grimmia with muticous leaves by the small size, and the shape and areola- tion of the leaves,— Base of Sperry glacier; Mt. Trilby. GRIMMIA MOLLIS B.S.—This European alpine moss is reported from Greenland, and should be found at intermediate stations in Canada.— Base of Sperry glacier. Grins SUBSULCATA Limpr. in Rabenh. Cryptog. FI., Laubm. 7-—New to North America. Cardot det.— Mt. Trilby. : WERERA CARINATA (Brid.). (W. cucullata carinata Husnot ; — "@Viculare Cardot)— New to North America. Cardot det.— Base 0 Perry glacier, Bryuy A forma t 75 ALPINUM L., var. denticulatum Card. et Thér., n. ee * ii ? Ypica differt habitu graciliore, foliis ovato-acuminatis, 124 BOTANICAL GAZETTE [AUGUST brevioribus, marginibus parum revolutis, superne distincte sinualo denticulatis, costaque longe ab apice dissoluta——On the way from Holzinger’s basin to the Rim. PSEUDOLESKEA RADICOSA (Miill.) Lesq. & James.—This specits was distributed as P. rigescens Lindb.: it is the P. atrovirens of Eu pean authors. Best det— Holzinger’s basin ; Mt. Trilby. | PSEUDOLESKEA DENUDATA HOLziNGERI Best, in Bull. Torr. Bot Club 27: 229. May 1900.—Holzinger’s basin; Mt. Trilby; Avalancht basin. | Hypnum Cardoti Thér., sp. nova.—-Polygamum, olivaceo-viridé molle, laxiuscule depresso-caespitosum. Caulis procumbens vel astél dens, irregulariter ramosus, 2-4 longus. Folia remotiuscula, patulo | Squarrosa, interdum subsecunda, e basi constricta anguste decurreitt late ovato-deltoidea, subito in acumen angustum breviusculum recurvll | protracta, circa 1.5™ longa et 0.75 lata, marginibus planis fete undique sinuato-denticulatis, costa simplici bifurcata vel gemella, crus longiore ad medium producto, cellulis laxiusculis linearibus subflexuo™ basilaribus brevioribus et latioribus, alaribus laxis majoribus sub | hyalinis. Folia perichaetialia externa ovato-lanceolata, breviter ae nata, subintegra, enervia, intima plicata, costata. Capsula in pedice | rubente valde flexuoso, circa 18™™ longo, subhorizontalis, arcuata, ope culo convexo apiculato.— Plate X/. a This species is near H/, ste//atum Schreb. and H. polygamum Sch. ee the first it is at once distinguished by the polygamous inflorescence es softer leaves with a shorter acumen and a looser areolation. The sh oe the stem leaves and of the perichaetial leaves distinguishes it from the forms of the second species.— Avalanche basin. mee HYPNUM FLUITANS L., var. BR von Ren. in Husnot : a Gall., forma Holzingeri Ren.— Voisin de la var. brachydiclyon Ree i n’en différe que par le port plus gréle, la nervure plus étroite, © tissu délicat. Dioique 92! Cette var., essentiellement alpine, n’avait pas encore, jé crois, & nalée en Amérique. ee A cause de la briéveté des cellules médianes, on pourrait cone forme avec flypnum aduncum Hedw.; mais le passage brusque ye a) foliaires de la base aux cellules superficielles de la tige permet ce confusion.—- Base of Sperry glacier. _ Hypnum ocuraceum uncinatum Milde.— A European alp! new to North America. Renanld det. Holzinger’s basin. — ne mos BOTANICAL GAZETTE, XXX coe as ES Ey a. a re ———— r++ (gumecms © SSS —— eas. —r — Sa Ga er he ° CC ——— ae gw coe a es! Coo — = ee ergs é aSSa (ea 9) . realy 1900 | BRIEFER ARTICLES 125 HYPNUM UNCINATUM Hedw., var. suBJULACEUM Sch., forma Hol- zingeri Ren Forme voisine de la forme orthothecioides Lindb.; en différe par la couleur verte, les touffes compactes encombrées de terre 4 la base, l’acumen plus court denticulé et le tissu plus délicat, non €paissé.— Base of Sperry glacier. Minor extensions of range will be noted in a fuller report on this collection.— Joun M. Houzincer, Winona, Minn. EXPLANATION OF PLATE XI. (Nachet’s objectives 3 and 6, oculars 1 and 3, with camera lucida. All drawings are reduced % in photo-engraving.) 1. Dicranowetsia subcompacta. a, entire plant, nat. size ; 4, 4, 4, 6, leaves, X 32; ¢, basal areolation x 135; @, marginal areolation in the middle X 135; ¢, point of a leaf x 135. the same of B. aciphylla from a specimen of Styria X 285. ‘yp~num Cardots. a, entire plant, nat. size; 4, 6, 6, leaves X 32; ¢, marginal areolation in the middle X 285; d, capsule X 13. NOTES OF TRAVEL. III. RIO AND PETROPOLIS, BRAZIL. ‘oe : Professional traveler and to a botanist Rio de Janeiro and "polis have more to offer than any other easily accessible place in Ny America, Mr. Barbour Lathrop, with whom the writer has the ae traveling as assistant, is familiar with many of the pictur- ] © spots in the world, and even to him the region about Rio was a Most agreeable surprise. . Pit ies together in 1897 the harbor of Sydney, Australia, which id pepared with that of Rio, and were able to draw Sydne oe ich are decidedly favorable to Rio. row, gy arbor is long and, in comparison with that of Rio, nar- and which reat number of small coves separated by sharp points of of t jut out into the stream. These points of land, each side nate with the coves opposite, like the teeth of a . - harbor, alter ark. Th, ae ese low and rounded hilly points are covered with Australian 126 BOTANICAL GAZETTE [AUGust scrub, which is composed of narrow-leaved acacias, eucalyptus, aid numerous Myrtacee of a gray-green color. To those who tire 0 the gray-green of Italian olive groves, these narrow-leaved Myrtacet soon become monotonous, and the scanty shade shed by the sickle shaped vertical leaves of the eucalyptus makes little in the landscape that is restful to the eye. There are scarcely any islands in Sydne harbor, and its principal charm to a traveler lies in these innumerable coves which on either hand pop into view from the steamer. Theyat often filled with shipping, and the shores are being rapidly denuded 0! their forest and scrub vegetation, though many quiet pretty views tl remain. A botanist finds in the curious Proteacez, the most pecilli grass trees (Xanthorrhoea), and the endless variety of Myrtacee a host of forms which tickle his morphological sense with their novelty. Rio harbor, on the other hand, is like a large inland lake, wit numerous islands scattered through it, and surrounded by tall curious! rounded sugar-loaf hills. Islands, hills, and low-lying swampy shoe are covered with a wealth of tropical vegetation quite as luxuriant 8 any to be seen directly under the equator. In place of the round gray-green of the hills of Sydney, Rio has dark imposing cliffs whic reach above the low-lying clouds. Their deep ravines and valleys . a tangle of creepers, bright purple flowered melastomas, proad-l “oe lilies, and innumerable epiphytic bromelias. Everything is steamins with moisture and the leaf tips are dripping with dew. ov The islands in the harbor, though very picturesque, lack o cocoanut palms and coral reefs, two features which give to the B® of the south seas their peculiar charm. The former lack could 3 be remedied, but the brilliant white coral reef would be diff supply. : Rio is the most picturesque city in South America. The Portug® architecture is a great relief after the monotony of Spanish Ams cities, and no suburb in any city that I know is more charming” Botafogo, the residence portion of Rio. Each picturesque hee yellow or pink stucco, trimmed with colored Moorish tiles and with red tiling, is set in a half-neglected garden of tangled creepers, bamboos, ficus trees, and foliage plants. Although not taken care of as the American gardens of Honolulu, they are . numerous and picturesque. In fact, I do not believe there 30 in the world where such an array of picturesque gardens a _ can be seen as here in Rio. + = 1900] BRIEFER ARTICLES 127 The characteristic feature of Rio vegetation is made by the avenues of royal palms. Although there are avenues of this palm in Java, Hawaii, Jamaica, Trinidad, and Ceylon, in none are they really impressive. The first leaf-sheath in young plants is always objection- ably prominent. The immense avenues of Rio are so tall that these leaf-sheaths are not noticeable. No more beautiful avenues of palms are imaginable than those of the Botanic Gardens and a double avenue near Botafogo. The Botanic Garden lies an hour’s train ride from the center of the city, in a locality unfortunately infested with yellow fever. Its most courteous director, Dr. Rodrigues, has made a special study of palms, and his collections are very tastefully arranged through the garden. Although containing many rare specimens, this collection does not compare in any respect with that in the gardens of Java or Ceylon. Owing to its situation so far from comfortable hotels and its lack of laboratory facilities, the garden will be a difficult place in which to prosecute botanical studies. In the winter months the danger from yellow fever would quite prohibit its being used except for a few hours a day, as no stranger who values his life risks living in the city, but spends his nights at least either at Petropolis, three and one half hours away, or at Tijuaca, much nearer, but not so free from fever. ‘The charm of Rio, botanically, is in its surroundings. Petropolis, a city of twenty thousand inhabitants, lies at an altitude of three thou- sand feet among the mountains across the bay from Rio, and is reached by two hours of ferryboat and railway travel. During the season of yellow fever, from January to June, over two hundred passengers, mostly business men, make this trip twice a day, leaving Petropolis re morning and returning in the evening. As the danger from ee ee Spending a night in Rj ae p night in Rio during its summer season. - oe i es is richer is panoramic effect than any sna ne ees: 6 in the tropics, surpassing the famous railway see bie end Poet to Caracas in Venezuela, Columbo to Kandy in ra vegetation m Padang to Padang Pandjang in west Sumatra. e bali ‘es a) the shore of the harbor, through which the railway - that, given the degree of ionization of an aci and ! : a a S* KUSTER, ERNST: Beitrage zur Kenntniss der Gallenanatomie. Flora 87 3. 1900. b : ; : jum =° FkUR, ROH: The toxic action of a series of acids and of thelr sodi on Lupinus albus. Am, Jour. Sci. IV. 9: 183-192. 1900. * 1900] CURRENT LITERATURE 141 salt, the toxic action may be analyzed into the effect of the H ions, the anions, and the undissociated molecules if any. The latter play a particu- larly important part, especially in the fatty and aromatic series of acids. In general the H ions of inorganic acids are powerfully toxic; the anions of organic acids are slightly toxic, often negligibly so as compared with the H ions; and carboxyl H is many times more toxic than hydroxyl H.—C. R. B. Mr. F. H. KNow.Ton has brought together our knowledge of the fossil plants associated with the lavas of the Cascade range, and has published the result in the Twentieth Annual Report (Part III) of the United States Geological Survey, in connection with an account of the Bohemia mining tegion of western Oregon. The species number about thirty, including but three ferns and three gymnosperms, and are said to point unmistakably to the Miocene age of the beds.—J. M. C PROFESSOR W. A. KELLERMAN, of Ohio State University, and his wife have published an account of the non-indigenous flora of Ohio (University Bulletin, Botanical Series no. 4). Inastate known to contain 2025 seed- plants, it seems that 430 of them are not indigenous. These introduced forms are from the following sources : 326 from Europe, 30 from Asia, 2 from Africa, 46 from South and West United States, 21 from Tropical or South America. PROFESsor K. Miyake, of the imperial University, Tokyo, finds that starch is present in the leaves of evergreens in winter, and that it is due to feeble photosynthesis occurring during that season. The mean temperatures of various days when this process was determined varied from 0.7-7° (mostly less than 3°). R, B, ; Dr. G. N. Best has revised the North American species of Paeudoleahys: He recognizes Seven species, with four varieties, of which three are new. One species, P. falcicus pis Kindb., is excluded ; and one, P. atricha Kindb., 's doubtful—__¢, R, B. ** Bot. Mag., Tokyo, 14: 44. 1900. “Bull. Torr. Bot. Club 27: 221-236. pl. 6, 7. 1900. NEWS. Dr. ROLAND THAXTER sailed for Europe late in June for about tw months’ absence. Dr. B. M. Davis is in charge of the botanical department of the Marit Biological Laboratory at Woods Hole during the summer. . THE Fern Bulletin for July publishes a sketch of Daniel Cady Eaton) Professor Setchell, accompanied by an excellent portrait. Dr. B. L, RoBINSON is now in Europe, where he will spend much of the summer examining types of American species in European herbaria, In Rhodora for June there appears a sketch of Edwin Faxon by . | George G. Kennedy, accompanied by an unusually well-executed porta ; . icago, has Mr. A. C. Moore, a fellow in botany at the University of ei . been appointed professor of biology in South Carolina College, Co S. C., the state institution. 1 Dr. P. B. KENNEDY, of the Division of Agrostology, U. S. mani Agriculture, has been appointed associate professor of botany aNC culture in the University of Nevada. Mr. GEORGE P. CLINTON, instructor in botany, University of Die a leave of absence for the year Igoo-I, to study at Harvard. His pis study will be the Ustilaginez and’ their allies. ae Dr. WALTER Busse, docent in the University of Berlin, i a journey through the steppes of German East Africa in search of t0™ and medicinally valuable plants and plant products. me ie PROFESSOR GEORGE L. GOODALE sailed in May for ei sabbatical year, and he will be away during the coming college y' work will be done by Dr. R. H. True and Mr. E. W. Olive. a a ; ical ex Mr. M. L. FERNALD will spend two weeks of July in a hoe ee tion of Mt. Katahdin, Me., with J. F. Collins, of Brown aa J. R. Churchill, Dr. G. G. Kennedy, and Emile F. Williams, of Bost® Dr. H. C. Cowes conducted a class in field study of on the Tennessee mountains during June, He has been aie Cold Spt charge of the botanical work at the Biological Laboratory # c Harbor during the summer. . 3 - [ave we 142 1900] NEWS 143 Mr. C. F. HOTTEs, assistant in the botanical department of the University of Illinois, went two years ago to Bonn to equip himself for taking up ee specialty vegetable physiology. His work has proved so satisfactory that he intends staying another year. He will return to the University in Septem- ber Igol. A PROSPECTUs of an illustrated work on the maples, native and cultivated, in North America, has been issued by A. M. Lochman, Bethlehem, Pa. The work is to be by Charles L. Lochman, and is to have twenty-four suM: Page half-tone plates made from photographs of shoots and scat sa with forty- eight pages of text. PROFESSOR Dr. E. Loew (Berlin, S. W., Grossbeerenstr. 67) and Dr. Otto Appel (Charlottenburg, Schloss-str. 53), who have undertaken to pre- pare the third volume of Knuth’s Handbuch der Bluthenbiologie, appeal to all who have made biological observations on extra-European plants to communicate their papers or notes. MR. B. T. GaLtoway, chief of the division of vegetable physiology and pathology in the U. S. department of agriculture, was elected a vice-president of the A. A. A. S. and chairman of Section G (botany) for the Denver meet- ing. Professor A. S. Hitchcock, of the Kansas Agricultural College, was elected secretary of the section. BACTERIOLOGICAL EXAMINATIONS of the Chicago drainage waters are Constantly in progress in the University of Chicago and the University of Illinois, Samples from a score of places along the sanitary canal and the Illinois and Mississippi rivers are examined each week with a view of eh" mining the extent and kind of sewage contamination, and the influence o natural conditions in the purification of the stream. IT 18 EXPECTED that there will be no changes at present in the staff at the Missouri Botanical Garden and the Shaw School of Botany. During the summer Dr. H. von Schrenk will make a study of the diseases of forest trees, im the interest of the United States Department of Agriculture, probably Spending most of h H. F. Roberts Will pass the summ J. F. Collins, B Pennsylvania : : THE F ing fo: iversity of rown University ; Professor J. M. Macfarlane, University and Dr. A, B. Seymour, Harvard University. . st- OLLOWING NOTE, written in response to our circular letter serine Freview a Copy of the book named, may be of interest to botanists 144 BOTANICAL GAZETTE [ read reviews of it. They will at least know the price which the publ offer for “a good review” and the ‘‘terms”’ on which, presumably, the b is advertised by such “scientific magazines” as receive copies: DEAR Sir: Referring to your memorandum of July 23d, no copies of Melh One Thousand American Fungi have been or will be sent out for review. sent a few copies to the editors of iaedade magazines, allowing them a special count of 50 per cent. off the list price in consideration of a good review. Should care to review the book on these terms we will be glad to send you a copy for $6. Very truly beets E BowEN-MERRILL C0. On AuGusT FIRST Dr. C. E. Bessey returned to active work in t botanical department of the University of Nebraska. He celebrated ls resumption of botanical work by a visit to the Yellowstone Park for the pu pose of studying the vegetation of that region, especially certain grou lower plants. With his return to the department Dr. August Rimbé connection will shortly cease, as he was engaged in the department a Dr. Bessey’s year of service as chancellor. Professor John L. Shel the Nebraska State Normal School, has accepted a fellowship in Bais the university, and will return to study at the opening of the year. Dr. again for further ecological studiesyof the alpine and sub-alpine v continuing also his collection of specimens for distribution. THE PRIVATE HERBARIUM of Mr, Harry N. Patterson, of Oqua containing about 30,000 sheets, has been secured by the Field s Museum, and will be installed with the rapidly growing colle institution as promptly as the careful cataloguing practiced in 4 ments will admit. The botanical department of the museum > gratulated upon this accession of one of the notable private h to its already excellent representation of the flora of that Antillean islands. Mr. Patterson’s herbarium is more or less museum will be of great value to botanical students and specials Horsford’s Acid Phosphate The most efficient remedy known for the relief of languor and exhaustion, so common in the spring and summer months. Taken after exhaustive illness it acts as a wholesome tonic, giving renewed strength and vigor to the entire system. Taken before retiring, quiets the nerves and induces refreshing sleep. Sold by Druggists. Genuine bears name Horsror Sozodont in a new size pA of the Liquid The event of the year in dentifrices. getting the genuine at the stores. It necessary send 25c. direct to the Proprietors, P. O. Box New York City. BORATED TALCUM : er Bathing El ” anc 4 all af: Odor of 008 of P April 5, 1900. POL PLAN WAREROOMS: Fifth Avenue and 16th Street, Ne : 268 Wabash Avenue, Chicago. tf 181 Tremont Street, Boston . Familiarity Breeds contempt as its acquaintances will at ENOCH MORGAN’S SONS CO. : ; = AN’ S5 ONS ote i = Vol. XXX SEPTEMBER, 1900 Nes 3: THE BOTANICAL GAZETTE JOHN M. COULTER np CHARLES. R: BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS FRITZ NOLL é University aes Bonn VOLNEY rs SPALD ais i University of Michigan is ee & B. icc, ROLAND THAXTER : University of Padua Harvard Cniversity AboLF ENGLER WILLIAM TRELE ASE tok . Site of Berlin i ssourt ‘hoi Garden sii Lox SUIGNARD H. MARSHALL WA fe £. Pe de Pharmacie, Parts po te fet oF fetes idee : T A. HARPER r : EUGEN. W ARMIN ~ | Oniversity: of Wi tsconsin Ont ai of Cape dagen ae Tene VEIT. WITTROCK et ms rial University, Tokyo _ Academy of Scio, aoe 7S. eee A zy ue University CASIMIR _DrcaNDOLE a a CHICAGO, ILLINOIS. — Bublishey by the Gnibversity of Chicago Che Aniversity of Chicage press — pata 1900: BY THE UNIVERSITY OF CHICAGO fil Hi ATA SSS S= Pots A See ati Aa Ht HA Hy iW NE EE eS ee OAS WR aR EN RS OE) eT ee ES ee, SEN Botanical Gazette A Monthly Journal Embracing all Departments of Hotanical Science per year, $4.00 Single Numbers, 40 Cents 4 The subscription am must be paid in advance. No numbers are sent etl the expiration a of the time paid for. No reduction is made to dealers or agen FOREIGN AGENTS: _ Great Britain—Wm. WrEsLEY & Son, 28 Esse Germany — GEBRUDER BORNTRAEGER, Berlin St. Strand, London. 18 Shillings. SW. 46, Schonebergerstr. 17a. 18 Marks "Vol. XXX, No. 3 Issued September 15, 1900 CONTENTS . - ORIGIN OF THE CONES OF THE MULTIPOLAR SPINDLE IN GLADIOLUS Pain PLATE XII). Anstruther A. Lawson - 145 THE DEVELOPMENT AND FUNCTION OF THE CELL PLATE IN HIGHER PLANTS a (WITH PLATES viII AND Ix). H.G. Timberlake - 154 peeemtocicar “Hehe BoB easly ON SOME PERENNIAL HERBS (vir PLATE ds s A. R E 171 tition: FROM THE ROCKY MOUNTAIN HERBARIUM. I, Aven Nelson - 189 TEFER ARTICLES. PHOTOGRAPHY IN vba AND IN HorTICULTURE (WITH TWO FIGURES). /. 4. Waugh J. 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Fa a ge ~~ we eeeeaeea—e Se ee IA Eee eee eee NSN NAS ‘ Eo ts Insurance Company of America Strength With over $33,000,000 Asses and over $6,000,000 Surplis Value Excelling in usefulnes Over $42,000,000 paid! | Policy Holders. Merit Fulfilling its obligations the letter. | Simplicity in i licy Contracts” © Benekis ‘plain stated. Liberality Insured and B ve ecogeh Dividends a0 ditional Benefits. Fair Dealing ee to all —an established putation. dential n./ at 4 iJ HOME OFFICE: NE Pee Vt), WARK: eet Nes NS NS ee el Net ee! eed ell VOLUME XXX NUMBER 3 BOTANICAL (GAZEIee ; SEPTEMBER, 1900 ORIGIN OF THE CONES OF THE MULTIPOLAR SPIN- DLE IN GLADIOLUS. y ANSTRUTHER A. Lawson. (WITH PLATE XI!) RECENT investigation in cytology has gone far to establish the fact that multipolar spindles are of very general occurrence that the origin and development of the karyokinetic spindle in these plants is diametrically opposed to that which prevails in : the lower plants and animals. These observers believe that the Spindle, instead of being bipolar from the first and under the Control of a kinetic center, passes through a series of multipolar Stages in the course of its development. The problem has thus. come exceedingly interesting and important, and calls for vig- °rous and careful investigation. Multipolar spindles have been found in Lilium by Farmer (1893), in Larix by Belajeff (1894) and Strasburger (1896), in Equisetum by Osterhout (1897), in Lilium, Fritillaria, Helle- borus, Podophyllum, and Pinus by Mottier (1897, a and 4), in Hemerocallis by Juel (1897), in Chara by Debski (1897), in amia by Webber (1898), in Sagittaria by Schaffner (1898), in ymphaea and Nuphar by Guignard (1898), in Hesperaloe, Hedera, Disporum, Smilacina, Gladiolus, Iris, and Cobaea by Lawson (1898), in Passiflora by Williams (1899), in Solanum 7 Nemec (1899), and in Convallaria and Potamogeton by legand (1899). 145 : + Ju hapod ba ~ at %n the higher plants. It is indeed held by many investigators. rs 146 BOTANICAL GAZETTE [SEPTEMBER Occurring in such a wide range of forms, and having beet demonstrated by various methods of fixing and staining, it seems extremely improbable that the phenomenon is abnormal or due to artifact. Such a conclusion is much strengthened by the fact that in none of the higher plants has the animal type of spindle formation been discovered. While it is true that some investigators (Guignard, Schaffner, Fullmer, etc.) have described and figured granules situated at the poles of the spit dle, and thus in some respects resembling centrosomes, in 10_ case have such granules been observed to take an actual partif the formation of the achromatic figure as it occurs in animals Assuming then that the multipolar type is the normal type of spindle formation in the higher plants, since Strasburget | and other authorities cannot reconcile the function of a kineti¢ ; center with such a process, we are confronted with two problems namely, the manner in which the multipolar spindle become : bipolar, and the origin of the cones. No explanation has yet been offered that will reasonably — account for the ultimate bipolarity of the spindle beyond ait mere statement that certain of the cones fuse and arramge themselves in two groups. Such an explanation was first sug- ; gested by Belajeff (1894), and was later confirmed by Osterhout (1897), Mottier (1897), and others. How this fusion is broup® about is still uncertain. Even Guignard (1898), who descim centrosomes situated at the apices of the cones in Nymp and Nuphar, states that he is ‘‘unable to suggest, at present what manner the multipolar spindle becomes bipolar.” So = with or without centrosomes the phenomenon is unacce for, and we must therefore wait for the light of future investig® tion. On the origin and development of the cones much encouraging results have been obtained. The investien Belajeff (1894) and Strasburger (1896) on Larix, pee (1897) on Equisetum, Mottier (1897) on Lilium, Juel (1897)' Hemerocallis, and more recently those of Nemec on ©” and Allium, and Davis on Anthoceros, have thrown comsi®® 1900] MULTIPOLAR SPINDLE IN GLADIOLUS {47 light on the subject. But while these observers all agree that the spindle arises from a weft of kinoplasmic threads, very few observations have been made on the very earliest stages of the development of the cones ; probably on account of the fact that these early stages are very difficult to obtain. In 1898, however, the writer was fortunate enough to obtain a very complete series of these early stages in the pollen-mother cells of Cobaea scandens. In this case it was found that, as division approaches, the nucleus becomes surrounded by a zone of granular substance which dif- fers in structure and staining properties from the rest of the cytoplasm. This zone is so constant that it was proposed to des- ignate it perikaryoplasm, and as it has been found in several other genera, the writer feels justified in using the term in the Present paper. Upon the breaking down of the nuclear wall, the perikaryoplasm forms a central network of kinoplasmic fibers which grows out into several projections. By their growth cutward these projections become the cones of the multipolar figure. The Spindle fibers are therefore formed by the elon- gation of the meshes of the net-work in the direction of the pro- Jections, ; Encouraged by these results the writer was led to pursue his a of these early stages in other forms. His observa- ph on Gladiolus proved so interesting that it was thought advisable to record them, believing that they will throw further light on this much disputed problem. ; The method employed in preparing the material was practi- Pet hes Awe as that adopted in my work on Cobaes, namely : with Pi tip: solution of chromic-osmic-acetic acid, diluted stain haa ama of water, was used for fixing, and the triple would te ey enevigkel and orange G, for staining. I rial in the aie x emphasize the importance of fixing the seer Was only in th oe ae the material I have examined it being ree as anthers which were fixed immediately after Were found : 2 3 the plant that the early multipolar oo washed in ‘an ‘Alter being fixed the anthers were thoroughly ning water from six to eight hours, and were then 148 BOTANICAL GAZETTE [SEPTEMBER dehydrated by being passed through various grades of alcohol. Bergamot oil was used to precede the infiltration of paraffin Microtome sections from 3.6 u to 6 mu thick were used. THE POLLEN MOTHER CELLS OF GLADIOLUS.’ Gladiolus affords very exceptional material for the study of spindle formation on account of the large size of its anthers pollen mother cells, and nuclei. The cytoplasm in the resting pollen mother cell appears in the form of a clear uniform reti- culum, with numerous small spherical bodies scattered irregularly through it. The nucleus is very large, containing a vacuolated nucleolus, and the chromatin in a very characteristic spirem As division approaches, the cytoplasm undergoes a remarkable differentiation in identically the same manner that occurs at this stage in the corresponding mother cells of Cobaea. While the chromosomes are being formed, there gradually accumulates 2 © complete and sharply differentiated zone of granular substance about the nucleus. In nearly every respect this zone resembles the perikaryoplasm so characteristic of Cobaea. It accumulates in the same manner, has the same structure, and stains in the | same fashion. In fact, the only difference that could be detected / was in the size of the granules of which the zone is com In Gladiolus these granules are very fine, while those in Cobaea J are comparatively coarse. We shall therefore continue to use = term perikaryoplasm in the following description. By the time the chromatin thread breaks up and assumes has form of curved rod-shaped chromosomes, the perikaryoplas™ | reached its maximum development. In many cas® -— ere scar observed that the small black spherical bodies which w tered irregularly through the cytoplasm had arranged the can in the form of a ring at the outer margin of the perikary°P me But this peculiarity is not as constant or as striking as it Cobaea. ee At this early stage in Cobaea it was observed that t membrane breaks down, but in Gladiolus this does 9° : prid & *One of the common cultivated garden forms, probably of the hy avensis (Hort.). ies # t hap 1900] MULTIPOLAR SPINDLE IN GLADIOLUS 149 until a much later stage, as can be seen readily in figs. z-8. This fact is significant, because it is quite clear that the method of spindle development depends much upon when the nuclear mem- brane breaks down. This is very well illustrated by the two types which we have before us. In Cobaea, where the nuclear wall breaks down at a very early stage, we have a central net- work formed which occupies the space of the nuclear cavity. In Gladiolus, where the nuclear wall persists, the network is not formed in this central position, but is formed outside of and immediately surrounding the membrane. This difference also has its effect on the character of the network. When the nuclear wall breaks down at an early stage the nuclear cavity affords a large space for the formation of the network. Its meshes are consequently quite large. On the other hand, when the nuclear wall persists, as it does in Gladiolus, the network appears in the form of a close weft or felted zone. Very careful observations were made at this stage to see if etic kinoplasmic threads penetrate the nuclear membrane, as recently observed by Wiegand in Potamogeton. Although hundreds of cells were examined no such penetration was detected. : When this weft or felted zone commences to form, one might st frst think that it arose by the fraying out of the nuclear mem- rane. But as it grows to such a width ( fig. 7), and the nuclear mm apparently loses none of its distinctness ( jigs. 1-8), “sms much more probable that it grows at the expense of Perikaryoplasm as the central network does in Cobaea. a ihe ce oe of its rst appearance the felted zone stains meshes es with gentian-violet, and as it increases in size , which it Se larger and the sharp kinoplasmic threads : weft oo wey readily distinguished. As soon #8 ; e but pushes hen size ( fig. I iF it ceases to grow uniform a fat ak the oe into several projections as shown in /ig. Ee i Number of rh ite make out, there appears to be no de . Was the lar €s€ projections. fig. 2 shows at least five. Six est number observed in cross-section. Just how 150 BOTANICAL GAZETTE [ SEPTEMBER many there are it is impossible to say at present. These projec- tions rapidly increase, and on account of their growth outward the meshes become decidedly elongated. By the time they have reached their full growth, the whole outer portions of the cones are composed of long distinct fibers converging to the apices, and it is only at the base that the meshes of the original welt can be distinguished. The apices of the cones taper out inte remarkably sharp points, but in no case was there a body observed at these points which might be considered as a col trolling center. _During the entire process of the formation of the cones the nuclear membrane remains intact. It is only after they have reached their full development that it begins to break dows. The breaking down of the membrane is shown in figs. 6 and 7. It will be seen from these figures that this takes place on om side first, where the identity of the membrane becomes lost i the network at the base of the cones. It will also be observed from these figures that the nucleolus still persists. It remaifls quite conspicuous until all traces of the nuclear wall are me when it suddenly disappears. What eventually becomes of | was not observed. oe In my observations on Cobaea it was thought probable the linin of the nucleus took part in the formation of the ¢! , network from which the cones develop. Now in Sei where the kinoplasmic network and the cones are fully - > oped before the nuclear wall disappears, it becomes quite ee that the linin takes no essential part in the formation eS achromatic figure. ao Upon the disappearance of the nuclear membrane oa va of the cones soon adjust themselves to the space offered | nuclear Cavity, and in doing so come in direct contact, ik ite first time, with the chromosomes. These latter bodies aie : lacking in interest; but as the writer, in future work, h make a more detailed study of the chromatin in Gladic a other forms, his observations will not be recorded at a ; Soon after this stage a series was observed showing * — 1900] MULTIPOLAR SPINDLE IN GLADIOLUS 151 cones first approach each other and finally unite in two groups in the form of a bipolar spindle, as shown in fig. zo. From the above observations, and from previous work on other forms, it would seem that there are several types of spindle development in the higher plants. It would be imprudent to classify all of these probable types until the early stages of many of the forms have been more thoroughly investigated. -At present we have at least three forms which have been thoroughly worked out and which differ from one another sufficiently to warrant us in distinguishing them as types. These are repre- sented by Equisetum, Cobaea, and Gladiolus. SUMMARY. The above observations may be summarized as follows: As nuclear division approaches, a granular zone accumulates about the nucleus. This zone in every respect resembles the perikaryoplasm so characteristic of the pollen mother cells of obaea. A close network or felted zone of kinoplasmic fibers is formed immediately outside of and completely surrounding the nuclear wall. This js probably developed from the perikaryoplasm. This network grows out into several projections which become the cones of the multipolar figure. The nuclear membrane persists until the cones are almost fully developed. The spindle fibers are formed by the elongation of the Meshes of the network composing the cones. : Neither the nuclear wall, nucleolus, nor linin take any essen- tial part in the formation of the achromatic figure. The Cones of the multipolar figure fuse and arrange them- Selves in two Stoups and form a bipolar spindle. THE Universiry OF CALIFORNIA. 1894 LITERATURE. | 3 BELAJErr, W. Zur Kenntniss der Karyokinese bei den Pflanzen. seg Bone a AVIS,B.M. The spore mother cell of Anthoceros. Bor. GAZ. 28:89. 18972, - NEMEC, B. Ueber Kern- und Zelltheilung bei So/anum when BOTANICAL GAZETTE [seprensen DesskI, B. Beobachtungen iiber Kerntheilung bei Chara fragili, Jahrb. f. wiss. Bot. 30: 227. FARMER, J. B. On nuclear division in the pollen-mother cells of Lilium Martagon. Ann. of Bot. 7: 392. Ueber Kerntheilung in Lilium-Antheren besonders in Bemg auf die Centrosomenfrage. Flora 80: 56. FULLMER, E. L. Cell division in pine seedlings. Bor. GAZ, 26:23) ——— The development of the microsporangia and microsporess! Hemerocallis fulva. Bort. GAZ. 28:81. GuIGNARD, L, Centrosomes in plants. Bot, Gaz. 25: 158. — Les centres cinétiques chez les végétaux. Ann, des St.” Nat. Bot. VIII 5: 178. Juet, H. O. Die Kerntheilungen in den Pollenmutterzellen Hemerocallis fulva, und die bei denselben auftretenden Unrege! massigkeiten. Jahrb. f. wiss. Bot. 30: 205. Lawson, A. A. Some observations on the development of te karyokinetic spindle in the pollen-mother cells of Cobaea scandens. Proc. Cal. Acad. Sci. III. r-: 169. i Mortier, D, M. Ueber das Verhalten der Kerne bei der Entwidt lung des Embryosacks und die Vorginge bei der Befruchtung. Jahrb f. wiss. Bot. 31: 125. —— Beitrage zur Kenntniss der Kerntheilung in mutterzellen einiger Dicotylen und Monocotylen. Jahrb. 30: 169. den Pollet f, wiss. Bot Flora 86: 214. der Wurth Spindel bei Equisetum. Jahrb. f. wiss. Bot. 30: 159- SCHAFFNER, J. H. Karyokinesis in the root tips of Ad OT. GAZ, 26: 225, STRASBURGER, E. Karyokinetische Probleme. Os: 861, oe Ch | Jahrb. 2 wiss. Bok : Itheilung. Jah | f Ueber Cytoplasmastructuren, Kern- und Zel . wiss. Bot. 30: 375. entude Wesper, H. J. Povatlas structure occurring in the — Zamia. Bor. Gaz. 23: 453. ‘ot WEIGAND, K. M. The development of the mic microspores in Convallaria and Potamageton. Bot. GA? * WILLiAMs,C.L. The origin of the karyokinetic spindle es : cerulea. Proc. Cal. Acad. Sci. II. 1: 189. : ——— ried 7 ATI. aa PLATE we peti Fie ee Fe 8 3S ty Oe oe tO : ZEITE, F ep : BOTANICAL GA 1900] MULTIPOLAR SPINDLE IN GLADIOLUS 153 EXPLANATION OF PLATE XII. Figures drawn with Abbe’s camera lucida, Zeiss homog, immersion objec- tive one twelfth, apert. 1.25, compensating ocular no. 6. Fic. 1. A pollen mother cell showing the cytoplasm differentiated into three zones; the outer cytoplasm stains a light gray-blue ; the perikaryoplasm stains a light orange, and the weft or felted zone of kinoplasmic fibers stains blue; the nuclear wall is int dth ] contains a large nucleolus and several curved chromosomes, with a small amount of linin threads. Fig. 2. The weft of kinoplasmic threads is commencing to push out into several projections, preparatory to forming the cones of the multipolar figure ; the network of the weft appears more distinctly. Fic. 3. A still later stage in the development of the projection, with distinct cones formed. Fic. 4. A still later stage showing the elongation of the meshes and the formation of fibers by the pulling out of the network. Fic. 5. A slightly older stage than that of fig. 4. Fic. 6. A later stage when the cones are nearly fully developed; the Outer portions of the cones consist now of distinct fibers, and it is only at the base of the cones that the network of the original weft can be distinguished ; the nuclear wall has as yet shown no sign of breaking down, and the nucleolus still persists. 1G. 7. The first indication of the nuclear wall breaking down, The -_ Mosomes now come in contact with the base of the cones. Fig. 8. Shows the same as fig. 7, but a little more advanced. Fig. 9. The nuclear wall and the nucleolus have disappeared ; several ed the cones have fused, and the chromosomes are attached to the fibers at the base of the cones. FIG. 10. The mature spindle, with the chromosomes at the equator and ut to move to the poles. abo THE DEVELOPMENT AND FUNCTION OF THE CELL PLATE IN HIGHER PLANTS. we H. G. TIMBERLAKE. (WITH PLATES VIII AND IX) (Concluded from p. 99) 2. the genetic stage. THE origin of the cell plate elements occurs in the equator of the central spindle. In the onion this brings them in the midst of the carbohydrate zone (figs. 18, 19). There seems to bee. : doubt that they are thickenings of the spindle fibers. In the onion, while they are often very difficult to distinguish at first, owing to the fineness of the fibers and the abundance of cattt hydrate material, it can be determined clearly that they ae swellings of the fibers (fig. 29). There is nothing to sugges the movement of cytoplasmic granules toward the equa” — plane to form the cell plate in the manner described by Tresh Working on living cells, Treub might easily have failed or the beginning of the cell plate. His statement that it appears as a fine line would indicate this. The same © tion would apply to the observations of Zacharias, although ; to be noted that Zacharias described special bodies as ell PY elements and not mere undifferentiated cytoplasmic 8” such as those of Treub. It is to be doubted, however © vu these bodies have any connection with a cell plate. poe think it unreasonable to suppose that what Zacharias sy substance destined for the formation of the cell wall ™m shown more clearly in the larch. Here the thicket much more pronounced, being elongated bodies insta” * 3 Cf. FARMER, p. 78 of this paper. 154 1900 } fe CELL PLATE IN HIGHER PLANTS 155 nodules (figs. 4, 27¢). In this case it looks as if the activity of a fiber previously described as beginning near either daughter nucleus and going toward the equator, has become localized at the latter point, producing a swelling on the fiber. The pro- duction of such a swelling is accompanied by the further short- ening of the fiber (fig. 27c), showing that there has been an actual transformation of the substance of the fiber in the for- mation of the cell plate element. While the above described process seems to agree with the older observations of Stras- burger that the substance of the cell plate elements has flowed in the fibers to the equator,37 it should be noted that there " nothing in the process I have described to indicate a collec- tion of smaller granules within the fiber, but that the fiber itself has changed... The whole process seems to indicate a somewhat plastic character of the fiber. The time relative to other phases of mitosis at which the cell plate elements occur seems to vary in the onion and the larch. In the former their appearance is concurrent with or closely fol- lowing that of the carbohydrate material. The chromosomes have begun at this time to form slightly denser masses, prepara- ‘ory to the reconstruction of the daughter nuclei, but separate chromosomes may still be distinguished. In the larch, the Process of reconstruction goes so far before the cell plate ele- rpg appear that the daughter nuclei are often clearly outlined a = 4). Whether this difference has any special signifi- Sais Si Sealed ries It would be interesting to compare a wider lety of forms in this respect. : a as the elements can be detected, they seem to form cating rie the thickness of the original spindle, indi- stages ma i ey are emultancous in their origin. Very early ful isch ave escaped observation, however, although Ae tise tight i made for them. That such a progressive forma- fend & wi the case would be suggested by such cue cell plate ele ere there are some peripheral fibers on whicl r ments have not yet formed. On the other hand, it &. p. 75 of this paper, ‘ 156 BOTANICAL GAZETTE [SEPTEMBER is probable that all the fibers showing cell plate elements in jig 4 are the original connecting fibers, while the peripheral fibers are the previously described radiating fibers. I have previously shown that the changes in the connecting fibers, before the for mation of the cell plate elements take place in all of the fibes at the same time, while the radiating fibers have apparentlys changed as to form peripheral connecting fibers. Whether all of the connecting fibers form cell plate elements was a poitl that I could not settle with certainty. It is possible that tho connecting fibers which did not show the changes described i® the preliminary stages do not form cell plate elements. The phenomena observed in later stages render such a possi ility more probable. | After they first become visible, the cell plate elements tinue to enlarge in their equatorial diameter until they come into contact with one another and fuse into a continuous plate. This process is accompanied by the further shortening of the fibers In connection with the shortening of the spindle fibers there a continued appearance of trophoplasm in the terminal parts the spindle (figs. 4, 5). ee The result of the processes just described may be brief summed up as follows. There is in the spindle a youn . plate formed simultaneously from the substance of the spi 4 fibers. In the onion it lies in the midst of a zone of resent carbohydrate material to be used in the formation of the ae cellulose wall. In the larch such a zone is not seen : ol stage. The cell plate may now be said to begin the next st in its development. a 3. The growing stage. poe mace This stage is marked by the following phenomen : central fibers continue to shorten, adding their substance net! cell plate until they have finally disappeared. The per ' fibers begin to appear more and more bent and to fo plate elements. As the central fibers disappear their eg partly taken by granular trophoplasm and the daught® come to lie nearer the young cell plate. While the Per 1900] THE CELL PLATE IN HIGHER PLANTS 157 growth is continuing, the older portion of the cell plate splits and the new wall is laid down between the halves. These processes continue until division is complete. During the period of growth the cell plate may so shift its position as to lie ina plane different from that in which it was first formed. I have thought it best to describe the above processes sepa- rately. The fate of the spindle fibers is a question of much interest here. From the phenomena that I have been able to: observe in the cells studied, I have become convinced that all of the fibers that form cell plate elements are completely used up in the growth of the cell plate. A comparison of figs. 4, 5+ 6,and 7 is instructive on this point. In fig. g the cell plate elements have just formed, the connecting fibers have drawn away from the daughter nuclei, and granular trophoplasm has appeared among the ends of the fibers of the central spindle. In fig. 5 the cell plate elements have fused into a cell plate, the fibers have shortened still further, and the trophoplasm appears. Eee Pigs is especially important. Here the fibers have become very short and the ends furthest away from the cell plates are no longer surrounded by the granular trophoplasm. These ends appear to be sharp pointed, while the portion of each fiber that is in connection with the cell plate is relatively thick. i stage the trophoplasm becomes distributed more oo the region originally occupied by the central spindle. se it oy appears in irregular rows of granules ( fig. 7)» ma % atter figure very small portions of the spindle fibers Pe oa They show the typical pointed structure above as and are distinct from the rows of trophoplasm. the ai ae all of ane processes involved in the formar of Strong] ck. the spindle fibers and the cell plate itself stain the Re the “iolet of the triple stain. I have not observed described transition from the violet to the orange color when th y Strasburger for the spindle fibers at the time € cell plate is formed. Not only do the spindle fibers. and the cell plate stain violet, but the plasma membrane of the: *Zelihinte. 158 BOTANICAL GAZETTE | SEPTEMBER mother cell frequently shows the same color. To be sure some preparations show the orange color in connection with the cell plate, but the violet of the cell plate itself could nearly always be distinguished. The orange color is probably due to the small amount of carbohydrate material that sometimes appeats at this stage. I do not think that the presence of the above mentioned irregular rows of granular trophoplasm is to be take as evidence that the spindle fibers break up into granular cyt plasm. My preparations show clearly that the spindle fibers a entirely distinct from these rows. The history of the fibers seems to consist of a gradual shortening and thickening until their whole substance is transformed into a cell plate, #.é@ membrane. While the above description will apply to thos fibers which take part in the formation of the cell plate, the fate of those radiating fibers which have no part in this process remains unaccounted for. My observations upon this latte point have been too limited to base any conclusions upon them. In a very early stage in the formation of the cell plate these fibers often seem to lose their characteristic radial arrangemet ie and to become more of a tangled mass in the cytoplasm (fs: : 3, 4). It is possible that they become separated from “ daughter nuclei and are finally absorbed into the rest if ‘ protoplasm. There is one other interesting case to be 20" In some ceils of the larch and some of the cells of the root of Fritillaria, after the cell plate was formed, a few clearly fibers were observed in the region between each daugi nucleus and the cell plate. These fibers are probably the ee that have been mentioned previously as not showing the ge Preparatory to the formation of the cell plate. Their wee not determined. Fig. zo is interesting in this conse represents a late stage in the division of the pollen mother © : when the permanent cell plates are complete, but ease still exist numerous fibers all around each nucleus, ome into the cytoplasm and reaching in many cases to thet : membrane, Whether these are a part of the connecting * radiating fibers that existed during the earlier stage eae Pe tire ier ay ante se teat Pe ee ae Ns iy San ea oO SRE PEE! SCAG 4 ete i Sea a ap eer av ae eae eee aS 1900] LHE CELL PLATE IN HIGHER PLANTS 159 fibers that have been formed after the division was complete, I did not determine. If the former supposition is true there would seem to be an unusually large number of spindle fibers which took no part in the formation of the cell plate. From the fact that in all cases observed of the formation of the first cell plate no such abundance of regularly arranged fibers was evident, I think it probable that these are mostly new fibers. Connecting stages, however, may have been overlooked. Their significance in either case is not clear. It may be that they build a new plasma membrane around each pollen grain inside of the plasma membrane of the special mother cell. Further investigation upon these late stages of the formation of the pollen grains is desirable, The process by which the cell plate grows in area is difficult te ee with certainty. By comparing stages represented in figs. 4-7, 1 have concluded that a part of the growth of - cell plate takes place by means of the continued addition to tof the substance of the original fibers until they are entirely used up. I have already described what seems to be the process ihc which the fibers go in the formation of the cell plate. eRe the evidence for the complete transformation of at ae a of the fibers into the cell plate conclusive. The most the 2 € interpretation of fig. 6 seems to me to be that all of ing s Soha fibers here shown are the remains of the connect- aa Tes shown in fig. g. This interpretation is further 5). hae by an examination of an intermediate stage ( fig. are in oh ngs fibers are not much shorter relatively than they while ea 4, but it ws noteworthy that they are further apart, facts gah plate is, of course, more conspicuous. These coliectin O the conclusion that the substance of the fibers, apart, a ie the middle, has simply pushed the fibers further While a ee that constitutes the growth of the cell plate. tion of the simple mechanical process may account for a por- Which seem wth of the cell plate, there are phenomena all of the ee indicate that other processes accompany “ in € figures, while there is a large area throughout in | 160 BOTANICAL GAZETTE [SEPTEMBER which the cell plate is in the same stage of development, it will be noted that the peripheral portions show earlier stages, and that there are fibers connected with this latter portion whose sub- stance has not yet been used in the formation of the cell plate In many cells these fibers show all gradations from very short ones toward. the center of the cell to those reaching nearly, i not quite, to the daughter nuclei (fig. 30). Some of the latter, instead of extending from one nucleus to the other, extend simply from the nucleus into the cytoplasm of the opposite daughter cell. These facts would indicate that these peripheral fibers are radiating fibers which are taking part in the formation of the cell plate. The fact that the peripheral fibers appear! be more numerous and in a more compact layer in the stage represented by figs. 6 and 7 than in earlier stages (jig: 5) may be explained by the crowding produced by the growth of the . cell plate by the transformation of the inner fibers in the fashio® described above, 7. ¢., the peripheral fibers in jig. 6 would repr . sent all of the radiating fibers between the boundary of the ce 4 plate in this figure and its boundary when it had reached a stag such as that shown in fig. 5. The appearance of shorter - on the inside of the peripheral bundles and longer ones of a outside shows that the process of cell plate formation goes 0 gradually in these fibers from the inside outward. This is ee farther by the fact that the cell plate elements of the Ss fibers have already fused into a cell plate, while the interme ones are often distinct, and the outer fibers often show ne plate elements (fig. 30). The foregoing observations see” show that the increase in area of the cell plate is due to @ fold process consisting (1) of a continuous transforma! the substance of the original connecting fibers, Tes expansion of the portion of the plate around each fiber i consequent wider separation of the fibers; (2) concltt the above method occurs the addition of new cell plate or to the peripheral portions of the cell plate, and theif ee! uit the same fashion as those previously formed. These” processes go on in the radiating fibers which have ©? 1900] THe CELL PLATE IN HIGHER FLANIS 161 such a relation to the central spindle as to continue the first process of growth. Another question of importance in this connection is, does the growth in area of the cell plate depend entirely upon the fibers already existing, or are new peripheral fibers formed f So far as my observations upon the larch go, I think it probable that there is no formation of new fibers, but that the whole growth takes place as a result of the changes in the existing fibers; yet the evidence on this point was not conclusive. If the hypothesis is true, whether the mother cell divides into the four cells which form the pollen grains by successive or by simultaneous division depends upon the number of spindle fibers existing in connec- tion with the first nuclear divisicn. If there are enough fibers " form a cell plate completely across the cell, successive divi- sion results, while if the cell plate does not reach across the = it is absorbed into the rest of the protoplasm, and the final division takes place simultaneously after the second nuclear division. In the onion the conditions are such as to indicate the necessity for the formation of new peripheral fibers. It will be remembered that at the time of the formation of the young “cil plate there were but few radiating fibers visible, and those that could be seen were relatively short, with the exception of sr Ay the periphery of the central spindle. In the later es Pe ter the fibers had entirely disappeared from the central tion ok .. relations of spindle fibers in the peripheral por- Viously ste could be distinguished that have bese pre- that there Sun ed in the larch (fig. 30). This relation indicates ing fibers sage a growth at least of the original radiat- There: ig s sa probable production of new ones. nother point of importance here. The fact that peripheral fibers may be traced to the nucleus uous production of new peripheral fibers from and the onion ; . The difference, then, between the larch '0n is possibly as follows: in the former the extent ate depends upon the number of exist- latter, where there is necessity for the 162 BOTANICAL GAZETTE [SEPTEMBER formation of a complete cell plate, the original fibers mayb supplemented by the growth of new peripheral fibers. The cases in the larch in which the cell plate is not completed after the first nuclear division are possibly accounted for by the fact tha! the nucleus has begun to prepare for its second division, até consequently new fibers necessary for the completion of the cel plate are not produced. Concurrent with the growth of the cell plate in extentai the disappearance of the fibers from a central portion of the spindle the nuclei come to lie nearer the cell plate (figs. 4 I} This phenomenon has often been observed, but no adequalt | explanation for it has been suggested. From the appearance the whole cell in the larch it would seem that such a migratie? of the nuclei is due to the mechanical pressure of the surround ing cytoplasm. While some granular cytoplasm has entered th region occupied by the spindle during the early stages, the lates stages show very little addition until the cell plate 1s complete and all of the spindle fibers have been nearly — | The fibers seem to form a barrier against its ingress, rar forms a denser layer around the whole spindle (fig. 7) | possible, too, that the growth of the cell plate by the a previously described has displaced the cytoplasm 7 ’ spindle, and that such a displacement results in a greater p Ss upon the nuclei. ae The history of the carbohydrate material during ie of the cell plate could not be followed with certainty. pind that it always appears plainly in connection with — fibers which are forming the cell plate. In the older po age the cell plate, however, where the spindle fibers had © oe) disappeared, this substance was often indistinguishable : ae those cases where the cell plate was split, a young cell w f nearly always be detected. It did not extend, so fat ea be seen, the entire length of the cleft, but was only : ie portions (fig. 37). A peculiar case was sometimes mens larch, where an incomplete cell plate appeared with 2 ee ous cell wall formed between the halves of the older P' = 1900] THE CELL PLATE IN HIGHER PLANTS 163 lying in the cytoplasm, with some of the original fibers still attached to it while the daughter nuclei were again dividing. Whether this cell plate and wall were finally completed, I did not determine. The same stage of development of the cell plate was often observed, in which no cell wall could be detected. The significance of the presence of a cell wall without the pre- vious appearance of the carbohydrate material has already been discussed, The evidence for the splitting of the cell plate before the formation of the new cell wall need not be discussed in detail here. Treub, in his experiments upon living cells, showed that the splitting occurs, and the new wall is laid down between the separated halves before the cell plate has attained its full growth. My own observations confirm this fact (figs. 79, 37). But one 83 t of difference should be noted between Treub’s obser- “ations and mine. In Treub’s figures the beginning of the Process is represented on one side of the cell where the cell plate ~ reached the membrane of the mother cell, while in my prep- eee it may’ be clearly seen that the splitting begins in the mrs region of the plate and extends toward the periphery. ne mechanics of such splitting is hard to explain. The fact had s evidence of it was found except where the spindle fibers i ‘appeared would indicate that the fibers have no part in Process. From his observations on Fucus, where the — which form the cell plate divide before fusion, Stras- Riss Songs that the splitting may be the result of a similar may ees the higher plants, 7. ¢., the separate cell plate elements two ie ; — they fuse. If such a division takes place the aos * Temain so close together that no evidence of it are so. hice much later stage. Where the cell plate elements expect eae as they are in the larch, it is reasonable to he lack of nd stages showing their division if it really oreurs comes late Such stages seems to me to indicate that the division Plate, ¢ nie the elements have fused into a continuous “ontinuous is be true, the problem is that of the splitting of a Protoplasmic membrane. Strasburger decided that 164 BOTANICAL GAZETTE | SEPTEMBER there is no change of a middle layer in the cell plate into a cel wall. This conclusion rests mainly upon the fact that the halves of the cell plate which appear after splitting are together equa in thickness to the original plate, 7. ¢., there has been no diminw- tion of the substance of the cell plate during the process of splitting. That the cell wall is not a differentiated portion of the cell plate seems to me to be further shown by those cases it which portions of the separate halves of the cell plate appeat with no cell wall between them (figs. 30, 37). | The splitting seems to be due primarily to a differentiation of the substance of the plate itself into two layers. Of whit this differentiation consists is by no means apparent. It is hard to conceive of a layer of protoplasm becoming differentiated into two separate layers similar in all apparent respects to each other. To be sure, the two layers form the boundaries of sia rate cells, a fact which may be taken to indicate a possi . chemical difference between them. This differentiation, iM itself moreover, would not account for the separation of the halves of the plate. A possible explanation for this latter phenometo® may be that there is secreted between the halves some non-stall® able substance, perhaps cell sap, which serves to separate thee The apparent disappearance of the carbohydrate material befor the cell plate halves appear separated may mean that there change in this substance, preparatory to its being deposited # wall, of such a character that it does not take the same ee before. It may be that it simply forms a less dense eee . and that in this form it is first deposited between the halves the cell plate. The appearance of a stained wall would take mean that the substance had again become dense enough eed | the stain. While the above explanations are purely hypothe! . I cannot see that they in any manner do violence to the facts. They are suggested only in order to help bring oe connection that the splitting of the cell plate is apparen” : quite analogous to the longitudinal splitting of tae is : thread, unless Strasburger’s view that the granules div! = : 1900] THE CELL PLATE IN HIGHER PLANTS 165 they fuse into a cell plate is to be accepted. In the splitting of the chromatin thread we may easily imagine the process taking place by constriction, but in the case of a continuous layer of protoplasm like the cell plate such a process would be impos- sible. The more so when we remember, as my figures show, that the splitting begins in the central portion and extends out- ward. The change in position of the cell plate during its growth is shown by figs. r2 and 27. The explanation of such a change is not obvious. The figures show that the nuclei do not move with the cell plate. Whether the cell plate ever changes from a true diagonal to a transverse plane I was unable to determine. The figures show only those cases where the plane in which the cell plate was first formed was not a true diagonal. Nemec” has shown that the position and form of the young spindle can be determined to some extent by means of pressure or tension exerted upon the tissues in which the dividing cells occur. It , Possible in these cases that the conditions of pressure or ten- ‘ion upon the single cells have so changed by growth of some of the surrounding cells of the tissue as to bring about the change in position of the cell plate. The most obvious signif- ase of the shifting of the cell plate is that the original Pesition of the spindle cannot always be taken to determine accurately the ultimate plane of division of the cell. CONCLUSIONS. 1. The most obvious and at the same time most important usin to be derived from the foregoing observations is ee the division of the cell body is due to the activity of the ies of hs cell. The splitting of the cell plate, OF at plants i: differentiation into separate layers, is in the higher not nae ey act in the division of the cell body, for it is Separatio such a differentiation has taken place that sea is a tinct at Be protoplasm of the mother cell into two dis- Parts. That the cell plate is kinoplasmic has already been — + Zelitheilung bei Solanum tuberosum. Flora 86: 214. 1899. 166 BOTANICAL GAZETTE [ SEPTEMBER insisted upon by Strasburger in two of his recent papers.® The difference between Strasburger’s observations and mine is that in the former only a small portion of any one fiber is used in the process of cell plate formation, while in the latter the substance of the whole fiber becomes transformed into a portion of thecell plate, z.¢., in the one case the cell plate is simply a product of the activity of the kinoplasmic fibers, the fibers themselves no! being used in the process, while in the other case the cell plate is a result of a change of form of the substance composing the fibers. The identity in character of the substance of the cell plate and that of the spindle fibers has an important bearing 02 the relations of the different methods of cell division occu in different groups of plants. The process of free cell formatio’ in the embryo of Ephedra, as described by Strasburget, is a interesting case of kinoplasmic activity, in which the division takes place around a single nucleus instead of between two nuclei in regular cell division. The fact that the membranes # formed simultaneously would show that this process goes 0 around all parts of the nucleus alike. It might be contrasted # | this respect with the formation of ascospores, as descr ais Harper, in which the growth of the membrane isa progressive om and the nucleus is consequently modified in form to esr necessity of kinoplasmic activity from one point alone. : method of division of the egg from the endosperm mother a and the synergids, as described by Mottier*, presents a Pr s intermediate between the method of free cell formation © Ephedra and that of regular cell division. Here, vie : are two nuclei lying adjacent to one another, formed in connecting spindles between them, the case of one synergid in Mottier’s jig. 24% - blindly in the cytoplasm, a cell plate is formed 10 such casts following the method of Ephedra. In all of the above ™ the formation of a membrane from the fibers is an est ee “ Ueber Cytoplasmastructur und Zellhiute. 4*Ueber das Verhalten der Kerne bei der Entwickelung des & + 123. 4 der Vorginge bei der Befruchtung. Jahrb. f. wissen. Bot. 31° 1900] THE CELL PLATE IN HIGHER PLANTS 167 fact. The method of cell division by constriction of the plasma membrane, which occurs in many filamentous alge, may be anal- ogous to the growth of the plasma membrane as the cell increases in size, with the difference that in the former case the growth of the plasma membrane is localized and proceeds without the accompanying growth of the cell contents. 2. I have already indicated what seems to me to be the rela- tion of the nucleus to the kinoplasm in the process of cell divi- sion. It is the center of the metabolic processes concerned in the production of the kinoplasm. Whether there is a transfor- mation of some previously existing substance of the cell into the filar form of kinoplasm I have not been able to determine. From the two facts that no such transformation is visible and that a great many of the fibers are used up in the formation of a membrane, I think it probable that they are formed anew in each cell. That the chromatin is the real center for their for- mation is shown by the formation of new radiating fibers around the daughter nuclei in the onion during the diaster stage, and by the formation of a spindle around a single chromosome in the manner described by Juel for Hemerocallis, The history of the kinoplasm in a single cell of the higher plants would ‘sem to be as follows. It is formed as fibers around the rete as acenter. In this form the kinoplasm takes part bi Ris Process of nuclear division, and later divides the cell ee or of the fibers being transformed into a membrane dinates a in splitting the plasma membranes of the s. a ag a cell division does not follow immediately pales cat lvision the filarplasm may be absorbed into the plasm of the cell, either to reappear or to be formed an i ce needed for cell division. The ultimate form of the bec Piasm of any one cell seems to be reached when it has Ome ONS Saat membrane. It is then concerned with the rom the fila Arenas 2 cell. The transformation of kinoplasm hypothesis as into the membrane form seems to show that the a permanent substance, which may be described 168 BOTANICAL GAZETTE [ SEPTEMBER by the term “ filarplasm,” lately proposed by Strasburger,* i not well founded. To be sure there are, in a great many cells spindle fibers that seem to take no part directly in the form: tion of membranes, but they apparently lose their filar form. The most permanent form of the kinoplasm seems to be of the membrane. It would seem better, then, to retain the older physiological term kinoplasm, since such a term would denotea substance having certain physiological properties without nett sarily being limited to any one form. The conclusion reached by Kostanecki #3 of the permanency of the spindle fibers in the cell, and that they are always reproduced by division of the individ: | ual fibers, ‘“‘ omnis radius e radio,” is, as Nemec * has pointed out, unsupported by the history of the fibers in plant cells. It would be impossible, moreover, to account thus for the manne! in which the fibers are transformed into a membrane. 3. The relation of the carbohydrate material to the process of division would seem to show, as already stated, that the sub stance for the formation of the cell wall is held im a reserve form in the protoplasm before it is actually needed for the process of wall formation. The fact that it appeafs in ee tion with the spindle would suggest, as Farmer and ee have shown, that the spindle fibers have for their substance a conductive function. These authors did not ascribe @ pe tive function to the fibers because they did not find any plate. It is not impossible that the cell plate may have? entirely overlooked in this case.. In some of my own ie tions it was often very difficult to discover the young cell a in the midst of the carbohydrate material. o or If the above mentioned relation of the carbohygay rial to the spindle be taken in connection with the facts 5s by Klebs,* and by Townsend,* that the presence of | 42 Zellhaute. 44 Od. cit. 44 Kerntheilung von Allium a 7 Beitrage zur Physiologie der Pflanzenzelle. Untersuch. a. d. bot Tiibingen 2: 500. 1888 “Der Einfluss des Zellkerns auf Bildung der Zellhaute- 30: 484. 1897. Jahr. f, wis: 1900] THE CELL PLATE IN HIGHER PLANTS 169 necessary for the formation of a cell wall, there would be some evidence for the hypothesis that the nucleus forms the cell wall substance. The investigations described in the foregoing pages were begun at Lake Forest University under the direction of Profes- sor R. A. Harper, whose helpful interest and valuable criticism have continued throughout their progress. The work was con- tinued and practically completed in the botanical laboratory of the University of Michigan during the years 1897-8 and 1898-9. I wish here to thank Professors Spalding and New- combe of the above laboratory for the liberal way in which material and equipment were provided, and for many helpful Suggestions received. Professor Jacob Rheigard, of the depart- ment of zodlogy of the University of Michigan, very kindly allowed the use of the photographic apparatus of that depart- ment, and Dr. J. B. Johnston aided me greatly in making the photographs. THE UNIvERsITy OF WISCONSIN. EXPLANATION OF PLATES VIII AND IX. _ Figs. 1-21 are reproductions of photographs made with the Zeiss photo- Ps aa apparatus. (Figs. g and 22 have been purposely omitted.) ig 23-31 are from drawings made with the aid of the camera lucida, Zeiss ae eiageioag objective, and compensation oculars 8, 12, and 18, The size €n reduced 4 in reproduction. Figs. 1-10. Larix ; pollen mother cells. Fig. 1, Equatorial plate. x 750. ohne ym diaster showing first stages in preparation for cell plate Fig i central spindle is differentiated into three zones. X 1200. % wag than preceding ; central spindle alike throughout. X 1200. : “tae plate elements just forming. x 1200. : a Bsc oung cell plate has begun to grow in area; connecting fibers § to disappear. x 1200. F 3 . "GS. 6-10. Later stages in the growth of the cell plate ; see explanation 00, Figs. 11 Fig, 13, —21. Allium Cefa,; cellof growing root-tip. Equatorial plate; spindle fibers arranged into strands or 500, 170 BOTANICAL GAZETTE [ SEPTEMBER Figs. 14-15. Metaphases showing granular appearance of region between the groups of daughter chromosomes. X I500, ' Fic. 16. Stage corresponding to fig. 2. X 1500 Fics. 17-19. Later stages showing the equatorial zone containing car bohydrate material. X 1500 Figs, 18-19. Pormation of cell plate in the midst of the zone of cir bohydrate material. x I500 Fics, 20 and 21. Brow of cell plate and disappearance of spindle bes xX 1500. See also fig. 7 Fics. 12 and 21. ee showing change of position of cell plate froma diagonal to a transverse position. X 1500. | Fic. 23. Larix; showing arrangement of spindle fibers and differenti tion of Bi ec into layers, X 3000. G. 24. Larix, details in relation of radiating fibers to the connecting ae In a stage represented by fig. 7. X 4500 : Fic. 25. Ad/ium, portion of spindle in fain metakinesis, showing “adh tions of fibers to one another and distribution of granules in the spine. : xX 4500. : Fic. 26. Larix; two radiating fibers lying adjacent to the cent! spindle and crossing in such a way as to appear fused. X 4500 Fig. 27. Larix, different stages in the transformation of the substance : of a spindle fiber into a portion of the cell plate. X 4500 ee Fig. 28. A/iium, relation of the connecting fiber to the carbohyd® material at the equator. x 3000. Fig. 29. Aldium ; cell plate elements _ the midst of the carbohydrate material. x 300 . _-FiG. 30. Addium ; late stage in the growth # the cell plate, ae relation of the outer radiating fibers to the growing portion ‘ ee X 4500. a FiG. 31. Altium ; cell plate with young cell wall showing ee where the cell plate is split. x 3000. a . | | | oe on the connecting fibers ERRATUM: on p. 85, line 16 from top, 7g. 10 should read fig: 3° PHYSIOLOGICAL OBSERVATIONS ON SOME . PERENNIAL HERBS. A. RIMBACH. . (WITH PLATE XIII) I. Arisaema Dracontium (L.) Schott.—During winter Arisaema Dracontium consists of a stem-tuber, the growing point of which, covered by several scale leaves, lies at about 5°™ below the sur- face of the earth. It is devoid of roots. Early in April the bud begins to elongate upwards, and at the same time from fif- teen to thirty roots break out in a ring like zone from the base of the bud ( fig. 4). They appear almost simultaneously and grow horizontally, radiating in all directions from their point of origin. They are 1 to 1.5™" in diameter throughout their entire extent, and attain a length of more than 20%. After a time some of them become somewhat transversely wrinkled at the base, because they undergo there a slight longitudinal contraction. At the end of April the scale leaves protrude from the soil. They are tightly appressed to each other and enclose a hollow, ‘n which the foliage leaf develops ( fig. 6). In the latter the end leaflet is vertically extended, the lateral leaflets are bent Mdieigd and the blades of all are involutely rolled up. After “ving reached the surface of the earth, the scale leaves stop re Stowth, and at the end of April or beginning of May the lage leaves and inflorescences break out from their coverings - 4nd unfold in the air. In the course of May the inflorescence : ‘ seus About the same time a second set of roots is formed Mediate] ee diffe y above the first one (fig. 5), but there is a striking rence between the two. The roots of the second set are cker, about 2.5mm £Y grow from and 1900 thi : in diameter, and taper towards the end. their origin vertically downwards or nearly So, ee ©ver into a horizontal direction only at their thin end 171 172 BOTANICAL GAZETTE [SEPTEMRER portion. They undergo very soon a considerable longitudinal — contraction in the thickened basal portion, which is usually? to g™ long. After having attained a certain intensity, this com traction manifests itself externally by a shriveling of the root surface. I noticed, however, that these roots of the second st are not in all individuals developed in the same degree, li specimens, for instance, which were located at a considerable depth, these roots differed very slightly from those first formel On the other hand, in individuals which were placed very super ficially I found the roots exceptionally thick and exceeding) numerous. ~“— The contraction amounts to about 4o per cent. within the Space of 5™" in the swollen basal portion. It diminishes ® intensity towards the thin terminal portion, where no contraction at all takes place. The whole contraction of the root amoul\” to about 15™™ or even more, the contractile region having ong inally an average length of about 5°. Of these facts E if myself by marking and measuring the roots during their dere? opment in the earth. For this purpose the plants were cultivate in specially constructed culture-cases, furnished with bea windows, which could be removed thus permitting access t0 = | roots. The active contractile tissue is here, as in other 18 of similar structure, the cortical parenchyma, The cen of vascular bundles with the endodermis is passively eaten and the same holds good for the epidermis and ent one or two layers of parenchyma immediately below se = These passive layers of the outer cortex very ce A folded, and form transverse wrinkles. During 0%” which process lasts in the whole root about three or fou ametet, ? outermost layers of the active parenchyma after @ time? and become tangentially compressed by the innet wees finally a rather wide area of crushed cells is found * a few layers of the still turgescent innermost cells. In dermis, as well as in the exodermis, there is also 4 Z 1900] PERENNIAL HERBS 173 result of the root contraction noticed. The radial longitudinal walls of their cells, being quite straight at the beginning, as con- traction sets in become marked with wavy foldings. In the endodermis the waves are strongest in that longitudinal band of the wall which corresponds to the well-known dark spot on the cross-section. ~ As each root is fastened at its tip to the earth and at its base to the tuber, in consequence of the contraction a tension is set up init, and the root must give way at the point less firmly fixed. Asa matter of fact, mostly the base of the root moves towards the tip and pulls the tuber with it. Thus, by the com- bined action of all the roots, since all in their basal part point steeply downwards and differ but little in direction, the tuber is drawn down into the earth a certain amount every year. Thus the contractile roots determine to a great extent the position of the tuber. The latter is found sometimes upright, sometimes lying horizontally, not seldom even turned over with the bud directed downwards, so that the leaf-stalks and stems have to make a strong curvature in order to attain an upright position. The situations are largely due either to a uniform or one-sided pull of the roots. I am not able to say with certainty which direction of growth the tuber would take up by itself, and whether its behavior in this respect would be the same under different external conditions. The annual prolongation of the tuber is in larger individuals from 6 to 10™™ ( fig. 6), and if the — of growth were always upright, the dragging action of the —— Moud of course be of essential importance for keeping Sfowing point in place and preventing its emerging from the earth. . Soe ok nee are furnished with root halts; and form oo the tim: a . at the first order in their terminal pee ; their scars the withering of the leaves all the roots die ) mp this dikes at then found in a ring-like zone on the surface In the a . : — S of the scale leaves lateral buds originate, four oF fiy : © every year, which persist after the leaves which support 174 BOTANICAL GAZETTE [SEPTEMBER them have disappeared, and are found in the fall of the sam year as lateral protuberances upon the fully developed portion of the tuber (dd, fig. 6). During the next spring they enlarge considerably, and later, as that portion of the tuber on which they are inserted becomes emptied and dies off, they are st free. Asa rule, they do not yet develop a leaf and roots in the same year, but remain dormant and sprout only in the second year after that in which their supporting leaves were vegetating. | The tuber always contains but one year’s growth ina fully deve! | oped state. The leaves and roots perish in July or August, and soon alt 3 the berries ripen. During the germination of the seed, whic takes place in the following spring, the cotyledon, growing downwards, elongates so much that the growing point of stem of the seedling is located 8 to 10™™ below the seed (fig.1)- The upper end of the cotyledon, which remains in the swells up so as to form an ellipsoid sucker, which finally becom | about 5™™ long and displaces in part the emptying yee” While this is taking place the first foliage leaf sprouts out ‘ penetrates the earth with a knee-like nutation of its sak | fig. 1). Already the first root of the seedling, reaching a length and 0.75™" in thickness, is a little contractile at its bo although it does not become wrinkled. During the su development the little stem of the seedling swells bene below the insertion of the cotyledon, forming a little tuber of the latter, during the first year, two, or mostly tr dom four, adventitious roots are formed (fig. 2). Each these grows longer and thicker than its predecessoh © ops al ; The fourth 10% ps also a longer contractile region. a instance, is usually about 10™ long and gh thick; aud traction amounts within the space of 5% 40 per® total contraction amounts to about 10™™. I obsertee Specimens that by the work of the roots the little drawn down from 8'to 10™ during the first vegetative F So we find, at the end of the first period of growth, ; nal bud of the plant from 1 5 to 20™™ deeper than it Wi 2 1900] PERENNIAL HERBS 175 in the seed before germination (fig. 3). In the following years this migration of the plant downwards continues in a similar way. The alternation of roots mentioned above shows itself from the second year onward. While the occurrence on one and the same plant of two kinds of roots differing in form and function is not very rare, the formation of these at different times, as in Arisaema, has so far been found only in a few species. I have noted the same fact, for instance, in Allium ursinum L., Fritillaria Meleagris L., Scilla bifolia L., and some other monocotyledons. Arisaema triphyllum (L.) Torr. resembles perfectly A. Dra- contiwm in the behavior of the underground organs. The contraction of the roots seems to play a great part in another American species of Araceze, in Spathyema foetida (L.) Raf. In this plant a stem-tuber is formed, which grows vertically upwards, dying off gradually at its lower end. This tuber attains 10° in length and 5°™ in thickness, and comprises the Products of several years. It forms yearly about fifteen roots hear its upper end. These grow obliquely downwards, tapering toward the tip, and produce from their thin end portion lateral rootlets of the first and second orders. They live several years, so that about sixty to seventy of them are found in ame Pent. All the roots are contractile, and apparently prevent gs emerging of the tuber from the ground, This has been i also by Foerste, who found the seedlings germinating ma of the surface of the ground and the top of Be ease C Several inches below the surface. I found saa Serminating on the surface of the soil and the growing pm OF the tuber in several larger specimens at a depth of about Ioc™, Just the (L.) Covi ining 3° in height and 2™ in diameter. All its roots, in a ring-like belt, are more or less contractile basal portion, which becomes transversely HAAG as they grow steeply downwards they are enabled same phenomenon may be observed in Hypoais hirsuta 176 BOTANICAL GAZETTE [SEPTEMBER to drag the tuber vertically into the ground. I found the grow ing point of older specimens mostly in a depth of 3™. In Jrillium sessile L. 1 found most of the roots possessing2 strongly contractile swollen basal portion, which soon becomes wrinkled. In this species the tuber very often by the power of the roots is drawn into a horizontal or even downwardly directel position. Mesadenia tuberosa (Nutt.) Britton, a composite growing of wet places of the prairie, has a stem-tuber about 2™ in length, which yearly grows from 10 to 12™™ vertically upwards, while i dies off in the same proportion at its lower end ( fig. ay The tuber comprises the products of two years, separated by a constrictiot. Every new member of the tuber, in May after its formation, sets out about twelve roots, originating in a simple ring. These 1008 grow obliquely downwards and attain a length of more that 30™. They are at the beginning about 2™ thick, but ze | secondarily in thickness, attaining at least a diametet of (a They exhibit considerable contraction, and pull the stem much back into the ground as it elongates upwards. The 1008 seem also to show reserve material. They persist through . vegetative periods and form later numerous long but very | lateral rootlets, branching sparingly in their turn. ee vegetative multiplication by lateral buds, which become ®Y”” by the decay of the mother tuber. oe The species of Arisaema, Spathyema, Hypoxts, Trill Mesadenia mentioned above can therefore be classed togethe one group, characterized by possessing a rhizome one 3 or less vertically upward, which becomes drawn down pF tractile adventitious roots. To the same group belong e following species: Ayacinthus candicans Baker, Lilium gee: L., Allium ursinum 1., Succisa pratensis Moench, 40 major L. ik. ee Erythronium albidum Nutt.— During sumine’ ee Erythronium albidum \ies hidden in the earth vit ring’ ant without roots. I found the growing point of fia me tee 1900] PERENNIAL HERBS ve ie of which I examined about twenty-five specimens in the month of April, mostly between 15 and 16™ distant from the surface of the earth, the extreme cases being 11 and 20™. The sterile specimens, the bulb of whichas a rule is smaller, were encoun- tered always at a less depth. Of the about 200 specimens of this latter kind which I examined, the larger ones were found atan average depth of 9.5°, the extremes being 5 and 15°", and the smaller ones at an average depth of 7.5°", the extremes being 4 and 13°". I took into consideration only those individ- uals which were found on an even ground and apparently had not suffered any considerable disturbance. Hence the smaller plants, as a rule, are found in a more shallow position than the larger ones, In the late fall—the exact time I am unable to give—the roots break out from the stem portion of the bulb. They appear all at once, their number being in the smallest specimens from four to ten, in the largest from twenty to forty. They are about 0.75™" thick, uniform throughout, become about 15° in length, and never branch. In the specimens I examined root hairs were aot present. These roots do not exhibit any contraction, Their direction of growth varies very much, and seems to be largely influenced by local factors. At the beginning of April the leaves emerge from the ground, penetrating the earth with the cartilaginous point of the erect ‘MWardly convolute blade. The young specimens develop only is mt the older ones a stem with two leaves and with or with- Hower. With the development of the aerial organs the i S . the old bulb-scales and the formation of a new bulb me ag “s € new bulb may be formed miOne to the old one, or we moved from it to a more considerable distance by ba ale enber. Of 200 sterile individuals I examined, tA aif ae arunner, the rest not, and of 25 flowering individ- ree had formed runners. — is solid in its basal half, but in its terminal half it on ik As eoabbeel longitudinal channel, the termination of which € is found usually a little below the middle of its 178 BOTANICAL GAZETTE ‘[sepremam length on the side opposite the roots. In those sterile spec mens which do not form a runner the corresponding opening is found on the same side of the bulb at the base of the foliage — leaf, about 5™" from the roots upwards. In both cases the opening leads to a hollow space, near the end of which the new bulb is formed. In the sterile individuals mostly two runnes are formed by each bulb, the one being about twice as long the other and also thicker. The longer runner reaches an aver age length of about 12, with a diameter of 2-3, the lon est I found being 17™ in length. In flowering specimens | found the runner 3~5™™ thick and 8™ long, but I do not know the definite length it may attain. Where the new bulb is formed close to the old one, one finds instead of the smaller runner oflf a little bud. The smaller runner, as well as the little bud met tioned, may be found sometimes at the right, sometimes at the left side of the main bulb. | Without entering into morphological considerations = the parts of stem and leaves which take part in the formation : these structures, we will consider only their physiological ie ior. At the end of April the elongation of the runner st grains are found in the whole extent of the runne quantity in the new bulb. During May the development new bulb is accomplished, and at the end of the month t age leaves, the old bulb, and the runner die off, so *# beginning of June the new bulb is isolated. It remains mant state until the fall of the same year, when the @ described cycle recommences. The difference between those specime new bulb from the old one by a long runner, #” ' form it close to the old one, lies only in the relative ¢ the stalk uniting the stout stem-portions of the e Those stem-portions in which the leaves are inserted which the roots start are rich in vascular bundles, au nants may be preserved several years. Sometimes 4 1900] PERENNIAL HERBS 179 or four of them are found still attached to a bulb, indicating the places where the plant had rooted in former years. In case no runner is formed, the bulb advances from 2 to 6™™ every year, this being the distance of the points of rooting of the subsequent bulbs. Through the formation of a runner, the distance of the subsequent bulbs amounts to 3 to 10, being asa rule some- what smaller than the length of the runner, as the latter usually is curved, By the growth of the stalk uniting the subsequent bulbs, whether this stalk be short or long, the plant may suffer a dislo- cation both in horizontal and vertical direction. In both cases the position of the rhizome is somewhat singular, the side from which the roots are formed being turned upwards, and the side on which the bud arises downwards. Where a short stalk is formed, it is directed obliquely downwards and advances the plant a few millimeters in this direction. Where a runner is developed, it grows horizontally at the beginning, but later turns more or less vertically downwards, thus placing the new bulb Several centimeters jower. I found in an examination of about 100 specimens the new bulb of the main runner on an average 4.6 deeper than the old one, while the bulb of the smaller runner was sunken but half as deep. The extreme cases of sink- ‘ng observed in the larger runner were 1% and 10, in the Smaller one o°™ and 6™, In full-grown flowering specimens a we to be formed but rarely, as I found in most of them de ae. es of several years’ growth close to the actual bulb, the yous aaa horizontally about 6™™ every year. in saints eek individuals also, even in a rather shallow posi- kind sh: orm a runner every year. Specimens of this latter wich I found had formed no runner during three years; others, which had devel ns: advo of it in tlh Grciase eveloped a runner this year, had been devol ast year’s sts Ing year, as was shown by the remnants of the Year, seemed 4 ; Others, furnished with a runner the present © have formed one also in the foregoing year, Since n ‘ E ‘ thei La the slightest indication of an old bulb was present in ‘T neighborhood. = it 1a a Nae Ai, waist oi ile 180 BOTANICAL GAZETTE | SEPTEMEER I do not know what rule may exist in this alternation or by what factors the formation of a runner may be determined Nevertheless, from my notes it seems that a shallow position a the plant favors the development of a runner. Probably all the shallow, small individuals are derived from seedlings, germinated near the surface of the soil. Their mode of growth must bring them gradually into the depth of the full-grown individuals. By the yearly duplication of their bulbs a rapid vegetative multipl cation takes place. | Erythronium mesachoreum Knerr.—This plant, which reset bies £. albidum very much, inhabits the open prairie. I found the growing point of the bulb in about twenty-five full-gow individuals which I examined between g and 13™ distant from the surface of the earth, the average depth being about ae In this species the small, sterile individuals occupy # ™%* superficial position, being found at a depth of from 308 They apparently have originated from seeds. In the part of April the formation of the new bulb begins. In the young shallow individuals I always found the new bulb remo from the old one by a runner, but the latter is short, _— est I saw being about 20™™ in length, and it grows _ beginning vertically downwards, thus placing the pa year from 3 to 20™™ deeper (fig. 8). In this species ‘ 1 noticed the formation of more than one runner by one b a “a result of this manner of growth is that the on subsequent years are arranged ina vertical row, and I fact, the remainders of the products of five years located way above the living bulb; the leaf of the present ee . its way through the long channel formed by all the dea? ® ments ( fig. 9). By such a movement of growth the plat : reaches a depth beyond which it does not ae oe indeed, that in the old full-grown individuals, S08" depth of about 11°, the bulb grows no longer pe horizontally or nearly so, the new bulb rooting about ally from the old one. . J 1900] PERENNIAL HERBS 181 The observations on Erythronium mesachoreum as well as E. albidum were made in the neighborhood of Lincoln, Neb. So faras the physiological behavior of the rhizome and roots is concerned, Lilium superbum and Medeola Virginiana agree almost perfectly with Erythronium. Lilium superbum 1. has a horizontal rhizome, in which stem and leaves are fleshy, and participate in the same degree in the storing of reserve material. Each year’s growth of the rhizome is in full-grown specimens about 4™ long, forming at first a funner-like stem portion beset with a few small fleshy scales, and at last a stout, bulb-like structure with numerous crowded fleshy scales, from the middle of which the aerial stem arises. From the under side of this terminal bulb-like portion of the thizome, just at the place where the aerial stem originates in June, about ten thin roots grow out horizontally, radiating towards all sides and sparingly forming lateral branches of the frst order. They do not exhibit any contractility, and last but one year, the rhizome dying off from behind very quickly. I ee the growing point of the rhizome in full grown specimens between 7 and 10% below the surface of the earth. The plant and keep this depth, as the roots are quite inactive, ey = movement of growth of the rhizome itself. This behavior, —— some other American species of Lilium seem to share, is rely different from that of Lilium Martagon and other species nis es World, in which the bulb grows vertically upwards, awn down by strongly contractile roots. rea Virginiana Li. has a horizontally creeping tuberous State. oo. but one year’s growth in the fully-developed bears about ernating, tuber-shaped, end portion of the rhizome attain Isem sean or twenty-five thin, thread-like roots, which first and a a ength and produce numerous branches of the tractile Ca the second order. These roots are not con- ing very i late ” all directions from the rhizome, meander- towing in a nae their course, On even soil I found the rhizome Ment is due t tee varying from 1 to 4™. Also here the move- © the rhizome alone, the roots taking no part in it. 182 BOTANICAL GAZETTE [ seprenset The roots of Erythronium albidum and E. mesachoreum, Lilium superbum, and Medeola Virginiana have only a nutritive function They are of no great importance in fixing the plant in the earth, nor do they exert any strain upon the parts from which they Start, nor do they store any considerable quantity of reserit material. In contradistinction to those of Arisaema, Spathyema Trilltum, and Hypoxis, we find in these roots but little cortical parenchyma, no compressed cortical cell layers, no wrinkling 0! the root surface, and a total absence of wavy foldings in the longitudinal walls of their endodermis and exodermis. The spe cies of Erythronium, Lilium, and Medeola above mentioned at representatives of a type of geophilous plants in which the rhizome, by its manner of growth, seeks and keeps a certaa depth in the ground, without any help of the roots. Deda bulbifera 1.., Paris quadrifolia L.., Colchicum autumnale L., One mascula L.,and Platanthera montana Reichb. f. are species which belong to the same type. IIl, 3 : Many perennial herbs develop a long tap root, which bere more or less thick and fleshy, and filled with reserys mab “ae To this group belong the following species: Kuhma eee des L., Lacinaria punctata (Hook.) Kuntze, Grindelia square 3 (Pursh) Dunal, Nothocaleis cuspidata (Pursh) Greene, Kuhns candida (Willd.) Kuntze, K. purpurea (Vent.) MacM., / esculenta Pursh, Astragalus crassicarpus Nutt., Peucedanum | laceum Nutt., Callirrhoé alceoides (Michx.) A. Gray; eg Carolinianum Walt., Aquilegia Canadensis L., Asclepias pe | Lithospermum angustifolium Michx., Physalis longifolia 5s _ a Allionia nyctaginea Michx. a In many of these plants the contractility of the roo! important feature. I have noted this phenomenon - nyctaginea. In seedlings of this plant it can be obser that the base of the cotyledons, which at the be itself above the ground, after some weeks disappea _ Surface of the earth. That this is really due to the © pot the root I satisfied myself by marking the as 1900] PERENNIAL HERBS 183 specimens, grown in a special culture case, from their base with lines of India ink 5™™ apart. These 5™™ spaces shortened to 4"" each within six weeks, which equals a contraction of 20 per cent. At the same time the uppermost line, that next the root base, moved about 6™™ downwards, placing the base of the shoot so much deeper. Also in Aguilegia Canadensis 1 observed a considerable short- ening of the root, which results likewise in drawing down the growing point into the earth. In seedlings of Aquilegia vulgaris L. I noticed that the growing point of the stem, which immedi- ately after germination stood 8™™ above the ground ( fig. 17) was found at the end of the first summer, in consequence of the contraction of root and hypocotyl, about 6™" below the certace of the earth ( fig. 72). The upper lateral roots become bent down during this process in a very characteristic manner, (7, fg. 12). The transverse wrinkling of the surface in the older oe Aquilegia and Allionia is due to the same cause, and “orresponds to the folding of the roots of Arisaema, Hypoxis, and others, There can scarcely be any doubt that in Lithospermum angusti- sah Nothocaleis cuspidata, Peucedanum foeniculaceum, Astragalus pam te Kuhnistera candida, and K. purpurea the root is also Pe a although I have not had the opportunity to measure the und A This seems to be indicated by the following points: beac aunty. cea of the innermost vascular bundles, the the up = of the outer bark, the disturbed position of bales we ateral roots, and finally the position of the esate prolon ee the surface of the earth, in spite of the continua in Kulnia be re: perennial stem-portion towards above. Also to occur a A oe and Grindelia squarrosa contrachee seems it possibl ough in . less degree, while in Lacinaria punctata eee sf cad - exist at all. In pieces of roots tree observed a = tum and Ki uhnia eupatorioides lying in water, In addition seh i a considerable shortening. a. rosa (L.) Hill might say something concerning Lacinaria squar- ; Which, although resembling Z. punctata very 184 BOTANICAL GAZETTE [SEPTEMBER much in its aerial organs and even in its germination, diffes considerably in the development of its underground organs Lacinaria squarrosa has an underground roundish tuber about 3* high with four lateral swellings, a partition that seems to cont spond to the tetrarchic arrangement of the vascular bundle is the primary root. I am not able to say, however, whether the tuber is formed only by the basal portion of the primary 100! or whether the hypocotyl also or the stem take part in it, & all events, the terminal portion of the main root does not pet sist, but is replaced by new lateral roots starting from the tubet Those roots originate in the older plant in four groups of from five to fifteen situated on the four swellings of the tuber (jf L3)i They break out from the tuber all at once in the latter pat | of April at the time when the first leaves appear. They white, soft, somewhat thickened in their basal portion 0% extent of 2 or 3°, attaining here a diameter of 2™ and tape ing very rapidly to a diameter of only 0.5™™. They grow sie : beginning almost vertically downwards, reach about 5° . length, and form very thin branches of the first and secot® orders, with numerous long root-hairs. The thickened portios of the root consists at the beginning mostly of soft, thin- tissue and is contractile, exhibiting according to My meas : ments in the culture case a shortening of 6-10 pet em ee a space of 5™". The whole root probably never shortens | than from 2 to 4™™, The contraction ceases after four " ; weeks of growth of the root, and during this time the pe istic wavy foldings in the walls of the endodermis and = ‘ad make their appearance. Later the root becomes ye BS on account of the development of thick-walled cells mee : tral axis. At the beginning of October, when the ie ig and the aerial shoots begin to die, the roots perish, am oe 6 winter there are no living roots on the plant. In old s noe of Lacinaria squarrosa the roots probably cannot progae | ment of the tuber, while in younger plants it possibly tae pen. But they help apparently to hold the plant in a strain hey exert. This contrivance seems to be 1° 1900} PERENNIAL HERBS 185 perennial organs of attachment are wanting, and since the tuber is very superficially iocated and the aerial shoot is relatively high, heavy, and exposed to strong winds. Physalis longifolia Nutt. does not show any contraction of its long fleshy tap root, but reaches a considerable depth in quite another way. The plumule of the seedling is raised 5 to 10" above the ground and transforms itself afterwards into a long shoot. The primary root grows vertically downwards, sending out numerous thin lateral rootlets, and becomes subsequently thick, fleshy, and filled with starch. Very soon on its surface adventitious buds appear, some at the upper end at the limit of the hypocotyl, rarely on the hypocotyl itself, others deeper down at a distance of 6 or 8 from the surface of the earth. At the close of the vegetative period the upper part of the plant dies down, but the root, at least a part of it, with its buds, hibernates, and at the beginning of the next year one or more of the buds, now much deeper than the plumule was, grow out by means of the reserve material stored up in the root. In older specimens I found the root 1° thick and extending to a depth of more than 50, 2 The seedling of Asclepias Cornuti Decne shows a development Toot shoots quite similar to that of Physalis longifolia. oo Carolinianum, Callirrhoé alceoides, and Nothocalets pias aa ia distinguished by the peculiar phenomenon that in in apie the root is slit into several longitudinal cords found aa of the dying off of certain tissue portions. I cena i root of Delphinium, for instance, divided into eight yorming’ a circle round a central hollow space and con- € and below. In Nothocaleis the root at a length four - Poi variously pierced and divided into two, three, or laticesiat¢ om cords, each of them about 6™ thick, united caused by sj GN heights. This fission of the root is not liar mode eee . decay of the older tissue, but is due to a set Acer 8towth in thickness. The details of the processes | Mi hee, ‘onhdi closer study. A corresponding phenomenon served and studied by Fost in Gentiana cructata L., 186 BOTANICAL GAZETTE [SEPTEMBER Corydalis nobilis Pers., C. ochroleuca Koch, Salvia pratensis Aconitum Lycoctonum L., and Sedum Aizoon L. 1 observed it in Scabiosa arvensis L. and saw a fission into four cords also inthe subterranean stem of Gentiana puberula Michx. In addition, I have noticed in MNothocaleis cuspidata a daily opening and closing of the flower heads. My observations wett made between May 1 and 15, in sunny, moderately warm, windy weather on the prairie near Lincoln, Nebraska, where this plant grows naturally. The heads begin to open at 7 A. ™., and are at 9 a. M. fully expanded. They remain so till 3p %.@ which time they begin to close again. At 4:30 P.M. they ae all closed, and they stay so during the night. All the plants mentioned, furnished with a deep tap rok confined to the place they occupied when germinating. Besides, they lack vegetative multiplication, their only means of propag tion being by seeds. Perennials of this type are exceeding!) numerous on the prairie. Nevertheless, other forms occur als, like Helianthus scaberrimus, with long, horizontal rhizomes, 5 means of which they are enabled not only to change thett place but also to multiply vegetatively. % Helianthus scaberrimus FE\l., as found in early spring, pi of a subterranean shoot 2 to 4™ long, horizontal or Be - at its end, furnished with about twelve long rigid roots, Gs obliquely downwards and forwards. At the end of Api terminal bud of the rhizome leaves out, forming imme™ pe long internodes, the first of which bear scale leaves. ™ middle of May, from the stem nodes between the roots: from the axils of the scale leaves above them, Tunners start. These take up a horizontal direction growing till August, attaining a length of from Like the leaves of the aerial stem, the scales 0 and coll 50 to protecting it during its way through the earth. The to 3mm thick and consists of about twenty interno ; of which may measure 8°", The last four or five intern the runner are very short, and this stout end-portion ' 1900 ] PERENNIAL HERBS 187 middle of September gives rise to new roots, while its terminal bud, having stopped its growth, remains quiescent until next spring. At the middle of November, when the aerial shoot has died down, the runner begins to decay from its base, while the roots have reached a length of about 20™ and are still growing. During winter the runner decays, only its end portion with the roots remaining alive, and the plant assumes again the form in which we find it in the spring. LINCOLN, NEBRASKA. BIBLIOGRAPHY. De Vrizs, H. Ueber die Kontraction der Wurzeln. Landwirtschaftl. Jahrbiicher 6: 927. 1877; 8: 474. 1879; 9: 37. 1880 WarminG, Euc. Om Skudbygning, Overvintring og Foryngelse. Natur- hist. Forenings Festskr. Kjébenhavn 188 eber perenne Gewdchse. (Botan. Centralblatt. 18: —. 1884.) ForRsTe, AuG. F. The development of Symplocarpus foetidus (L.) Salisb. Bull, Torrey Bot. Club 15: 154. 1888. Fost, L. Die Zerkliiftungen einiger Rhizome und Wurzeln, _ Bot. Zeitung 48: —. 1890. ARESCHOUG, F. W. C. Beitrage zur Biologie der geophilen Pflanzen. Acta Reg. Soc. Phys. 6: —. 1806. RImBacu, A. Ueber die Lebensweise des Arum maculatum. Berichte d. Deutsch. Bot. Ges. 15: 178. 1897. Lebensverhaltnisse des Altium ursinum. Ibid. 15.248. 1897. oe Beobachtungen an Colchicum autumnale. Ibid. 15: 298. 97- Die kontractilen Wurzeln und ihre Thitigkeit. Beitrige zur wissen- Schaftl, Botanik. 2: 1, 1897. Das Tiefenwachstum der Rhizome. /did. 3:177. 1898. a Lilium Martagon. Berichte d. Deutsch. Bot. Ges. 16: 104. I . Beitrage zur Physiologie der Wurzeln. did. 17: 18. 1899. EXPLANATION OF PLATE XIII. oo ie drawn from nature, and, with exception of fig. 70, are Fig, ; j S gist horizontal dash-lines indicate the surface of the ete - Seed ; Meer ts of Arisaema Dracontium still in connection wit Fig, 2 a. cotyledon ; Z, first leaf : yr, first root. — developed a v€ same, near the end of the first period of growth, having Its roots; 7, tuber ; ~', contractile adventitious roots. 188 BOTANICAL GAZETTE [SEPTEMBER Fic. 3. The same, in the resting state after the first period of growh Fig 4. Small specimen of Arisaema Dracontium in April, forming the first set of roots; 7%, thin roots of the first set ; sc, scars from the rootsofthe — preceding years. ee Fic. 5. A similar specimen, in May, forming the second set of roots; r', thin roots of the Urst Serr", enick contractile roots of the second set. portions emptied in former years; dd, lateral bud; the shaded part of tuber is to be emptied this year. Fic. 7. Tuber of Mesadenia tuberosa, in May ; longitudinal section; roots formed in the present year; 72, roots formed in the preceding year. Fic. 8. Young descending specimen of LErythronium mesachoreum April, with the runner developed; 4, bulb; /, leaf-stalk; % roots; f, 3 runner. . ie g. Young sterile descending a of Erythronium ne with its et of the five preceding year: 3 Fig. 10. Tip of the runner of Rylan mesachoreum ,; cian A tudinal section; 4, new bulb. 5. ou Pia, tt. Becdiing of Aguilegia vulgaris, soon after germination; & hypocotyl; ¢, cotyledons ; 7, primary root. . FIG The same, at the end of the first period of growth; ae where the growing point is situated; 7, primary root; 7, lateral roots ®° down by the contraction of the main root. , FiG. 13. Lacinaria sguarrosa, subterranean part of a full-grown men in summer; ¢, tuber; 7, contractile root. : asi yh Dae et BOTAN. ICAL | GAZETTE, XXX PLAT. EX, Ps CONTRIBUTIONS FROM THE ROCKY MOUNTAIN HERBARIUM. I. AVEN NELSON, Draba Yellowstonensis.— Annual, with stems of two kinds; the principal stem slender, erect, scape-like, 2-3" high includ- ing the raceme,,simple or with long-peduncled racemes from the uppermost axil or axils; the one or more accessory stems from the base slenderer and shorter, ascending or erect: leaves mostly basal; root-leaves rosulate, from broadly linear to narrowly elliptic, sub-acute, entire or nearly so, I-2°™ long; the few (2-4) stem leaves mostly near the base, narrowly ovate, generally smaller than the root-leaves ; pubescence on the leaves finely stellate, on the stems and peduncles sparse, the hairs more or less branching : racemes long in fruit, usually more than half the whole height of the plant: pedicels shorter than the capsules: flowers small; the sepals elliptic, obtuse, about half as long as the petals; the petals cuneate-spatulate, barely emarginate, » 2-3 long: capsule linear-oblong, tapering slightly to the apex, 10-13™" long, finely pubescent; the style very short (less than 0.5™ long) but evident; the stigma 2-lobed. Avery distinct species, having its nearest ally in D. montana Wats., a plant aA of More southern range. \ Two Collections of this were secured in Yellowstone p&rk where it occurs A Siete rane ee L operidium pubicarpum.— Annual, with slender vertical tap -gcthe stem paniculately branched from near the base, in well “evelo, ped plants the branches similarly branched, only O15 high ams | ding the racemes, obscurely puberulent: leaves small, Slabrous yor nearly so, linear or somewhat spatulate, acute, the broader ¢ .ynes remotely cut-toothed: beginning to blossom when Nery smal, fd, the fruiting raceme crowded: pedicels short, hardly Pe jong asY\ the capsule : petals wanting, the sepals purplish : ae : XN 189 190 BOTANICAL GAZETTE [ SEPTEMBER capsule permanently finely pubescent, from broadly oval to orbicular ; stigma sessile in the narrow shallow notch: cotyledo incumbent. eo Most nearly allied to Z. afeta/um Willd., from which it differs in being lower and more divaricately branched from near the base, while Z. apefalun has a stem simple at base and branched above. L. pubicarpum is wh restricted below the flower cluster, since this is very short and the pedicels become gradually divaricate. Its puberulent capsules separate it at once frm L. apetalum. Two collections secured: no. 6235, Nez Perces creek, Yellowstone path July 30; no. 6793, Dwelle’s, Mont., August 31, 1899. Arabis densicaulis.—Biennial or possibly more enduring, the tap root producing several or more often numerous crowded stems from its crown: stems ascending, 3—5% high (including the raceme), leafy only toward the base, simple or some of the larger ones sparingly branched, glabrous or slightly hirsute neat the base: root-leaves crowded-rosulate, oblanceolate, alee petioled, 2~3°™ long, finely stellate-pubescent ; stem-leaves rather numerous, glabrous, broadly linear or tapering uniformly from a broadish base to an acute apex, auriculate-clasping, the lobes short: flowers small, the petals white or purplish, linear-spatua® about 5™" long and nearly twice as long as the oblong sepals: a fruiting raceme very long, often two thirds of the whole lengts : the numerous pods arcuate and widely divaricate oF sone drooping but not pendulous, 4—5™ long, about 2” wide, | Fe obscurely I-nerved at base; the pedicels about fee! long: * oval, in one row, scarcely winged; the cotyledons obliq! approaching incumbent. em The cotyledons seem to indicate this as : member of the section i BRINA (Syn. Fl. 1: 159), but I am unable to find in any of the known ™® | a close ally. It may be recognized easily by its numerous stems fT" the crowns, and by the numerous, widely divaricate, arcuate long, naked racemes. The type Specimens were secured on partly wooded, hard, grav slopes, near Undine falls, Yellowstone park, July 6, 1899, no. se Arabis fructicosa.— Similar in size and habit to of nee ing, glabrous throughout except for some fine stellate ane at base: the stems even more numerous, with cate! . ee f f AOR et 1900] ROCKY MOUNTAIN HERBARIUM Ig! bases: crown leaves less crowded, some of them sparsely den- tate; the stem leaves oblong to ovate, the larger ones dentate: fruiting racemes shorter, the pods broader, scarcely arcuate, divaricate-ascending ; the flowers larger as are also the seeds. No other Arabis is known to me that has the habit of this. A single plant sometimes has fifty or more assurgent stems and forms a hemispherical mat several decimeters in diameter. In this respect the preceding species most nearly approaches it. The two were found in the same locality, but they are at once recognized as different. The type is no. 5681, Undine falls, July 6, 1899. Arabis lignipes.— Short-lived perennial, simple-stemmed or more rarely with two or three stems from the summit of the tap root; the woody base of the stem persistent, apparently of as many internodes as the plant is years old, leafless, more or less Covered with the old petioles ; the internodes variable, usually only 1-3" long, the whole forming a naked woody foot sur- mounted by the crown of rosulate leaves at the base of the herbaceous part of the stem; herbaceous stems ultimately 3-5°™ high, begirning to blossom when quite low, erect, finely stellate- pubescent below, glabrous upward, becoming smooth throughout in age: the rosulate leaves small and crowded, entire, minutely but densely stellate pubescent, narrowly oblanceolate, tapering to a short petiole, r—2-™ long ; the stem leaves numerous, sessile, almost linear, tapering to an acute apex from an auricular sagit- tate base, slightly longer than the rosulate leaves: raceme Crowded in anthesis, open in fruit : pedicels sharply deflexed €xcept in the youngest buds, at first minutely pubescent as are also the sepals, 5-7™™ long : petals purplish or sometimes white, _"atrowly spatulate, 5-6" long, nearly twice as long as the Sepals : pod pendent, straight or curved, smooth, 1-nerved, cs long, about 2™ broad: seeds in one row, broadly oval, meatcely wing-margined, about 1™™ long. : | This finds its nearest ally in 4. Holboe//ii Hornem., from which its naked oody foot, its invariably simple stems, its smaller entire leaves, and its per- character separates it. The following collections of it were secured on dry, sandy or stony bottom lands in Yellowstone park: no. 5503 and 5505, Madison river, June 23, Te n0. 583, Glen creek, June 30, 1900. 192 BOTANICAL GAZETTE -[sepreven Arabis pendulocarpa.—Perennial (probably short-lived), a short, simple or branching, woody caudex surmounting a slender _ taproot: stems I-3, only one from each crown, simple, ascending, : rather weak, about 2 high, nearly glabrous except at the base eee the remains of those of former years, closely and finely stellate pubescent, narrowly oblong to elliptic, tapering into a short pe- — iole ; the stem-leaves crowded toward the base, linear-oblong, 4 sessile, not auriculate, 5—10o™™ long, usually longer than those of the crowns: flowers few, small, nearly erect at anthesis but the siliques soon pendent: pedicels 6-8™ long, glabrous or neatly so: petals white or tinged “with purple, about 5™™ long, dis tinctly longer than the sparsely hairy sepals: pods 4-6™ long, about 2™ wide; the seeds narrowly wing-margined. The key of the Synoptical Fhra throws this into close proximity to 4. Pulchra Jones, but it is probably more nearly allied to A. Hodboellii Homen t occurs among the rocks on exposed or partly wooded hilltops. page ve twice, only in Yellowstone park: no. 5504, Madison river, June 23,,1899; : no. 5728, Yellowstone river near Junction butte, July 9, 1899. : , : Arabis elegans.— A tall biennial from a vertical tap 10h — 6-10 high: stem simple and strict (rarely a branch a from the base), a pubescence of branched hairs below, becoming glabrate upward, leafy up to the inflorescence : leaves cro on the lower part of the stem but not rosulate at the > mostly entire, more rarely some of them remotely em” finely pubescent or the uppermost almost glabrous; the: oblanceolate, petioled, passing into the oblong-lineat,, ; auriculate ones of the middle stem ; the uppel oa smaller, linear, Sagittate-clasping : raceme either few- bio ere (ae 1900] ROCKY MOUNTAIN HERBARIUM 193 This species bears in its habit a marked resemblance to A. confinis Wats., but its narrower pods, which are not beaked, the shorter pedicels, and less glaucous appearance will aid in distinguishing it from its eastern ally. It is of frequent occurrence in the open woods on moist slopes. No. 5601, Mammoth hot springs, June 30; nos. 5676 and 5680, Undine falls, July 6, 1899, the latter being the type number. Arabis divaricarpa.— In habit resembling the preceding, pos- sibly sometimes perennial, 4-6%™ high, glabrous except on the rosulate root-leaves, somewhat glaucous, more or less tinged with purple throughout : root-leaves petioled, slenderly oblanceo- late, crowded on the crowns, the pubescence minute and branched; the stem-leaves linear-oblong, sagittate-clasping, 1-4 long: inflorescence glabrous ; the flowers purple to white, smaller than in the preceding: pedicels about 5™™ long: pods uniformly divaricate ascending, straight, 3-5°™ long, about 3™™ broad, most of them conspicuously I-nerved from base to apex. Like the preceding species, I can compare it only to A. confinis, from which it differs in its broader pod, shorter pedicels, mostly entire leaves which hie stem are rather acutely lobed at the base. The seeds also are quite ifferent ; j etn secured in two localities on the open, sandy hillsides overlooking ne lake, August 1899, nos. 6352 and 6622. eee Viola Thorii,— Root mostly simple, semi-fleshy, rather large for the plant: stems several, short, about 15™™ high: leaves on a long, broadly ovate, truncate at base or abruptly os te the petiole, very coarsely and bluntly dentate, Sparsely Puberulent below or entirely glabrous, 12-20" long: Ses Sore sur passing the leaves, very sparsely puberu- leak < ie os * Sepals linear-lanceolate, glabrous, 4™™ long or weicia *s pt ie long, glabrous, yellow, the two upper _ OWN on the back, Fey — Greene, which I have not seen, is wtb ees a ee and has TR agg e Proposed lacks the cinerous cngret of - — the other, which oc In Contrast with the hastate or lobed leav € type eB as thickened or flattened petioles. Near the sums, 7 20 5816, were secured on the moist, open slop Mmit of the Thunderer, Yellowstone park, July 13, 1899. I has 194 BOTANICAL GAZETTE [SEPTEMBER also been collected by Dr. Blankinship in Montana and in Yellowstone park, but I am unable to cite his numbers. Epilobium Wyomingense.—Perennial, spreading by filiform remotely scaly subterranean shoots which end in ovoid winter bulblets with few fleshy scales: stems slender, 2-4 high, strictly erect, mostly simple, more rarely with slender, erect branches from the axils of the opposite leaves; wholly glabrous below, towards the summit of stems and branches (if any) am obscure puberulence: leaves thin and glabrous, linear, tapering from the middle to both ends, sub-acute, from 3—-5™ long (rarely even 8™ long), 2-5™" broad, the uppermost not noticeably reduced, midrib evident, the lateral veins obscure, plane, oF the margin barely revolute, opposite except the floral, the few (4-7) pairs nearly equidistant, often shorter than the internodes ; those of the branches similar: flowers several, erect, small; the calyx cleft nearly to the base; the petals white, ovate, deeply triangu lar-notched at apex, 3-4™™ long, a little longer than the sepals: capsules linear, 4-7 long, minutely cinereous-puberulent; th pedicels variable, from much shorter to even exceeding the nail sule: stigma oblong or short-clavate, barely notched at ape*: seeds numerous, fusiform, smooth, scarcely beaked; the com white, persistent. This species is not very closely allied with any of the species re me, though in a few respects it suggests . Oregonense gractllimin” Slaberrimum. In the more essential characters it seems to be evn palustre, from which its longer smooth leaves and smooth stems an smooth almost beakless seed separate it. +o gtreall ring It occurs in dense patches, on the grassy, boggy banks of SP Snake lets. Yellowstone park, no. 5902, Yancey’s, July 16; no 6428, neat river, August 12, 1899. d from the Cryptanthe multicaulis.— Several to many: stemme sliate- crown of the vertical tap root; the stems rather slender, pe hirsute, erect from a mostly short-decumbent base, Sp 5 numerou> paniculate-branched above, 1 5—20™ high: leaves rather id hatts broadly linear, 2-3 long, the unequal, whitish, mere with pustulate bases: spikes slender, moderately ct ve t fruit: sepals setose-hispid, the stouter setae yellow! pa rae — EO EE nay 2) 2 I 1900] ROCKY MOUNTAIN HERBARIUM 95 midrib not evidently thickened, very narrowly we nate, about 5™™ long: nutlets obscurely roughened under a lens, ovate with sub-acute Sue, tess than 27° long, the narrow Bore forked at base but without conspicuous open areola, similar, usually only three maturing. a Sikes This species is allied to C. Pattersoni and to C. ramulosissima. ints it is intermediate between them, but in floral and fruit characters it differs ROSE fan i 6440, from Snake river, Yellowstone park, August 13, 1899, . CRYPTANTHE AFFINIS flexuosa,— Evidently allied to C. affinis, buta larger plant, 3—54m high: stems loosely branched from near the base upward, the branches long and flexuous: leaves ee rowly oblong, oy long: fruiting spike long and open ; ep s lanceolate-acuminate, densely hispid at base, the tips Te *pen and spreading, 6—gm™ long: nutlets mottled, smooth an shining, Ovate-acute, about 2™™ long, the ventral groove nearly Closed, forked at the base but without areola. “oe This May prove worthy of specific rank, but until further material is i : ; ther “cured it may best Stand as a variety. The species, I think, occurs fur toward the west and northwest only. Secured in Jackson’s hole, near Jackson’s lake, August 17, 1899. Mertensia amoena.— Root somewhat woody, more or less branched, surmounted by a branched caudex: branches of the “audex few to several (3-8), crowded, erect, closely covered with dead leaf-bases : stems one or more from each crown, sim- Pie “scending, ;—2am high, pubescent with short spreading or Ctisped hairs : Crown-leaves oblong-lanceolate, sub-acute, glab- Je eink softly hispid-pubescent above and on the s Oe ots long, about one fourth as broad, on slender petioles which | = hows longer than the blade: stem leaves rather crowded, €ar or narrowly oblanceolate, sessile, 3-6™ long, milar to that of the root-leaves; inflorescence Rio calyx-lobes lanceolate, sparsely ciliate-hirsute, about é as long as the tube of the corolla; corolla about 15" a € tube scarcely longer than the campanulate limb, the lobes the limb abo ; . 4 ents ut half its length, crests inconspicuous; filam 190 BOTANICAL GAZETTE [SEPTEMBER broad as the anthers, inserted in the throat, the free portion about as long as the anther; the style nearly as long as the corolla. This is probably a part of the 47. Zanceo/ata (Pursh) DC. of Gray in Proc. Am. Acad. 10:53, and of the Syn. Fl. 2: 201, though it is very distinct from what Pursh and De Candolle understood by that species. The original seems to have been that glaucous, glabrous (at most slightly scabrous) plant which we know from the eastern slopes and foothills of the Rocky mountains, and which extends eastward toward the Missouri in the hill regions. That has rather thick fleshy leaves and has fewer stems. I have for years been familiar with it in southern and eastern Wyoming. During the season of 1899 Mon- tana and northwestern Wyoming, where the species now proposed is common, came under my observation. It did not occur to me that anyone could pos sibly have called it J/. lanceolata, so different are they in the field. M. amoena may be recognized by its cespitose habit, its hirsute (almost cinereous and never glaucous) leaves and stems, and the more crowded inflorescence, which in young plants reminds one of J/. oblongifolia, It has the habit and leafiness of J/. foliosa, and is more clearly distinct from J. Janceolata than M. Fendleri is from that species. Collected at Monida, Mont., June 1 5, no. 5413; Glen creek, Yellowstone park, June 29, 1899, no. 5556; at both of which places it was abundant. Solidago dilatata.— Perennial, from a woody root surmounted by a branched caudex bearing a few surculiferous branches which terminate in a fascicle of leaves: stems single from the crowns simple, or branched above, rather stout, somewhat se glabrous, 4—64™ high: leaves glabrous, conspicuously retic- ulate-veiny below, minutely scabro-ciliate on the margins; ©" leaves oblong-spatulate to elliptic, tapering into a broad 28 gined petiole which is sometimes as long as the blade, ¢t ; closely or remotely serrate, mostly obtuse at apex, 8-15" ye the stem leaves numerous, sessile, mostly small (2357 en Fescence nearly glabrous (some ciliate straggling hairs), gee lately corymbose, either compact or quite open, the ue pedunculate branches leafy bracteate: heads numerous, ne “ pediceled; the disk about 6™ high; involucral bracts in te three rows, the shorter outer ones very few, the two we sub-equal, minutely ciliate on the margins, linear, most © obtusish and slightly dilated upwards: rays 8-10, iat akenes short and lightly pubescent. EEE _ picuous - FEN a ee eee SDPO nee ee Nin ee ae ee ae ee) ee aa 1900 | ROCKY MOUNTAIN HERBARIUM 197 This is to be compared with S. mu/tiradiata scopulorum, but it is a much larger plant, with larger root-leaves and larger more open inflorescence. In being practically glabrous it also differs from that, and the upwardly u. lated. bracts especially distinguish it. It was abundant in loose gravelly soil in the open woods in the southern part of Yellowstone park, no. 6586, August 21, 1899. Machaeranthera superba —Probably only biennial, very numer- ously branched from the crown of the slender tap root: stems decumbent at base and widely spreading, 8—-15™ long, each simple below, but paniculately corymbose as to the inflorescence, their purplish hue masked by a minute cinereous puberulence ; root-leaves (mostly wanting at flowering time) oblong-lance- olate, with minute spine-tipped teeth, cuspidate-obtuse, tapering into a slender petiole somewhat shorter than the blade, whole length 4-6: stem leaves rather numerous, broadly linear to narrowly oblanceolate, entire or remotely denticulate, the teeth and apex cusped as in the root-leaves, minutely and softly sub- cinereous (scarcely canescent), 3-5°" long, smaller in the inflo- rescence: heads moderately large, disk about 1°™ high, nearly as broad; bracts of the involucre oblong, acute, tips mostly erect, decidedly tinged with purple which is only slightly obscured by the thin puberulence, very rarely a few gland-tipped haits on the Margins: rays 12 (more or fewer), a deep blue an yom . OF this Species, which was submitted with a number of others, Dr. Greene Writes as follo S: “A subalpine looking, too showy form of J/, canescens. - it cannot be referred to M/. subalpina, however much it looks like it at rst glance.” . a : i Species, of which | have typical specimens (the type number ia ag lam Satisfied that the species now proposed is amply distinct from thal egy as_understood (evidently) by Pursh, Nuttall, — i“ _ essentially erect plant even though branched from ee one misege : hg leaves are distinctly serrate or toothed, the bracts are evi oe is more Pped, the Pubescence is canescent rather than cinereous. AM. super Nearly allj 4S Compared With A, canesc en tis distinguished from it by the almost entire absence of viscid ince that was written I have given much study to both of the 198 BOTANICAL GAZETTE | SEPTEMBER or glandular hairs, and from both by its less erect habit and by its broad, colored involucral bracts. It occurred in the greatest abundance in one locality only, an open, sandy hillside near Yellowstone lake. he depressed, mat-like plants, with their relatively large, showy heads, were singularly attractive and invited the closest attention. The type number is 6337, from the Thumb, August 6, 1899. Erigeron Yellowstonensis.— Biennial, or probably many of the plants more enduring, with a strong vertical tap root: generally only one stem from the enlarged crown (more rarely 2-5), sim ple, stout, striate, erect, paniculately branched as to the inflo- rescence, 3-6™ high, purplish, glabrate, the whitish hairs very straggling, obscurely granular (scarcely glutinous): leaves numerous, pubescence nearly wanting, similar to that of the stem; crown leaves oblanceolate, petioles 3-6™ long; lower stem leaves similar but with short winged petioles; uppe! leaves sessile, narrowly lanceolate, not much reduced, the short branches of the panicle from their axils; bracts small, linear: heads numerous, on rather slender peduncles ; involucral bracts dark green, in two rows, subequal, very narrow, acuminate, shorter than the 1-™ high disk: flowers very numerous; rays filiform, purplish, only moderately numerous, largely concealed by copious pappus : akenes linear, appearing glabrous but sparselj pubescent under the microscope, less than 2™™ long; the soft dirty-white pappus nearly three times as long. ) Blytt) a into the Plant now proposed as a species. Neither does it seem probable th other usually accepted synonyms of £. Drocbachensis represent th of they also refer to European or arctic forms, except the £. a . ooker’s Flora. The latter seems to have spatulate root-leaves, wel = Ones almost linear; a racemiform inflorescence, with very long lower P cles, making an approach to a corymb; the pappus of a more yellowish : This plant was found in abundance near Yellowstone lake, 0 o _ in loose sandy soil: nos, 6348 and 6615, the Thumb, : ERIGERON MULTIFIDUS incertus.—-Caudex densely cespiton” its branches comparatively long and woody, roughened with is plant, for the a at 1900] ROCKY MOUNTAIN HERBARIUM 199 old leaf-bases: stems usually two or more from each crown, rather slender, curved ascending or erect, 12-18™ long, sparsely ciliate and obscurely granulo-puberulent, monocephalous: leaves crowded on the crowns, green and appearing glabrate but ciliate on the petioles and often on the blades, obscurely granulo- pubescent, simply three-parted, or more often each segment again three-parted ; ultimate segments linear-oblong, 4-6™ long ; petioles slender, 2-3°" long; stem leaves few (2-4), bract-like, mostly linear, entire, the lower occasionally trifid: heads 8-12™ oe ne (apparently sometimes a few filiform ones); S very numerous; involucre ciliate-pubescent and oT the longer hairs, bracts linear-acuminate, fasts ie : coat akenes flattened, narrowly obovate, B lone ae fh nt, about 2™" long; the soft pappus about twice 8 e akene. i. oo. about the same relation to F. mudtifidus Rydb. (Fl. two varieties . rage glabratus does to the same species. The glabrous in ee har “" e habit and leaf outline. They are both fradiate. The variety oe " Ae e one is wholly glabrous and the other is larger size, in being more a differs from E. multifidus discotdeus in its Nos. 5538 and 6066 oe ea sh in its cespitose caudex. 2343 from Dome ro. ps - one park belong here, as does also no. stony hills and AS faily at g Horn mountains, 1896. It occurs on dry A y abundant. coe aaah ped etseorus longinodosa. Stocks fro Toots, erect, 4— much exceedi Pubescence of Stems single from hori- ee sie which spring numerous, thick, fibrous high, the internodes long, usually 10-15™, ng the leaves or rarely only equaling them: (especially on a kinds, a short, dense, glandular puberulence hairs (especiall e peduncle), and some scattering white crisped the lowest ne on the involucre): leaves 3-5 pairs, denticulate; Petioles often : RS Nap 6-10™ long, on slender ast, rarely oe ong, disappearing early in the season or, at Tower, taperin eae the next lowest mostly obovate or nar cither shorter rag a short margined petiole, variable in size, Sessile by a 4 onger than the lowest pair; the upper pairs road base, very variable in size and shape, from 200 BOTANICAL GAZETTE [SEPTEMBER ovate to oblong, 5—15°™ long: heads 1-3, mostly single and then long-peduncled; if more, then from the axils of the uppermost leaves which are often much reduced, occasionally only one of the leaves and lateral peduncles developing; lateral peduncles — equaling or even exceeding the terminal, 1%™ (more or less) in length: heads large (if more than one somewhat reduced), disk 15—20™™ high, 20-30™™ broad ; rays about twelve, 20-30 long, 6-8™" broad; involucral bracts 14—20, lanceolate, acute, shorter than the disk: akenes striate, nearly linear, but tapering to the base, obscurely short-hispid on the angles, about 5™™ long, equaling the sordid, sub-plumose pappus. I have described this in detail, for I believe that it will ultimately be shown that this is a good species rather than a variety. The A. Chamissonis Less., from Unalashka, seems to be a much more pubescent plant, with nat- rower leaves and more pubescent akenes, the internodes, as compared with the leaves, relatively shorter, ” The following numbers represent some of the collections of this species in Wyoming : 1702, 1785, 3571, and 6379, the last from Yellowstone park, August 1899. M. E. Jones no, 5883, from Utah, is this species. Arnica Columbiana.— Perennial, 4-8 high: stems rather stout, erect, striate, simple below, paniculately corymbose above, with some lanate white pubescence: root-leaves not know! stem leaves several (six or more pairs not counting the rameal), the lower apparently petioled, the middle and upp ample, sessile, clasping, entire, rather conspicuously nerveds oblong, sub-acute, finely pubescent below, obscurely s0 above, 8-14" long, 3-52 broad; rameal leaves and bracts ovate, the smaller ones acuminate: inflorescence an ample panicled cymose corymb of 15-30 or more rather unequal heads: 1nV0 pan lucral bracts 29 (more or less), oblong, sub-acute, oe the disk: rays 12 or | nase -4™ broads ( y r less, 10-14™™ long, 3-4 der bas F small, 5-nerved, linear-subcylindric, tapering toa slen only about 3m long, shorter than the sordid pappus- Of this unusually distinct species I have seen but two specimens, which are in the herbarium of the Montana State College. The type is Mrs, J. J. Kennedy’s no. 24, Columbia falls, Mes Williams’ no, 1049, from the same locality, June 14 1894» Mont., 1894 hE al, poth of : js the | EY ee ee, ey ee Le Ee a we ee Fear Pee eee 1900] ROCKY MOUNTAIN HERBARIUM zol same as to the larger specimen (on the sheet before me). Both were dis- tributed as A. amplexicaudis Nutt., to which they bear but little resemblance. Mrs. Kennedy's specimens are included by Dr. Rydberg in his 4. amplexifolia (Nutt.) of his Flora of Montana. Arnica ocreata.— From slender horizontal rootstocks: stems slender, 3-42" high, erect, very leafy, nearly glabrous, more or less finely granular-glandular, occasionally some straggling woolly hairs: leaves 6-10 pairs, ascending or erect, from broadly ‘o narrowly lanceolate, entire, sub-acute, gradually smaller "pward and becoming bract-like, all but the uppermost much *xceeding the short internodes (even overlapping two or three of the internodes), the lowest petioled, the uppermost sessile: petioles slender, dilated at the base and connate, the pair form- "8 *1 Octea or sheath which in the lowest leaves is 2-3™ long, ne sheath and petioles gradually shorter upward to about the middle of the stem where both become wholly absent: heads as 2 several; if three, corymbose, the two from the axils of sai bracts ; if more than three, mostly somewhat racemose Se of alternate bracts ; terminal head largest, a hier 1g ts the oblong, obtusish bracts, about: 15 pedicels nee se 1™ long; lateral heads somewhat reduce shins ate : rather slender, very lightly woolly-pubescent : Tow, taperin y glabrous, lightly striate, somewhat flattened, nar- ae 8 to the base, nearly equaling the soft pappus. hot "ag at 3 as specimens in the collections go, A. foliosa. It is tions of that om ois See: which is probably a rarer plant, though port description and as ma : Sais That, as may we seen by ee ete ie ay ! € e gathered from Hooker's Flora and from ae Chamissonis) is a = : (w ere 1t was compared with A, montane . : y different plant, and nearly allied to A. Chamtssonis. What equally leafy “ote Strict habit of that, is tomentose pubescent, meee , ughout, the leaves callous-denticulate, evidently nerve dias ieee a Sa It may be recognized by its Ces pitose habit an whic bah , urbinate head (the rays are ascending also), and the pubescent akenes, Collections of it ar no. 6717, € no. 5785, Druid peak, Yellowstone park, July 12; Teton mountains, Aug. 16, 1899. Laramie, Wyo. eeterER ARTICLES: PHOTOGRAPHY IN BOTANY AND IN HORTICULTURE. (WITH TWO FIGURES) ALMOsT every working scientist knows something about photogra phy, and probably no one will deny that a camera is one of the indis: pensable equipments in every well-regulated botanical or horticultural laboratory. Yet a casual acquaintance among various horticl 9 and botanists, and the repeated publication of unnecessarily inadequate (not to say atrociously bad) photographs, lead us to believe that the use of the camera in horticultural and botanical work will bear some discus sion. We.are convinced that the value of the camera as a piece _ scientific apparatus is not generally appreciated. We believe tha camera is not used as often as it ought to be, and we believe yet more strongly that the photographic methods with which many science ve €rs content themselves are not really creditable. We believe that 3 ® help in daily work, the camera ranks next to the microscope fo! a botanist, and far above the microscope for the horticulturist. i while every laboratory has a row of books on microtechnique and while every botanist and most horticulturists work hard to ii themselves. in microscopical methods, a book on photograp pee rarity, and men are satisfied to blunder along with almost any ge camera, and with plates and developers of which they know po nothing. Some even are willing to “press the button ” and ET company in another state “do the rest.’ In the matter of photographing plants, fruits, flowers, objects," we may perhaps offer a few observations as the result ’ ments covering several years. The particular piece of apparatus required for this ner rti some support which will hold the camera approximately in a ve dis- Position, and will provide a transparent horizontal shelf some es tance from the floor. The objects are laid on the latter sanhese 1s focused down upon them. and similar exper . : : ril 1900 : ; Photographing flowers and trees,” Zhe Photo-Minialure, N 5 ss SE 204 1900] BRIEFER ARTICLES 205 Any sort of construction which provides for the vertical camera and the horizontal shelf will answer, and several different forms are acimally inuse. Fig. 7 is sketched from a photograph of the appa- ratus in use at the Vermont Experiment Station. /ig. 2 shows amore elaborate form, designed by Mr. McFarland and used at the Mt. Pleasant Printery. The construction will be obvious in either case, and may be modified to suit individual needs. Lightness, rigidity, and portability are the chief requirements in an appa- — for field use; while in the studio the per- missible increase of weight makes a somewhat 7 convenient construction available. Fig. 2 renaoomed a studio camera-stand, adaptable for vertical : : as and horizontal work, making lan- es, enlargements, etc. ahe advantages of such an apparatus are: ty) It does away with nearly all trouble in rh eg the subject. Simply lay a plant me oO : 3 ? Ou th Wers, specimens of fruit, or other objects bat Pia shelf, and they stay where they are ut. In th . ; ‘ : REE Soe of si which will not always »,One s Fic. I. cine, -, robber rings : the fruit ; washers, z. ¢., rubber rings 3-4" in diameter. Set it into the ting, and i A ost any positi ’ it may be moved anywhere and put in desired. ‘Sheet on. (2) It allows one to arrange any background be success; . . cardboard in white, black, and neutral tints may I lvely tried behind th j t does away with sh e subject and the best one chosen. (3) disastrous in — shadows which are almost inevitable and frequently Considerably ae other method of plant photography. (4) It facilitates or all sorts of 4 work of making exact size photographs of objects. Ve all photogr i and fruit photography it is much the best way to This jg Oa (with a few necessary exceptions) made exact size. Negatives n y ¢esirable when one begins to have a collection of For ogarahegs into the thousands. orta : full Natural size = Plant, flower, vegetable, or fruit photography in tial, To obtai. camera with considerable bellows extension is essen- between the alec natural size image the lens must be equidistant focal len t and the ground glass of the camera at twice its sth. Thus iff »if fora plate of 614 x 8% inches a lens of 7% inches 206 BOTANICAL GAZETTE [SEPTEMBER focal length be used, sufficient bellows must be at hand to permit the ground glass to be fixed at 15 inches from the objective. The modem “long-focus”’ cameras provide very fully for this need and others which will appear upon trial. The best work will be possible if a lens is chosen of rather short focal length, one technically known as a “wide- angle” lens. Not only is the range of the camera thus in- creased, but the sharp focus over a considerable depth which is absolutely requisite in morphological work is far more easily obtained. In focusing, it will be found a decided advantage to select a spot in the object or composition upon the glass shelf a little above its vertical center, and there to affix tem porarily a white card or paper with fine lines. An ordinaty visiting card 1 When this is focused "po and the lens then « stopped down,” the whole object will be sufficiently shatp. We urge extreme and cleanliness in 4 graphic operation care ll photo enemy ; every grain is willing to be photographed. A skylight is most undesirable for vertical photography, P&€ the reflections upon the glass platform. Indeed, the worker will : . Imost essential to guard against reflections from the ceiling of a light room, and particularly from the poli front and lens, by preparing a wire frame or hood, covere Nec or the like, and extending about the camera SO 4S to cu tmmediate top light—F. A. WaucH and J. Horace because of find shed camer 5. In work: | ing full size, dust 1s 4 oe : skylight, d with black zo t off a . 4 CURRENT LITERATURE. MINOR NOTICES. T. Ivo aND J. Matsumura‘ have begun the publication of a catalogue of the flora of the Latcha Islands, the archipelago between Japan and Formosa. The present work, written in English, is intended to be a preliminary con- tribution to the knowledge of the flora of these islands, containing merely the bibliography, synonymy, and local and general distribution of the plants, with occasional descriptions and critical notes. The Leguminosae, including about 125 species, are specially worked out by Professor Matsumura, a number of new species being described. The flora is one of exceptional interest, and a " aaa and complete presentation of it will be welcome to botanists. region indicated by the title. The Fungi are by P. Hennings, Phacolimacium fea As pshegage ces (Hysteriacee), Phacorhytisma (Phacidiacez), tein, 3, and Discocyphetla (both Thelephoracezx), Pseudotrype (Hypo- Macez), J - heed (Melanommacee), Schizacrospermum (Acrosper- pie anseella (Stictacez), and Phacomacropus (Pezizacex) being new “1 Java. The Alge by F. Heydrich, and the Hepaticae by V. : ist, the latter group naturally having a very extensive abound in The et pteridophyte groups are presented by blertas especially oat Species and critical notes. The genus Selagine 4 A dectibed 4 ered, being represented by 184 species, forty-seven of w ie §ymnos s = author, and its geographical distribution 1S pare x Part is ful] pale sini close the volume, are also by Warburg, and t Z isttibution of tens oe as it does descriptions, fine paige 'Te any of the little known and more critical forms.—J. M. ©- Serene fom yoo Lutchuensis. Sectio I. Plantae Dicotyledonex Polypetalae. aad i ~ Coll. Imperial Univ., Tokyo, 12 : 263-541. 1899- i Monsungebietes ta zur Kenntniss der Vegetation des siid- und ostasiatischen M40, Oty pp. 207. pls. 2-17. 1900. Leipzig: Wilhelm Englemann. 1900] : 207 208 BOTANICAL GAZETTE [SEPTEMBER BRYOtES FOR STUDENTS, THE ‘“‘PROTEID VACUOLES”’ of gymnosperms have recently been rein- vestigated by Arnoldi. These structures were described by Hofmeister nearly fifty years ago as “ Keimblaschen” (germinal vesicles) and he makes the statement that the archegonium of the vascular cryptogams differs from that of the gymnosperms in having only one nucleus (Keimblaschen) while that of the gymnosperms has many. Schacht, who wrote at about the same time, regarded the structures as cell-sap vacuoles, while Strasburger, at @ ' considerably later period, regarded them as proteid vacuoles. Goroschankin (1883), who made some study of their development, found that they differ from the sap vacuoles, and make their appearance before the cutting off of the ventral canal cell, disappearing during the formation of the embryo. The present writer, with the aid of modern technique, has attacked the old problem of the origin and development of the bodies and finds that they are genuine nuclei. The nuclei of the cells of the tapetal layer immediately Surrounding the oosphere put out amoeboid processes which penetrate the cell wall,and soon the whole nucleus passes through the opening and into the cytoplasm of the egg, where they become more or less modified so that while they bear a striking resemblance to nuclei they could not be positively ident fied as such except by trating them back to their origin. Nucle! were als? observed to pass from the next layer into the cells of the tapetal layer. ™ paper is only a preliminary one, but the stages figured are strong “— favor of the author's view. These bodies serve only for the nutrition of \ embryo and take no part morphologically in its development.— CHAS: CHAMBERLAIN, es usin tele ive dicotyle hesporium; e me; ; , e func mother cell organizes a row of three, the lowest of which becomes th Honing megaspore ; the ante-fertilization development of the sac fertil '$ as usual; the antipodal cells are evanescent. After the ante Zation stage is reached the sac grows rapidly, especially in length, oadens below, remaining narrow at the top, thus appearing of the necked flask, with the primary endosperm nucleus lying at the = mains ip neck. At the first division of this nucleus one daughter nucleus . a 3 ARNOLD : Beitra . i We Weeca 1, W.: Beitrige zur Morphologie der Gymnosperme - Abiet nee? FI mblaschen”’ oder ‘“ Hofmeister’s Kérperchen” in der eT Ora 87; 194-204. pi. 6, 7. 1900. *On the development of Sawrurus cernuus L. Bull. Tort. Bot o 365-37 \ Bl. 23. 1900, o 1900] CURRENT LITERATURE 209 the neck, the other moves down into the body of the flask, and a wall is formed across the base of the neck, giving rise to two endosperm chambers. The upper endosperm nucleus divides and forms a‘ compact tissue in the upper chamber; while the other enlarges but never divides, the lower and larger endosperm chamber apparently being related to the adjacent peri- sperm as an absorbing organ. In the upper chamber the embryo is organ- ied, and the endosperm about it encroaches upon all the nucellar tissue adjacent to it. The germination of the seed is also very peculiar, the endosperm emerg- ing first, and retaining hold of the cotyledons and supplying nutrition after the cotyledon tips have carried the old seed well up into the air. The author concludes that there is no evidence that Saururus is more primitive m character than many other dicotyledons.— J. M.C. 9 ceca has devised an apparatus for research on the effect of air pressure on plant growth,’ in which the plants can be culti- ee eiereble time without being in a stagnant atmosphere and ee cS subjected to repeated evacuation of the culture chamber, Te ak are ls the use of a tubulated bell jar to which air is admitted §" a capillary tube, a water pump in continuous operation serving to exhaust the chamber more rapidly than the air can enter through the capillary passage, ' He find specially that of the leaves, while germination is retarded and shed O pressure. The acceleration of growth, however, is not (which really tends to retard growth slightly), nor to les- 2 Pressure, nor to the altered light, heat, or moisture. The only Seems to be that the plants absorb water more rapidly Sure (isolated cylinders of live pith do so), and conse- reach their definitive size more quickly, thus attaining earlier The copious guttation observed both by Débereiner aib ; © also accords with this explanation.—C. R. B. ITeus OF 8: 385-510, Fegion, bing numerous new species. NVeogoetzea Pax (Euphorbiacee), (Gesneriacex), Megalopus K. Schumann (Rubiacez), and and J. y. * Rania (Composite), are the new genera.— HENRY Deans are publishing in Proc. Linn. Soc. of South Wales a series *Fiinfstuck? i 82: 52, thas. S$ Beitr. z, wiss. Bot, 4:93-148. 1900. See also Bot. Centralbl. 210 BOTANICAL GAZETTE [SEPTEMBER of papers entitled “Observations on the eucalypts of New South Wales.” The last one issued (Proc. Linn. Soc. 4 : 612-630. pls. g8—50. 1900) is the sixth, and includes descriptions of three new species.— E. P. BIcKNELL (Bull. Tor. Bot. Club 27 : 373-387. 1900), in continuing his “Studies in Sisyrinchium,” has revived Salisbury’s genus Hydastylus to receive the yellow-flowered S. Calé fornicum, and has associated with it 11 other species, 9 of which are described as new.— K. M. WIEGAND (id. 388-391) has described two new species of Saxifraga and one of Primula from the northwest.—C. V. PIPER (tbid. 392- 401), in continuing his ‘New and noteworthy northwestern plants,” describes new species of Amedanchier, Potentilla, Saxifraga (3), Townsendia, Erigerm, Castilleia (2), and Salix (2).— KATHARINE BRANDEGEE (Zo€ 5 : 31-35: 1900) in her third paper entitled ‘“‘ Notes on Cactez,” describes two new species of Mamillaria, besides giving critical notes on several other forms.— C. WARS storF (Hedwigia 39: 100. 1900) has described four new Sphagna from Vit- ginia and North Carolina collected by T. H. Kearney.—J. M. ©. Mr. HAROLD WAGER has published the results of his study of the fertil ization of Peronspora parasitica,’ his conclusions being as follows : the prot: plasm of the oogonium differentiates as usual into vacuolate ooplasm and granular periplasm ; a receptive papilla is formed on the oogonium in contact with the antheridium and is penetrated by the antheridial tube: the nuclei o! the oogonium and antheridium undergo mitosis before fertilization ; soon alte! the delimitation of the oosphere the central body appears, which seems '° play some part in bringing the sexual nuclei together ; a single nucleus from the periplasm travels towards the central body, coming in close contact wit it, and towards it the antheridial tube advances and discharges # - nucleus ; fusion occurs while the nuclei are in the resting stage and ne a the oospore is nearly ripe; the central body disappears before fusion, ripe oospore is uninucleate ; no difference is observable between gs " : nucleus and those that remain in the periplasm, all probably being .. tially sexual; three types of fertilization and oospore-formation . oe €ronosporee may be distinguished, as follows: (1) uninucleate it binuclear fusion, and uninucleate oospore (?. parasitica) ; (2) unin é Sosphere, binuclear fusion, and multinucleate oospore (¢- eee Pett and P. Ficariae),; (3) multinucleate oosphere, multin idl pak and multinucleate oospore (C. Blitt).— J. M- C. a A PRELIMINARY paper by Lidforss’? records some interesting eae upon the chemotropism of the pollen tube. Varcissus Tasetta fu : material for most of the work. da piect If the pollen be placed in a 5-15 per cent. sugar-gelatin solution an ail : of stigma be added, the pollen tubes soon turn toward the stigma, a : tt . of Botany 14:263-279. p/. 16. 1900. — (vor Mittheilung.) 1 B.: Ueber den Chemotropismus der Pollenschlauch® | 8-) Ber. d, deutsch. bot. Gesell. 17: 236-242. 1898. SE ERS ey ee et ee IN Se ge ee Fe PE eg en ee ee Ma Pe Were tee eee Mere sr eee see igo] CURRENT LITERATURE 211 known. Lidforss finds that organic acids, formic, acetic, succinic, lactic, tartaric, malic, etc., as well as amides, glucosides, and tannins, do not produce any undoubted effect upon the direction of growth of the tubes. Diastase, however, produces an almost immediate effect, and experiments show that it is the proteid constituent of the diastase which attracts the pollen tube. Further experiments show that carbohydrates and proteids are the substances which influence the direction of growth and indicate that the movement of the pollen tubes is for the purpose of securing nutrition. The pollen of most Liliacez: is more sensitive to mineral salts than Narcissus pollen, the same diastase preparation which attracts Narcissus pollen quickly killing the pollen of Fritillaria, but if the salts be dialyzed out the proteid exerts a strong influence upon Fritillaria pollen.— CuHas. J. CHAMBERLAIN. In AN ADDRESS before the Niederrheinische Gesellschaft fiir Natur und Heilkunde zu Bonn,® Dr. F. Noll? makes a suggestion which is likely to prove fruitful, because it not only groups together some previously isolated facts, but —_ likely to be the starting point of further investigations. He has been led to believe that the form of the plant body itself is a source of both orienting a stimuli. As evidence of this he appeals to the fact that in aoe vesiaes the concave side of roots remains entirely free of lateral ie at se are limited to the convex side; or, if they appear at all upon nks, they bend more or less sharply toward the convex side. alae have been observed in all the plants studied, including are Naa all the great groups which develop roots. fe 4 Pelagia, — Noll correlates Wiesner’s ‘exotrophy,” a pics nh fanks i ee, ereby the growth of external members or their externa Picuously promoted as compared with that of corresponding Members 5 : P Polarity have a fields a : iso need exploring from this new point of departure.—C. RB; is “oskiigay by A. C. Hill that the hydrolysis of maltose by maltase tose from 9] process and that this enzyme is also capable of forming mal- seems to be supported by the results of M. Cremers - finds that the glycogen-free expressed extract of yeast is cing glycogen in a 30 per cent. solution of fructose, which 1s eg d. Sitzungsber. d. Niederrhein.. Gesells. 1900; Sitzung von Jan. “apable of produ *Sep. Abd "51900. pp, Noll’ : withichefticke 1 researches are announced to be published in Thiel’s pa nig € Jahrbiicher early in this year. ans. Chemical Soc. 1898 : 634-658. ha Berichte © d. deutsch, chem. Gesells. 1899 : 2062-2064. 312 BOTANICAL GAZETTE [SEPTEMBER a still more complex process than the production of maltose from glucose, Meyer suggests * that similar relations may exist among other polysac- charides which are hydrolyzed by enzymes. Holding that the enzyme which digests starch has its origin in the plastids,3 he now thinks it probable that this enzyme acts like maltase. When a concentrated solution of sugar enters the stroma from the cytoplasm, amylose is formed, which on reaching a cer- tain concentration crystallizes out in the form of a starch grain; whileif sugar is wanting in the cell a relatively active inversion of amylose occurs, and a rapid solution of the starch grains. The suggestion is interesting and deserves careful testing.—C. R. B. Mr. W. H. Lang has published the second number“ of his series entitled “Studies in the development and morphology of cycadean sporangia,” deal ing with the ovule of Stangeria paradoxa. His results are as follows: two ovules are developed on each sporophyll; the archesporium consists of a mass of cells, one of which is selected, enlarges very much, and forms 2 row of three, the lowest cell of which organizes the megaspore ; at the time of pol- lination the sac is full of endosperm (but without archegonia), the pollen chamber is fully developed within the beak-like process of the nucellus, but the breaking down of nucellar tissue between the pollen chamber and te embryo-sac has not begun ; as in Cycas and Zamia, the sperms are large, Spirally twisted, and multiciliate, with the blepharoplast band evident ; dis- organization of endosperm tissue between the pollen-chamber and embryo-sac gives free access to the latter, and the sperms are discharged into the archegonial chamber of the nucellus, reaching the neck by sem ming.— J. M. C, ‘ y be explained by double fertilization. H : | grain whose endosperm showed the characters of the pollinating os a hybrid embryo; and that every grain whose endosperm showed © acters of the embryo-sac parent had an embryo of pure race, and was fore self-fertilized. In a few words he puts it that double f corroborated by double hybridization.— J. M. C. ™ Bot. Zeit. 577: 313. 1899. "3 Untersuch. iiber Starkekorne? 169. ™ Annals of Botany 14: 281-306. pls. 17-78. 1900. ‘SSur la fécondation hybride de l’endosperme chez le mais. Rev. gén- 4 ea Mique 12: 129-137, bl. 15. 1900. i ertilization 5 Se ee 1900] CURRENT LITERATURE 213 GrorGE MASSEE, in discussing the origin of the Basidiomycetes,” presents the following summary: (1) in the conidial condition certain ascigerous fungi bear their spores on structures morphologically indistinguishable from the basidia of the Protobasidiomycetes; (2) some members of the same form genera as those described in (1), as St/bum vulgare Tode, have lost the ascigerous condition from their life cycle, and are accepted as true Proto- basidiomycetes ; hence we are justified in concluding that the Protobasidio- mycetes as a group originated from ancestors that represented the conidial condition of ascigerous fungi; (3) there is no evidence in favor of the Suggestion that the Autobasidiomycetes are descended from the Protobasidio- ees on the other hand, the evidence in favor of the Autobasidiomycetes hrs been derived by gradual modification of the spore-bearing organs, or basidia of conidial forms of certain ascigerous fungi, is not lacking.—J. M. C. he W. Toumey has published the results of his study of “crown-gall” as “ ag 33 from the Arizona Agricultural Experiment Station, under the title ia Pate into the cause and nature of crown-gall.’”’ The name is applied fruit pao appears as fleshy outgrowths on the roots of deciduous , sually at the crown. The author concludes that the cause 1s a rc a one of the Myxomycetes, parasitism among which has shin Causes « xis pinned to a single species (Plasmodiophora brassicae, which effect upon i. in cabbage and allied plants). The plasmodium and its study of the or ie cells are fully described, as well as the sporangia. The ganism is made the occasion of the establishment of a new snus, Dendrophagus.— J. M. C. moo E FUSION has been discovered in connection with the Plying. First gai endosperm nucleus, illustrations are rapidly multi- for the aaa by Nawachsin for Lilium Martagon, it was Con- other species of "te Species by Gui gnard and Miss Sargant, the former adding a aS a os and also Fritillaria and Tulipa. Now Miss Ethel N. Palustrig x7 CO nuclei wrapped about the polar nucleus in Te embryo-sacs 5 awaschin has described and figured ® triple fusion in the (an orchid), eee annuus, Rudbeckia speciosa, and Phatus Blumet Warn ST. F te ber of the : ag 8ives a description of the anatomical characters of a num- With the Seti of Sphagnum which have not previously been examined this group egg and minuteness which modern taxonomy demands !n . e age a characters are drawn from types, and the examination of "7 Jour. Linn ’ - Soc, 34: *438-448. pls. 15-76. Annals of : 448. pls. 75-76. 1900. "Ber. dp any 14: 318-319. 1900. if eutsch ” Bot. Cont. * bot. Gessell. 18: 224-230. pl. 9. 1900. 82 :7 Sq. 1g00. 214 BOTANICAL GAZETTE [SEPTEMBER these has led to the reduction of many of the recently proposed species, par- ticularly those of C. Miiller, whose later work was so prolific of “new” species. Some of the critical remarks touch American and Antillean species. R. B Von Derscuavu has studied carefully the process of wall thickening in the formation of the teeth of mosses.” He finds that preceding the true thickening process, the activity of the cytoplasm consists only of its prelimi- nary accumulation on the membrane to be thickened. In this the nucleus exerts no clearly recognizable control, but does do so in the thickening process itself. This consists of the apposition of materials early produced in the cytoplasm, of which cellulose is the primary one, the other substances being such as promote hygroscopicity and resistance to decay.—C. R. B Dr. GeorG Goetz has restudied the development of the egg in the Characee,** which leads him to consider the group as independent of the alge and derived from the primitive type of the archegoniates, just as the mosses and ferns, The Wendungszellen of Nitella he regards as a reduced archegonium wall, and the peculiar separation of a portion of the nuclear substance reminds one of the formation of a ventral canal cell.—C. R. B. Dr. F. NoLu proposes™ the use of the scape of dandelion for demon Strating the mechanics of tendril coiling. By cutting out from the scape # long strip not much wider than thick, and, after fastening the two ends Vd that they cannot rotate, immersing the preparation in water, the inner tissues elongate so greatly that the strip is thrown into a spiral coil with one or more points of reversal, thus imitating very closely a huge tendril.—C. R. B. ACCORDING to Palladine,*3 though light is not requisite for the regenerate of proteids, if cane sugar is supplied to the etiolated leaves of Vicia under experimental conditions, the regeneration goes on more €n in light than in darkness, and for this the more refrangible light is ¥ efficient. Such leaves cultivated on cane sugar solution in light respire more than twice as actively as when kept in darkness.—C. R. B. . Dr. Oscar Loew brings together a useful summary of the pit knowledge of the physiological réle of mineral salts.™ Unfortunately Sreater part of the earlier experimentation in this direction has been more lees misguided, and one must hold very loosely the conc This summary, however, will be useful as a guide to the liter subject.—C, R. B, ergetically s the more » Bot. Cent. 82: 161-168, 194-200. 1900. *Inaug. Diss. Freiburg. 1899. See Bot. Cent. 81 : 366. 190 ™ Flora 86 : 388. 1899. 73 Revue gén. de Bot. 11:8 O.: “The physiological réle of neineral nari een EW Dept. Agric., Division of Veg. Physiol. and Path. 8vo, pp. 60. Printing Office, 1899. 0. 1-105. 1899: 1900] CURRENT LITERATURE 215 OF GREAT INTEREST to all botanists is the report of Henry Gannett*> on the forests of the United States. The author as chief of the Division of Geography and Forestry of the U.S. Geological Survey has had long and intimate acquaintance with the subject. It seems that on July 1, 1899, there were thirty-seven government forest reserves, aggregating 72,139 square miles, composed mainly of mountainous, rugged country, of no value for agriculture, but especially favorable for tree growth. The states containing these reserves are Arizona, California, Colorado, Idaho, Montana, New Mexico, Oregon, South Dakota, Utah, Washington, and Wyoming. Of these Washington has much the largest proportion, 19 per cent. of the total area of the state being thus reserved. The bulk of the publication consists of RS of reports on forest reserves by special agents sent to examine them. —|, as To THE List of seeds whose germination is affected by light, Heinricher adds our common Veronica peregrina” Light, even weak light, accelerates germination, as a difference of five to eight days between light and dark cultures strikingly shows. The effect of the light depends eer upon its action in promoting the digestion of the reserve foods.— B. OsWaLD RICHTER recommends” a concentrated solution of ammonia, ‘sed boiling, at 4o° C., or cold, as a maceration fluid. He finds it much Nperior to the usual acids, because the cell wall is always intact. In many “ses, also, the cell contents are preserved and even made clearer. —C. R. B Pei le Dr. S. Rosrowzew has devised,* apparently independently, Siderab] cide table with trapezoidal top, identical in form with that con- first 5 Yused in this country when window space is scant. The form was “ggested, we believe, by Dr. C. E. Bessey.— C. R. B P “Extract from Aiea Ann. Rep. U. S. Geol. Surv., Part V, Forest Reserves. 2 37, with 7 map: 900 pe d, Misa bot. Geséils 17: 308. 1899. terr. Bot. Zeitsch. 50: 5-11. 1900. *8 Bot. Cent. 81 : 361. 1900. NEWS. ; Dr. E. B. COPELAND, assistant professor of botany in the University of West Virginia, has been advanced to a professorship. Dr. HuGo ZvuKAL, professor of plant pathology in the Hochschule fir Bodencultur in Vienna, died on February 15. (Bot. Cent.) THE OFFICERS of the Botanical Society of America for I1g00-I are follows: President, B. D. Halsted; Vice President, R. A. Harper; Zreasurer, C. A. Hollick; Secretary, G. F. Atkinson; Councillors, B. D. Halsted, R. A. Harper, C. A. Hollick, G. F. Atkinson, B. L. Robinson, C. E. Bessey, and F. V. Coville; Custodian of Library, W. Trelease. TuE Division of Vegetable Physiology and Pathology has just completed : an extensive series of experiments at Halstead, Kan., in connection work on the development of new forms of cereals by breeding. The wor was planned by Mr. M. A. Carleton, but owing to his absence abroad it was carried on by Mr. D. B. Swingle, a graduate of the Agricultural College a Manhattan, Kan. Wo. J. Fox, of the Philadelphia Academy of Sciences, has had the § fortune to discover in the library of the academy a copy of that work of Rafinesque entitled Western Minerva, or American Annals of ee edge and Literature. Rafinesque proposed to publish a journal pie Science of August 10, 1900, It isa small quarto and contains vit oo and is of interest to botanists in that it contains new names for plants have not yet been noted in Synonymy. It is an interesting quel - a work suppressed by the printer, and presumably never distributed, : counted as a publication. In the copy discovered by Mr. Fox some © Pages are orginal proof sheets, being printed on one side only, © i jon of ee hot come within the definition of a publication when the ee : ority is concerned. Only four pages are given to botany, and Mr. F pe very fully their contents, Horsford’s Acid Phosphate The most efficient remedy known for the relief of languor and exhaustion, so common in the spring and summer months. Taken after exhaustive illness ‘ acts as a wholesome tonic, giving renewed strength and vigor to the entire system. Taken before retiring, quiets the nerves and induces refreshing sleep. Sold by Druggists. P senuine bears n a me Horsrorp’s on the eee ’ — in a new size 23. of the Liquid The event of the year in dentifrices. Beware of counterfeits getting the genuine at the stores. if necessary send 25c. direct to Se Proprietors, P. 0. 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Do not spoil their plenty ¢ of Quaker Oats. t porrid dee in the world is made from Quaker Oats, es = ee that Quaker Oats also makes Our Cer ead, oa ‘Cea. ups and Puddings. At Grocers in 2- Ages a wet eaiteren re ook Book, edited by Mrs. Rorer, gives hundre . ua innovations and valuable recipes. Write for i tas 9 ES CEREAL Co., Monadnock Bldg., Chicago, Ill. 2 a | a Pencil matters little, So long as it does not brea or crum- he quality is smooth and yielding, DIXON’ Sirk PENCILS i smooth, tough not b school wor the ee mples wor Souble the money will a sent on receipt of AM cents, if you mention this par a Catio Joseph Dixon Crucible Co., SEY ¢ nds of k, = ie are indispensable ; in iv }| Call on nearest representat | THE HAMMOND TYPEWRITER CO., Every Clergyman and | Every Public Speaker should ag Hammont 7 ypewriel their special ust e intended for da cha perfect satisfaction. Type other size a u shes vate : ast. 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A. upon application. ‘eit. a Allyemeine Botanische Zeitschrift Stematik, Floristik, Pflanzengeographie etc. . mit i nische rt bringt vor allem Abhandlungen tber schwierige Pflanzen- CRAKS inteross Riadas ¢ ten Formen und Bastarde, Seg gl frristisch und pflanzen- Mancengeogra sy nier Gebicre, botanis me golf genst Refera tiber wetriomgs 0 florists ae - “oo br abhische Arbeit : det, deren Ziel u. a. “lie yrs usgabe won ye cation werken rbeite ahrgang 1899 wurde iter Mitwirkng von waikern herausgegeben, ron 8 bot. Verei n, 31 Referate, Inhaltsangaben von 20 bot. ng Pitch re "berichtet iiber spate en » Anstalten etc., ther 32 Tauschvereine und Exsiccatenwerke, tber 15 bot. zur Kenntnis der Redaktion gelangende Personalnachrichten von stand eucine botanische Zeitsch f d mit erse ift” erscheint piinktlich am - jeden monet gehe tet un Abonne® Yersehen in minde ea pi J hrien nten ” portofrei unter > seta cet ae er ee ee as rael oe * Der Herausgeber: A ucker 3. Karlsruhe in Baden (Deutschland), gre 4 @ Keke Glumaceae Exsiccatae. Glum, ae oH den ‘Carices exsiccatae,’’ werden auch Mitarbeiter aus ~ apap thea ; iche he Eat, o Siedern sich in Gramineae, Cyperaceae et Juncaceae xs. er ITO ge : ; : ner 1a einsendet, erhalt 1 Lief. als Aequivalent; bei den Carices sin ‘5 etzt sind hie ra ges : A ief. ist te br Schiive beige od en, und die E é ); Prof. pr. Ite Material wird ausser dem Her : = usgeber kritisch bearbeitet von = nen noch Buchenau th gnome Dr. Ps ails (Cyperaceae), Dr Dr. Atterberg (Getreide- or ~ Toate pe und ckowi Wer einsenden wi wende sich n Karlsruhe in Baden (osu nat Werder atz 48. Every Botanist Should be familiar with the prominent works of GEBRUDER BORNTRAEGER Publishers. Symbolae Antillanae Seu Fundamenta Florae Indiae occidentalis, edidit Ign. Urban. 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HE object of the ELemenrary Scuoot Recorp is to make possible for use in other schools the details of « subject-matter and method in the application of mod- sit fe relegy i in education, as demonstrated by the Univer- iv oo eet School. Each number contains a recor con r& done by a group in the school, and also an article cerning the work of one department in all grades. Subscription per year, $1.25; single copies, 15 cents Sent postpaid on receipt of price by the publishers aE NAA COLL ELORET The Botanical Gazette Baie KES ME \ ci tr Ot ee bitahenents throu Rite bio INK WELL 6O., 131- University Press Dj vision, Chicago, Ill. Pw Pp 4 John Pe Coult | inions think this daily. Don’t you? You © wilateays Arthar uiter, C, R, Barnes and J. C. have clean ink and a clean pen Monthy Merican and f atio nal ys at st oreign associates. . d. etce of Bo 0 pages, Devoted to the Automatic Ink Stan ‘ining tany in all its d Gravity w orks it, never ou odie’ nN epa order, always ready, non-evapo a of Tese rch book P aici com tae ene Gant proof. Lift erg , rev coy, ises instantly free fro: 50. i S$ items. ee notes for ate rene pier: be ins Ee ; hi} j en automatically eats m Copies, 40 ct: ear; foreign, peat dust tight. The most perfect and a good reason for aa | for sa Ss. All idicbotieas and ¢ inkstand made” A for Your Petiment and leading commercl ™mple Copies h it in its place. In use by U. 8. vor of Aluminum Roman Should be addressed to establishments Working P: al because of Univers: to tay and automate perfection. feet finest Si its uty and autom4! roo By return orward ty. of Chicago — Us 506 im stand complete guaranteeing § sate will rem guine expec’ ie THE SCHOOL AND SOCIETY. I THIRD EDITION. A MOST IMPORTANT BOOK FOR PARENTS AND TEACHERS By JOHN DEWEY. PROFESSOR OF PED- AGOGY IN THE UNIVERSITY OF CHICAGO ee N this book, which has been characterized as “one of the most noteworthy books of the year, in the field of pure pedagogy,” the educational situation is clearly stated and graphically illustrated, and new light is thrown upon some interesting phases of educational reform. The problem of elementary education is one of special importance to teachers, parents, and school boards, and Proressor Drewes clear exposition of the relation of t school to the larger society and the every day world cannot fail to be of interest. The “fads and frills” of the Public School main despite their critics, but they cannot be a : ; ost assimilated. PROFESSOR DEWEY gives 2 of these Tt has luminous statement of the meaning branches for the school and for life. been his good fortune to have stepped out of the field of theoretical pedagogy and t0 stand upon the successful results of four years’ experimentation in the Elementary School of the University of Chicago The ideas behind it and the methods of applying them are presented here in a stylé neither abstruse nor technical. “ . 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A special monthly ir ee, size, is issued the first full week in eac “ UNIVERSITY OF CHICAGO PRESS CHIC ALO. 13 TABOOS Issued through its Ordinary Department Alone LAST YEAR OVER Fifty-six Million Dollars of Life Insurance. Its policies are Pe Ss O in amounts With premiums ses yearly, half- aa or quarterly. PLANS INCLUDE ; PROVIDING FOR Cash Values Cash Dividends, Cas a Policies, Partner- Cash wage fr aid-u “Fasurance Ship Policies and Annuities. Extended T Many other attractive and profitable features. Write for el HE PRUDENTIAL INSURANCE COMPANY OF AMERICA OHN F. DRYDEN, President, Home Office: NEWARK, N.J. ; NUMBER 4 SOTANICAL GAZETTE ; OCTOBER, 1900 AND NUCLEAR DIVISION IN FULIGO VARIANS. R. A. HARPER. (WITH PLATE XIV) knowledge of such important vital phenomena of the Cetes, as the formation of the spores and the capilli- im, and nuclear and cell division, is still based quite largely the data given in the single paper published by Strasburger (23) in 1884, ‘Strasburger Studied Trichia fallax, and his main conclusions Mat the nuclei divide karyokinetically just before spore med by simultaneous breaking up of the multinucleated eg hyaline lines into uninucleated spores. Strasburger «Ss in the development of the sporanges simultaneously. Mentions also that the sporangia require several days to after their first appearance. a describes the capillitium, which in 7richia fallax lon 217 ‘Material on decaying stumps, and speaks of finding ‘on, the capillitium is formed in vacuoles, and the spores 218 BOTANICAL GAZETTE [OCTOBER microsomes in spiral lines about the so formed thread leads to the formation of the spiral ridge-like thickenings of the mature capillitium. The whole process is identical with that which takes place in the formation of a cell wall, according to Stras- burger’s earlier accounts of that process. Whether or not the microsomes are the units in cell wall formation, we have here excellent evidence that the interior protoplasmic surface, which lies next to the vacuole, is equiva- lent to the exterior surface which forms the peridium of the entire sporange. Each surface is able in essentially similar fashion to deposit on occasion a resistent membrane over its whole extent. The doctrine of the equivalence of plasma-mem- brane and vacuolar membrane as developed by Pfeffer and De Vries finds strong support in this method of capillitium forma- tion. Strasburger describes a period of nuclear division as pre ceding spore formation. The division is karyokinetic and the equatorial plate, separation of the daughter chromosomes, and development of the daughter nuclei are figured. The spindle fibers are inclined only slightly toward each other at the poles. The nuclei all divide at the same time, so that each section shows thousands of karyokinetic figures. Strasburger S4J° little as to the method of spore formation. He figures the cleavage as producing the one-nucleated spores directly, describes the boundaries of the spores as at first consisting . granules and then of clear lines, and notes also that the . : entiation proceeds for the most part from the periphery eer a the center. The young spore is at first polygonal, then ro itself up and becomes enclosed by a wall. Zopf (27) adheres to the view that the capillitium repre : plasma masses (Hyaloplasma, Gerustplasma) which are ie ‘ used for spore formation and have become hardened. He 4 not pretend, however, to have verified this statement for a 5 self. Later (p. 63) he accepts Strasburger’s account vegatd ) formation of the capillitium in Trichia, and is inclined po it as true for all the forms with hollow capillitia (Coco while still holding that the forms having a capillitium com of 190] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 219 of solid threads (Stereonemata) form the latter from strands of protoplasm. As a matter of fact, both forms of capil- litium may arise in vacuoles, as I have been able to determine in the cases of Stemonitis and Lycogala. The account given by Strasburger is correct, and I have not thought it necessary to give further figures at this time. Whether the thread is hollow or solid, simple or branched, free or connected with the peridium ora columella, are entirely secondary conditions, depending on the extent and form of the vacuoles. Massee neglects Strasburger’s account of capillitium forma- tion altogether, and advances the extremely loose and erroneous view that generally a surplus portion of the protoplasm takes the form of a more or less complicated network mixed with the pores, and homologous with the strands described as being Present in the sporangium of Mucor, inasmuch as both struc- = are made from a substance separated from the protoplasm during Spore formation. I have shown (4) that the so-called - “tsporal protoplasm of the Zygomycetes is merely excreted slime, and Strasburger’s account of capillitium formation is true ae all slime molds that have as yet been carefully investigated. Lister has discovered karyokinetic division of the nuclei in i. oF = considerable number of genera, and has “a a aryokinetic figures in the dividing swarm spores and © plasmodia, though he also figures direct division cell formats at this latter stage. He concludes that whenever Mucle; ay eer in the life history of the Mycetozoa, the de by karyokinesis. Lister also observed in certain Se a pe i-tasces containing six to ten nuclei in the” into a. ll and describes these masses as separating division, “Sig . during the succeeding stages of nuclear Sr acca = va Genera he confirms. the account of Stras- = SPore formation ¢ ss which nuclear division is complete before : B egins. formation ae, follows Strasburger in describing the spore € whole group of slime molds as taking place : by simu] | : taneous breaking up of the protoplasm of the sporange 220 BOTANICAL GAZETTE [ OCTOBER into uninucleated masses, and this is the current statement of botanical text-books. Rosen (17) made a very thorough study, as he describes it, of Fuligo septica. He finds nuclei of two kinds, one poor in con- tent and containing a so-called middle-body, and the other so densely filled with stainable substances as to appear almost homogeneous. The relative number of these two types of nuclei vary at different stages in the development of the slime mold. Nuclear division occurs prior to spore formation, but the process is described as much simpler than in the higher plants. Rosen thinks it belongs to the karyokinetic type, but it is doubtful whether a spindle figure is formed, etc. The cleavage 1S post tively stated to be simultaneous, and to take place by the depo- sition of a network of granular plates which cut the protoplasm up at once into polyedric uninucleated spores. These plates are said to show microsomes very plainly, the latter being placed at right angles to the plane of the plate. As will be seen below, my own observations on Fuligo have led to entirely different results from those of Rosen. es convinced that his two forms of nuclei are due to inequalities in fixation such as sometimes occur. As to the method of nuclear division and spore formation I am certain that Roset failed to find material in the stages when these processes ce His description of nuclear division must have been based 0" resting nuclei whose contents happened to be somewhat yer ally placed. As for the network of granular plates with mict” somes such as he figures, 1 am convinced that no such struct , “are to be found in Fuligo at any stage of development. i iy difficulties in the way of obtaining accurate results in ee 4 of fungus cells and nuclei are great, but not sufficien such slipshod results as those of Rosen in the paper sideration. a A summary of our present knowledge of the Myxomy""" has recently been published by Jahn (g), and agai h be made to it for a further account of the literatur® : group. » under cof 1900] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 221 My material was fixed in Flemming’s solution, weaker for- mula, sectioned, and stained with Flemming’s safranin, gentian- violet and orange. The formation of the aethalium of Fuligo has been very well described by De Bary so far as its grosser structure is con- cerned. The plasmodium which is ready to form spores creeps to the surface of the substratum, and there forms a reticu- lum which is similar to that of the vegetative condition except that it is more dense. It becomes a rounded cake-like mass, the meshes of the reticulum being relatively small and the protoplas- mic strands very thick. We have in fact at this stage a con- vented reticulum, the interprotoplasmic spaces having become minute. At this stage the solids which have been imbedded in the protoplasm are all thrown out upon its surface. Large amounts of water containing salts in solution are excreted, and, the Water evaporating, the salts are deposited as crystals along with the rejected solids. These waste materials are found in all the meshes of the protoplasmic reticulum, and form a sort of fragile framework piercing the ripe aethalium in all directions. The yellow coloring matter of the plasmodium is also trans- ferred to these waste materials, so that the protoplasm is left “pparently homogeneous and colorless. _ A further step in forming the aethalium consists in the con- a contraction of the protoplasmic reticulum, so that its 6.55 cial strands are withdrawn toward the center. In this reanle of the protoplasm from the peripheral parts of the bible ° phage wastes are left behind, and form thus a ae Muipins of ees the surface of the protoplasmic mass. on e ‘i Bes oi e aethalium this waste material frequently a forth. oats thin membranous almost papery border. In es atinic « a of the protoplasmic reticulum the interpro . - Or plates - become reduced in many cases to mere Q other cas € yellow colored waste materials described a _ lineq ith «thi the spaces remain as oval or angular ce : section th in yellow crust of the same excreta. Fig. 1 shows rough a portion of an aethalium relatively free from 222 BOTANICAL GAZETTE [october such cavities. In fig. 2, from another portion of the same aetha- lium, parts of several lacunae are shown, and their relative size and distribution is thus partially indicated. In addition to the solid particles and solutions thrown out, the protoplasm excretes over its whole surface a thin fragile membrane, in which the crystals of lime are frequently partly imbedded. This membrane is by no means as thick as in the case of the slime molds which produce sporanges, but it appears very clearly in microtome sections. In most regions it is hardly more than a cement to hold together the lime crystals in a continuous film. In other regions, where these are less abun- dant, it appears as a very thin homogeneous membrane. It lines all the interprotoplasmic cavities mentioned above, as well as covering the peripheral portions of the protoplasm. It: is always next to the protoplasm itself. I have never found crystals or other solid excreta between it and the plasma membrane. At a stage when cleavage is just beginning, such af tit shown in jig. 1, the nuclei are generally in the resting condition, and are distributed rather unevenly through the cytoplasmic mass. Frequently they appear aggregated in rather dense Sroups in certain regions, while in adjacent regions of the bees . plasm they are less numerous. Spore development now begits : with the formation of cleavage furrows, which usually arise first _ on the external surface of the entire aethalium and cut down @ all angles into the homogeneous protoplasm. These furrows a very narrow and sharp in some cases, and quite widely opens’ — in others ( fg, rT). This latter condition may be due, at leas - partly, toa slight shrinkage in fixation. Very commonly 6! are curved and forked so as to cut off a superficial layer “ fi ments. Almost simultaneously with the formation of these ae FOws on the surface of the entire aethalium, similar furrows #° : formed on the surfaces of the lacunae of the contracted ahs plasmic reticulum as described above. These surfaces, of mee are in reality external surfaces of the protoplasm, and . formation of cleavage furrows from them is not in any Sense ~ Ig] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 223 be compared to the cleavage from vacuolar surfaces as I have described it in Pilobolus (4). As noted above, the first cleavage furrows commonly do not cut through the entire mass of protoplasm in which they are formed, but curve and fork so as to cut off one or more super- ficial layers of segments. Further furrows, not continuous with the first, then cut through the central mass, dividing it up into large blocks, each with many nuclei. Meanwhile the superficial segments have still further divided, so that we regularly have one or few nucleated masses at the surface, while the central protoplasm is relatively undivided. The segmentation is very peny.» progressive process proceeding from the periphery toward the center. There is no such thing as a simultaneous breaking up of the protoplasm into uninucleated fragments. The protoplasm which thus segments is quite homogeneous, as noted “ove. There is no differentiation of hyaline zones or other peed regions prior to the formation of the cleavage furrows. Furthermore, the nuclei show no special distribution about the tleavage planes. As seen in figs. z and 2, it is quite common to find a group of nuclei on one side of a cleavage furrow while — oe dead a considerable area on the opposite side. any fee indication whatever at this stage that the nuclei exert influence on the orientation of the cleavage planes. ‘we examine the protoplasm immediately in front of one of any > diag pitrows also, we find it without differentiation of will one et ag indicate the direction which the furrow to advance al ide from the fact that it is common for the tiie’ it is quite 8s the same plane or curve in which it has ~~ : OW, i. ¢ « scgeateesheg predict in the case of any yer e in which Mic which has not yet cut through the protop si able that iy sas direction it will take. It is very notice Which is se x ates currows do not necessarily cut the pn through RoR one through its shortest axis, any more : . Cut off oar & = It is very common to see a strip or Ss - Plane of ag ae of 4 larger mass in such a fashion that : age lies in the long axis of the mass which is thes 224 BOTANICAL GAZETTE [ocTosER divided. I have even found some evidence that sausage-shaped Masses are cut out of the center of larger masses by means of a cylindrical cleavage furrow. It is very common to find semi- cylindrical masses cut from the surface of the protoplasm by two furrows which curve toward each other so as to form a trough-shaped cleavage surface. All of the above varieties as to form and direction of the cleavage furrows are illustrated in jigs. 1-4. At and immediately prior to the time when cleavage com- mences in Fuligo, its nuclei are all in the resting condition. None of the nuclei indicated in jig. t were dividing. A very little later, almost simultaneously with the formation of the first superficial cleavage segments, the nuclei throughout the entire aethalium begin to divide karyokinetically. In some cases it may be that the peripheral. nuclei commence to divide earlier than those which lie deeper. But the difference, if it exists, generally is a very slight one. On the other hand, the process of division seems to begin progressively rather than simultane ously in different parts of the aethalium, regardless of depth from the surface. This is shown by the fact that in examining sections different stages of karyokinesis are found in different parts of the same section. For example, all the nuclei in a cer tain region a few hundredths of a millimeter in diameter may be in the equatorial plate stage. Moving from this region in one direction one will find a gradual transition to the anaphase stages. Moving in another direction one may find prophases, or one may find nuclei in anaphase on all sides of a region oe ing only equatorial plates. There is no constancy in the order of stages which will be found in moving from the peripheral ” the central or deeper portions of a section cut radially to the ” face of the oval, cake-shaped aethalium. It is an absolute peo . however, that widely separated stages in division Bae found in close proximity to each other, at least in a“ , masses of protoplasm; and generally, passing over one : alium humerous lacunae, which, as noted above, pierce the aeth ‘- a in all directions, does not involve any sudden transition 1? 4 i900] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 225 stage of nuclear division. We may assume that the nuclei begin to divide at numerous isolated points in the aethalium, and that the nuclei of adjacent regions begin their division progressively inall or only in certain directions. If we consider that the division begins in response to a stimulus either external or internal, we should imagine the stimulus being propagated in one or several directions, from the point of its first effectiveness. Strasburger has remarked upon this same wave-like progress of the tendency to nuclear division in Trichia, and has compared the phenomena there with those in the division of the nuclei in a young endosperm of Fritillaria, in which the nuclei at one ead of the embryo sac begin to divide first, and the process is ‘een taken up progressively by the successive nuclei through the whole length of the endosperm layer. Nuclear division proceeds thus during the whole process of eranege, but without any relation whatever to the latter process. itanai figures can be found oriented in all possible ways ‘cc iad furrows described above, and any stage in divi- shape or : cep in segments of the dividing cytoplasm of any eh a as will be seen from figs. 4-9. As a further exam- ha oa can be found with a single nucleus in any stage of divisions Ang TI, 13, 15, 16). The details of these nuclear described bel ar as I have been able to work them out, will be ow, —- geal of nuclear and cell division in Fuligo are described b a puitely different from those in Trighi as Plete ieee at In Trichia nuclear division 1s com- “SSS are carried en begins, while in Fuligo the two proc- sociated with = simultaneously. The difference may be ¥ of Fuligo a ies for more rapid ripening of ne sae aethalium oe ecording to my observations, the building : tWenty-four ay formation of the spores takes place within State just bow “ ae is case of Fuligo. Strasburger does oe © Sporange of eg time is required for the development te) richia, but Lister found that they require from two to f ‘four : : days to ripen after their first appearance. 226 BOTANICAL GAZETTE [OCTOBER We may return now to the further consideration of the cleav- age processes by which the aethalium is cut up into spores, We have noted above that the primary cleavage furrows cut into the surface of the protoplasmic mass at varying angles, that they may be curved, may branch, etc., in the most irregular fashion, with no reference whatever to the distribution of the nuclei This much of regularity, however, can be seen. The furrows are so oriented with reference to each other and to the surface of the mass that cleavage at first progresses more rapidly at the Surface than in the center. Thus, one or more layers of very , irregular one- to several-nucleated segments are cut off on the periphery, while the central mass has been cut through by only a few furrows. What is true of the exterior of the aethalium as a whole is also true of the surfaces of the interprotoplasmic gaps or lacunae. It can be seen from jig. 2 that the surface of each such lacuna is lined by a layer of one- or few-nucleated seg ments, while beneath them larger multinucleated segments are found. The peripheral segments are very irregular in shape, 8 are also the larger central segments. Frequently broad thia plates are found; elongated sausage-shaped masses are also common. As noted above, the cleavage planes follow no such simple rules as cutting through the short axis of the mass yee divided, or always dividing a mass successively in planes th intersect at right angles. Hofmeister’s law, also, that cell oe sion always occurs transversely to the axis of most ia 38 . growth, has no application here, since no growth is taking ae : at the time when these divisions occur. ,3 i If we study the cleavage of any one of these central m << of protoplasm we shall find the orientation of the furrows - tially similar to that of those which cut in from the Bee is the entire mass. Fig. 5 shows such a mass with its yee the the equatorial plate stage, both polar and profile views : 4 latter being shown. It is to be especially noted also that # ‘ furrows in Jig. 5 are not directly continuous with - seg : appeared first on the surface of the mass from which ee ment in question was taken. No single furrow can be i900) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 227 for any great distance in an unbroken plane or curve. The fur- tows which are to further subdivide the segment represented in fig. 5 have been formed independently of those by which the block itself was delimited. The plan seems to be that each fur- fow may cut through the mass in which it originates, but may fot continue across the furrows with which it intersects so as to cut through successive masses in any specific direction. There ae no general planes of cleavage for the whole mass. As noted already, the first furrows that form do not asa rule cut deep | down into the mass toward a center. Rather they branch or are curved so as to cut off irregular blocks on the surface. © New furrows forming on the surface of these, and at very varying angles with them, continue the cleavage into the deeper portions of the mass, : is well shown in fig. 5 that the furrows at this stage also ae into a perfectly undifferentiated mass of protoplasm, there being absolutely nothing by which to predict the path they will take except the general direction which they have already a. 0 The protoplasm: is singularly homogeneous, this i vacuoles or inclusions of any sort, and through further © ce aabaie mass these furrows are formed. It is Various angle a they may be either plane or curved, and lie at ~:, © > With each other and the surface of the mass. a to eile results in no ‘ee definite conditions When the Saar pemients formed. Still we can find a stage uninucleated oo segments of the mass are quite regularly Mensions nA the deeper portions have been cut to vanoue far the ies. containing from eight to sixteen nuclei. So at this ad esembles that already described for Synchitrium ; division tg a very noticeable difference nr the method of ey. ... = con This difference is shown in figs. tiation Of the - itherto, there has been absolutely no differen- . Cleavage “eee otoplasm to mark the path to be taken by the between each ae nOW: broad hyaline areas are formed midway : line zones of 6d dividing nuclei. These are not at all hya- : €qual thickness, but furrows broader at the surface oe 228 BOTANICAL GAZETTE | octoser and narrowing toward the centers between each pair of nuclei. The surface is generally slightly depressed in the line of these hyaline regions, indicating the beginning of the furrow which is actually to sever the portions thus preliminarily marked off by the hyaline regions. The appearance is as if all the denser portions of the proto- plasmic mass had contracted about each nucleus as a center, thus leaving irregular, furrow-shaped, less dense spaces in the middle region between each pair of nuclei. These areas contaif very little or no stainable material, and seem to be filled with a watery liquid merely. They are, however, not in any —_ rounded vacuoles whose cell sap shows surface tension where it comes in contact with the denser protoplasm. The surface of the rounded mass of protoplasm aggregated about each nucleus is by no means smooth and even, as is the surface of the prot plasm about a vacuole. The denser protoplasm. passes over i insensible gradations into the less dense material in a fashion very hard to reproduce in a drawing. Peripherally these hyaline areas are bounded by a very thin protoplasmic film consisting of little more than the plasma membrane itself, which can here be more perfectly recognized as a distinct membranous film than in any other condition of cell development which I have a observed. The plasma membrane is in these stages never broken through. It always forms a perfectly continuous enveloping layer surrounding the entire segment which is being divided, a'° shown in figs. 6-9. In addition to the existence of these hyaline areas which predetermine the future cleavage planes, another striking condition is to be noticed. The hyaline regions Fuligo bound off inf every case a single nucleus and ees a group of nuclei. This nucleus may be in process of i but the daughter nuclei are never completely reconstructet ‘a : stage when these hyaline regions are present cutting a se : pair. The segregation is about the nuclei as units, 4? ie cleavage thus predetermined is to be a cleavage by whicr entire mass will be cut into uninucleated segments. beet hitherto the cleavage planes in these central masses have 190] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 229 oriented without especial reference to the distribution of the nuclei, thus cutting off larger and smaller segments with more or fewer nuclei, they are now to proceed midway between each pair of nuclei, so that equal and uninucleated masses will result. As already noted, the impression is very strongly given by these dense rounded masses separated by relatively watery regions that a contraction has taken place about each nucleus as a cen- ter. Such conditions might be produced by a contraction origi- nating in the structure of the cytoplasm itself, or in a pull exerted upon the cytoplasm by the nucleus. After the forma- tion of these hyaline areas, the cleavage is completed by the furrowing of the plasma membrane along the lines marked out. A later Stage than that shown in fig. 6, in which certain furrows have already cut deeply down into the hyaline regions, is shown in fig. 7. The furrows are apparently formed just as they were in the earlier stages, but they follow the hyaline areas, and thus the separation of the uninucleated masses is completed. Such Cleavage as this is progressive in the sense that both the hyaline “sions and the constriction furrows are developed gradually from the surface inward. When complete, however, it results in the simultaneous production of a number of uninucleated cells “qual to the number of nuclei in the original mass. And in this Sheet it differs markedly from the cleavage in the earlier wa 2 which larger multinucleated masses are progressively ae. oy Sapa with fewer and fewer nuclei. I have “ag a similar differentiation of a hyaline region pre- of ani. the plane of cleavage in the formation of the aporne own in a (4) 2. 25, jig. 21). In this figure resting nucle are throuch ° groups, between which a less dense zone is formed, 8) which later, as is Shown in jig. 22, a cleavage furrow he appearance was little regarded in my description oe ae of Clea ‘ i nee Stage Ohm a Pilobolus, but its appearance at a similar late . ‘Siderable <; ©avage of Fuligo indicates that it may have con- Ruclej © Significance in connection with the relations of the at least % Cleavage Phenomena. The hyaline areas, in Fuligo @ppear at about the stage in cleavage when the furrows 230 BOTANICAL GAZETTE [ocrorer seem to be more definitely oriented with reference to the dis- tribution of the nuclei. At the stage of embryonic growth, when they are found in Pilobolus also, cell division and nuclear division are proceeding in a somewhat definitely correlated fashion, It can hardly be questioned that whereas in earlier stages the cleavage was largely independent of the nuclei, it comes later to be directed solely with reference to their distribution, and it seems not unnatural to assume that in this latter stage the nuclei control the orientation ot the cleavage planes. If this is the case, it is quite possible that the formation of the hyaline zones is the visible expression of this activity of the nuclei. On the other hand, it is quite possible to assume that cleav- age throughout is controlled by the cytoplasm, at first with little reference to the distribution of the nuclei, but later with special reference to the formation of uninucleated cells. The formation of hyaline zones preceding the cleavage furrows might in this case also mark the transition from the earlier irregular to the later more definitive Stage of cleavage without implying any special activity of the nuclei, I have already noted that it is quite as easy to assume that the cytoplasm itself contracts about the nuclei as that it is drawn together by a tension exerted from the nuclei. Either view is consistent with the assumption that mate rial for the growth of the plasma membranes is formed in the nucleus and passes outward from it to the newly-forming cel boundaries. As is seen from jigs. 6-9, the nuclei are dividing while the cleavage just described is going on, so that the unin cleated Segments formed become almost immediately binucleated- ; Cell division then follows either before or after the compl j Teconstruction of the daughter nuclei (figs. 26, 17): aie a the end, uninucleated spores are produced. The formation ‘ hyaline region and constriction furrow for the division of — : cleated cell whose nuclei are already in the anaphase nt 4 shown in fig. 9. The beginning of the constriction for the # c division of a binucleated cell to form two uninucleated ee : the hyaline region having not yet appeared, is shown” - 226. S 1900] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 231 Transition types of cleavage between that without and that with a preliminary formation of a hyaline furrow are abundant. Fig. 8 shows a six-nucleated mass of protoplasm dividing by furrows, which in three cases are preceded by hyaline differen- tiation, and in the other two cases are cutting directly into the undifferentiated protoplasm. With the formation of uninucleated segments whose nuclei divide no more, the process of cleavage is complete. As noted above, the cleavage results in uninucleated segments at the periphery of the protoplasmic masses much earlier than in their interior. The nuclei in these early formed segments are always found dividing. The definite spore cells with a single resting nucleus are probably formed first in those regions of the aethalium where cleavage first began. The final delimitation of the spores seems to proceed progressively from these regions in all direc- tions through the aethalium. Fully formed spores may ulti- mately be found throughout the greater part of the aethalium, while in certain regions here and there cleavage may still be in Progress. There is, however, in the later stages of cleavage no “Saeerae difference between peripheral and central regions as eo during the early stages. Whether this is due to “ation in the cleavage at the periphery during the later Pei a a the nuclei there divide repeatedly to prolong eaEN have not been able to determine. ae aa we may characterize the whole process as one : as on : ravage by means of furnows which cut through ing angles ia Mass in very many directions and a very ave: that the as other. The process is progressive both in toward the . pueinate on the surface and proceed gradually are first Sag and in that larger multinucleated segments Shade e which are by further divisions reduced to the ., 1 Of uninucleated spores. It may perhaps be distin- Buished from bj artiti =e fe i Itiparti- ok i ition as a process of successive mu tipar Protoplasm sir, — furrows may invade any particular portion 0 the — aneously from a number of directions. At first of the cleavage planes shows no evident relation” 232 BOTANICAL GAZETTE [OCTOBER to the distribution of the nuclei. Later the furrows proceed in every case so as to cut off uninucleated masses approximately equal in size. This later period of cleavage is characterized in many cases by the aggregation and rounding up of the denser cytoplasm about the nuclei so as to leave hyaline regions midway between each pair of nuclei, thus predetermining in each case the plane of cleavage to be followed later by the cleavage fur- row. This type of cleavage results immediately in every case in the formation of uninucleated cells whose nuclei, however, may still be in a state of division. In the end, the entire pro- toplasm of the aethalium has been cut into uninucleated seg- ments which are at first naked bits of protoplasm. Later each cell becomes surrounded by a wall and constitutes a spore. Turning now to the phenomena of nuclear division, we may note first of all that the structure of the resting nucleus com forms, in spite of its small size, to that of the nuclei of other fungi and the higher plants. Nuclear membrane, chromatin (nuclein), and nucleole are present, and are differentiated by staining with safranin, gentian-violet and orange, just as sha rply as they are in the pollen mother cells of the lily. The nucleole frequently lies in a clear space (fig. 70), a& is so frequently seen in the nuclei of the root-tip of the onion: The nuclei, however, are too minute for the successful study of the prophases in spindle formation. In the equatorial plate stage (figs. 3, 5, 11) the spindle is sharply ‘differentiated. shows rather sharp-pointed poles which may be more peat stained at their tips. Broad-poled spindles, such as those figure by Strasburger for Trichia, are not found in Fuligo. The ea somes stain deeply in the equatorial plate stage. In polar ued it is possible to count the number with considerable speaginte? The great number of these figures to be found in “— Fuligo at this stage make the material especially favors such study. The chromosomes are relatively short and and form a very regular equatorial plate, all of them bie om tically in a single plane, so that in polar views they are pra a cally all in focus at once. From a study of a large nanan Er 190] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 233 such figures, I am quite certain that the number of chromosomes is twelve. All stages of the separation of the daughter chromosomes and their migration to the poles of the spindle can be found in the greatest abundance (igs. 6, 8, 72-15). The spindle becomes slightly elongated during this process. Connecting fibers are © present and form a figure closely resembling that in the corres- ponding stages in the lily or larch. As the chromosomes first separate, relatively large gaps are seen between the connecting fibers, which appear bunched together in a few large strands (fig. 13). The whole connecting spindle is markedly barrel-shaped at this stage. As the chromosomes approach the poles, the Te fibers become more evenly distributed, and are straightened so as to form a cylindrical series extending between the groups of daughter chromosomes (fg. 74). A marked Pi e between the nuclear divisions in Fuligo and those in ic rh I have studied is seen in the arrangement of the i e se chromosomes as they are drawn back. to the poles. Re asci these chromosomes are widely scattered on the while a this Stage, some having nearly reached, the poles, lit os ae are much nearer the equatorial region. This con- chicas, €s this stage the most favorable for counting the tena of the nuclei in the ascus. In Fuligo, on the toward the all the daughter chromosomes retreat simultaneously inte de, * as is seen in figs. 6, 13, 14. They also become Somes are “a ond together, so that the individual chrome: a8 in the ¢ Ae easily distinguished in polar views at this stage i quatorial plate stage. ; Te is etter chromosomes reach the poles, the whole (fig. 7 oe elongated, the connecting fibers being drewn appears, ag a a long slender strand, which gradually dis- the groy Poles of the spindle can be distinguished beyond 9 15), Out id of daughter chromosomes till a very late stage (/igs- The n ” Tather ucleole of the parent nucleus in Fuligo disappears ata Car . . Y Stage as compared with other fungus nuclei. It 1s 234 BOTANICAL GAZETTE [ocronER never to be found lying midway between the daughter nuclei near the old spindle, as is regularly the case in many asci. The daughter nuclei are apparently reconstructed in the ordinary fashion. The figures, however, are too small to show very characteristic details at this stage. : The similarity of the whole process of nuclear division in its main outlines here to what is found in higher organisms is cer- tainly very striking, and shows clearly enough that simplicity of structure and life history on the part of the whole organism is by no means to be taken as indicating a corresponding reduction in the complexity of the nuclear structures and activities. The capacity of the slime molds to become encysted at any stage in their life history when conditions become unfavorable is very well known. A condition which I have sometimes found, and which is represented in fig. zg, indicates that this may occur midway in the process of cleavage. The aethalium i question was made up of rounded, two- to several-nucleat masses, each provided with rather a thick wall. Whether ae with a return of favorable conditions such masses would continié their cleavage, and form normal uninucleated spores, of whether they would themselves function as spores, I have not been able to determine. A normal uninucleated spore is shown in fig: 18. The aethalium and the sporanges of the Myxomycetes differ from the sporanges of Synchitrium, Pilobolus, and Sporodin® whose method of spore formation I have already described (4), in that the multinucleated condition in the former originates ® least in the formation of the plasmodium. The plasmodium a product of cell fusions without nuclear fusions, $0 far as knoe at present. Physiologically considered, in nutrition, growth, and response to external stimuli, me equivalent of such multinucleated masses of protoplas™ ee a formed simply by growth and nuclear division without pie ‘ sion. The plasmodium itself increases its original ee formed by fusion, by this same type of growth. Fundam ycley considered, it is the physiological equivalent of the ae ae ted mass formed in Synchitrium by the division of the ° 190) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 235 single nucleus of the vegetative body. The internal relations of nuclei and cytoplasm cannot be conceived as different, merely because in one case the nuclei and separate cytoplasmic masses were brought together by fusion, while in the other they were formed from a uninucleated cell by growth and nuclear division. Morphologically, however, the two structures must be re- garded as entirely distinct, and the possession of the plas- modium, and the capillitium formed as a deposit in vacuoles by the slime molds, is probably sufficient reason for regarding them as constituting a separate developmental series running back to an origin independent of any of the existing groups of alge or fungi. Sachs (18) has quite recently expressed the © opinion that they are to be classed with the fungi, but he brings ho morphological evidence to support his view. There can be fo question that the Acrasiez represent simpler forms out of which the Myxomycetes have developed, and we thus have a developmental series leading from simpler to more complex forms. The plasmodium and capillitium, appearing only in the oN specialized members of the group, are plainly secondarily acquired structures developed as additions to the structural features of the Acrasiez, and are not to be directly homologized 208 Physiologically equivalent structures in other groups. muling Yological equivalence of the plasmodium and the : fe a. masses of protoplasm found in other fungi, such : where ( We. Pilobolus, etc., can hardly be questioned. , Else- i ‘multinuclesg ee discussed the question as to whether these _ ae ‘ Masses should be classed as single cells, or as ee of @ slime a isang tissues or organisms. The plasmodium On this Saas 53 well calculated to furnish further evidense amoeboid Ae n its method of origin by the fusion of distinct Jaen oo. it would seem to testify to its multiple in in an sea above, the whole physiology of the plas- nost oa reactions to stimuli, and growth, shows i fice a that it is a unit in exactly the same sense as ; OF one of the swarm-spores which combined to ' form it d ng fusing, the Swarm-spores gave up their individuality 236 BOTANICAL GAZETTE [ocropER to become parts of a larger mass. That this fusion did not involve a fusion of their nuclei cannot be considered as altering the result so far as the question of individuality is concerned. Where sexual cells fuse and their nuclei unite there is no ques- tion that the resulting fertilized egg is a single cell. If, as Hacker (2) has shown is the case in Cyclops, the pronuclei remain distinct through the early cleavage stages of the egg, this cannot be taken as evidence that the two nucleated bodies thus produced are not single cells rather than the equivalents of tissues. These binucleated cells, functionally and morphologi- cally considered, are the equivalents of the later cells of the ‘ Cyclops which appear with a single nucleus. The conclusion must be, as I have already pointed out, that the individuality of the cell is independent of the number of nuclei which it contains. Hertwig argues for the potential equivalence of multinucleated cells and tissues. The word potential here of course may mean much or little. In support of his view he urges the case of the insect egg, whose nucleus divides to form hundreds of daughter nuclei before cleavage begins. Later the multinucleated yolk mass is by cell division separated into a blastoderm of as mam) cells as there were nuclei present. It is quite plain, says Hert wig, that the apparently simple egg could not with a single stroke, as it were, have become a multicellular organism. he question here, of course, is how great a change is involved in transition from the one-celled to the many-celled condition, on this point it is interesting to note that up to the stage wns cell division takes place in the insect egg there has been 9° we ble differentiation of embryonic structures in the egg- The a t division simply transforms the one cell into a mass of equivalem cells, and this need hardly be considered as a change too of to be due entirely to the cleavage process. The relation multinucleated and uninucleated cells is well shown i” the ee fact that: the visible differentiation of the insect embry ost’ perhaps from the determination of its axes, which da plished even earlier, begins after the division of the eg8 oa numerous cells, and not while it remains a single cell, althous™ the and 190) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 237 it has meanwhile become multinucleated. It would seem that differentiation was dependent to a certain degree, at least,on the interaction of individualized protoplasmic units, each capable of receiving and reacting to independent stimuli as the parts of a multinucleated cell cannot. Hertwig, in his doctrine of biogen- esis, himself insists on the importance of the interaction of separate cells for the production of physiological differentiation and division of labor. o be sure, we have abundant evidence that the multinu- cleated cell can achieve a certain degree of differentiation, as is shown in the numerous Siphoneze which mimic in their root- like, leaf-like, and stem-like structures, the analogous parts of the higher plants. It is perfectly apparent, however, that this differentiation is on a far simpler scale than is seen in the com- plex mechanical and other tissue systems and organs of the higher plants, Indeed, the relative unimportance of the Sipho- hee as a part of the earth’s vegetation is to be regarded as very strong evidence that the type of structure which they show in their multinucleated cells is by no means well adapted to develop “omplexity and differentiation of structure such as is necessary es meet the manifold variations in environmental conditions to Which all plants are subjected. Thesé Siphonez are after all a more differentiated than the infusorians, which are typi- : y unicellular, _ Pfeffer (16) puts the case very strongly when in Sp ee that we can conceive of no such independent units multinucleated cell as the energids of Sachs are defined to egy energid is a nucleus with a portion of cytoplasm the Somes control, there can be no such structures in pionee, since the protoplasm of their cells is constantly i one point to another, with the exception of the hin. ee : a which remains fixed. No nucleus could thus Di . nite relations with any particular portion of the int heise, Neg and it is hard to conceive that in this stream- - CONstantly ; ‘3 Portion of the semifluid cytoplasm should remain Says, w YM connection with any particular nucleus. As Pfeffer _ ome must conclude that any specific mass of cytoplasm in a 238 BOTANICAL GAZETTE [ocroser multinucleated cell will be simultaneously influenced by various nuclei which are in contact with it. There can be no invisibly bounded units in which the same living substance remains united. Pfeffer also justly objects to Sachs’s characterization of the Siphonee as xoncellular plants, and regards them as both mor phological nnd physiological units. If we compare the method of spore formation in Fuligo with that which I have described elsewhere (4) for Synchitrium, Pilobolus, and Sporodinia, it will be seen that the processes in all these forms are identical in their main features, while differ- ing in a number of important details. In the four cases the cleavage is progressive from the surface inward, larger segments being first formed, which are later cut up into uninucleated cells, except in Synchitrium taraxaci and Sporodinia, in which the multinucleated segments function directly as spores. In the earlier stages of cleavage in Pilobolus and Fuligo the furrows pierce through perfectly undifferentiated and quite homo- geneous protoplasm, while in the later stages the differentia- tion of hyaline areas, wedge-shaped in transverse section and cutting through the masses to be divided, predetermine the planes of the cleavage furrows. Such hyaline areas were not observed in Synchitrium or Sporodinia. In Synchitrium and Sporodinia nuclear divisions precede cleavage. In Pilobolus nuclear divisions occur during the later stages of cleavag® and in Fuligo nuclear divisions and cleavage proceed simultaneously throughout, Fuligo is the only one of the five forms in whi cleated segments formed by the completion of process, and which I have called protospores, become the ra tional spores directly without further growth or nuclear divisio™ In this respect perhaps the cleavage of Fuligo represents 4 more simple primitive type than that of either of the others. : In all forms the orientation of the furrows with reference the surface of the dividing mass and with reference to © other is extremely varied, and it can be laid down as @ pes tule for the forms studied that no one furrow can be ch the uninu- 900) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS — 239 continuously through the entire sporange or aethalium which is to be divided except, perhaps, in the case of the very thin layer of spore plasm in Sporodinia. On the contrary, by the curving and branching of the furrows, segments irregular in their size, shape, and number of nuclei are cut off successively from the periphery toward the center. These segments in turn, and also progres- sively from periphery to center, are cut up by new furrows into smaller segments, until finally in Synchitrium decipiens, Pilobolus, and Fuligo the uninucleated condition is reached. No general system of cleavage planes, either parallel or radial to the surface of the dividing mass, can be discovered. The path of the cleav- age planes as division progresses becomes an inextricable con- fusion of zigzag lines, branching and intersecting at almost every angle. The occurrence of such similar types of cleavage of the multinucleated mass as are found in the aethalium of Fuligo and the sporanges of the Phycomycetes must be regarded as another example of parallel development in structures not phylogeneti- cally connected. The explanation of’ the similarity in these forms of cleavage is to be sought in the fundamental physiologi- cal properties of protoplasm, and not in hereditary transmission to me different branches of a series of genetically related forms. With the above account of fusion in Fuligo representing the Myxomycetes, types of all the main groups of fungi producing “Sexual spores in the interior of mother cells have been described “xcept the Oomycetes, and while it will be necessary to investi- ee atives of all the genera, at least, in these groups, ae BN es fairly justified that some form of progres- ey ae than simultaneous cleavage by cell plates will be alia a case. Klebs’s (15) investigations of Hydrodictyon simultane e met the formation of zoospores is at least not by us division into uninucleated segments, and the whole Process in this alga should b i igated ially with telerence oR ga shou e further investigate ; especia y Sethes ic c. occurrence of the nuclear fusions which Klebs Roted here a in the developing zonspores. It may he of “gpa that Bachmann (1), in describing a new species a, has observed incidentally the marking off of the 240 BOTANICAL GAZETTE [OCTOBER surface of the sporange into irregular polygonal areas at the time when spore formation is beginning. The lines marking off these areas are doubtless the beginnings of the cleavage furrows as I have described them for Pilobolus and Sporodinia. Bach- mann made no sections, but concludes that the spore formation must be a progressive process. A sufficient number of forms has been investigated to show that progressive cleavage is a widely spread phenomenon among the lower plants, occurring in very many cases when multinu- cleated masses are to be cut up into smaller cells. That simul- taneous cleavage may also occur is quite possible, but the evidence for it is not strong except, perhaps, in the case of the sporange of the Saprolegniacee. I have already shown elsewhere (4) that the progressive cleavage of the sporange in spore formation is in principle the same process as that which has been described as division by constriction in Cladophora and the conidiophores of the mil- dews, and it will be of interest: to attempt a comparison of this progressive cleavage with cell division as found in the growing points of the higher plants, especially from the standpoint of the more general theories of cell division. Schleiden’s (21) doctrine that the form of a plant is deter mined by its cellular composition, including as the two important factors the arrangement of the new-formed cells in growing regions and the subsequent varying growth and enlargement of these cells in their three dimensions, was first opposed y Hofmeister. Hofmeister (7 and 8, p. 129) advanced the vee that cell-formation is.subordinate to the growth of an enlarging organ taken asa whole. He held that the growth of the single cells of a vegetative point is controlled and conditioned by some formative principle which determines the growth of the comme organ, this latter being directed toward simple enlargement OF the development of some predetermined form. According this view, growth of the vegetative point cannot be interpret as determined by the sum of innate growth tendencies of individual cells, 390] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 241 Hofmeister believed that cell division takes place according toasimple mechanical law. The new cell wall always cuts the axis of most intense. growth at right angles (8, p. 127). It is plain that such a principle as this can have no application in interpreting the division of the masses of protoplasm in sporanges whose growth is complete, the accompanying tensions, as may be fairly assumed, having also reached a condition of equi- librium, Sachs (20 and 18, p. 22) follows Hofmeister in regarding the growth and division of the single cells as subordinate to the growth of the vegetative point as a whole. From a study of the arrangement of the cells in the growing point of the higher ‘ryptogams and flowering plants, he has developed the law of the rectangular intersection of cleavage planes in the successive cell divisions. He regards this as the most universal law of Sttucture in the plant world, and holds that it is independent of all phylogenetic relations, and not a result of natural selection. As is well known, he holds that the outer form of the growing organ is the primary determining factor for the orientation of the Cleavage planes. The periclines conform directly to the surface of the embryonic organ. The anticlines cut the peri- one at right angles, and if division is to occur in three dimen- ns the transversals also appear in a third plane at right angles ot. Sachs believes that the relations thus expressed es undamental a nature as to be comparable to those Hs ing the relations of the axes of a crystal. The cellular Reccca, ystal to the arrangement of its faces and their S angles. The form of a growing point is determined and selection when given the direction of the 2 th Planes, is at once known. Such fundamental relations ese, m Sach, La hanomorphoses. The principle thus developed by Rent of sh generally accepted as explaining the arrange- € cell walls in the growing points of the higher plants, 242 BOTANICAL GAZETTE [ocToRER and, as Sachs himself notes, the investigation of the apical cel- and its divisions has ceased to be regarded as affording a key to the explanation of the development of shoots. Assuming the correctness of Sachs’s law for the higher plants, if we attempt now to apply it to the case of the cell-division in sporanges we are confronted with many difficulties. First of all, these sporanges do not divide by successive bi-partitions as do the cells in the growing points referred to. Nor do they divide by simultaneous delimitation of the energids which, according to Sachs’s view, compose them. Their cleay- age is progressive, and the cleavage planes, as shown in sections, form no great series comparable to the anticlines and periclines which Sachs finds in a section of a root tip. In the cleavage of the sporange the principle of rectangular intersection is violated constantly, the angles of intersection of the cleavage planes showing no constancy whatever. It may be objected that the Sporange is not a growing organ, and hence its method of division should not be expected to conform to that of grOnms points. Growth is complete in the sporange before division begins, though it may recommence in the later stages of cleav- ‘age in Pilobolus. The process in the sporange consists 1 cutting up into cells a mass of protoplasm whose form has bet? already determined and its growth completed. Still, although Sachs states the principle of rectangular intersection for oer ing points, and conceives it as determining the arrangement © the cells in the growing point as it pushes forward in the elonga tion of the shoot, he always conceives the divisions as sre in these growing points after the essential embryonic growth 0 . the cell concerned is at an end. Growth of cells subsequent © division may, and generally does, in his opinion, distort the relations of the cleavage planes. : pe Sachs makes no attempt to include the multinucleated sp” th of the rangia in his discussion. Still he specifies the grow Siphonez (19, p- 100) as exceptional when compared irregular growth of many thallophytes, and considers night opment at their growing points as typical of that the Me with the 0) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 243 plants. He also specifically states that typical mechanomor- phoses are the cell nets in growing points of young organs and structures whose cells show no individual growth after cell division is complete. All the sporanges mentioned above would be included in this latter type of structures. The sporange of Synchitrium isaspherical mass of protoplasm which divides into cells which show no further growth, at least till after cleavage is complete. At this stage, then, the cleavage planes shouldiillustrate typical mechanomorphosis. It would seem that in such a spherical mass of undifferentiated protoplasm the opportunity for the law of rectangular intersection to come to full expression would be especially good. We might expect the periclinal and anticlinal Cleavage planes to be extremely conspicuous in suchacase. On the contrary, as noted above, and as is well shown in figs. 2, #14, 15, pl. 24, of my former paper (4), no regular periclines and anticlines are to be observed. The surface furrows cut into the mass at very varying angles, frequently also becoming curved and branching so as to intersect near the surface, and thus cut off Superficial cell-masses of the most irregular shapes and mes. There is apparently the greatest irregularity in the Srentation of the cleavage planes both with reference to each other and the fiour to the surface of the dividing mass, as a glance at bes i, referred to above will show, and as I have described Cleava Ae ie scape of Fuligo. As this method of progressive the ie _ is of wide occurrence among the thallophytes, ie "a e of Tectangular intersection loses that universal fo division upon which Sachs so strongly insisted. . we vase of rectangular intersection so as to further Sion to . meister’s doctrine of the subordination of cell divi- Tesults ie of organisms as wholes, and in this sense his theoretical] anor the “asked part, been taken up and utilized for its division purposes in discussions on the nature of the cell and the oa : a 26). With Hofmeister, Sachs consider? elops “eis point of such a structure as the pine shoot Stow} ply as a mass of protoplasm analogous to that at "98 end of a Vaucheria filament. The shape of the end 244 BOTANICAL GAZETTE Loctoser of the shoot has been determined by selection during phyloge- netic development; its growth is the enlargement of a proto- plasmic mass along lines predetermined. The putting in of the cell walls, being a purely mechanical process following the simple rule of rectangular intersection, can in no way be regarded as determining the shape of the shoot. The tip grows just as does the Vaucheria tip, the formation of individual cells being a purely secondary matter. The independence of the single cells, their growth and division, is entirely subordinated to the growth of the entire organ. It is an open question whether the rectangular intersection of cell walls in vegetative points, assuming that the facts are as Sachs describes them, is sufficient basis for so important a conclusion as that above stated. Jennings has characterized the law of rectangular intersect tion as ‘‘hardly to be considered as more than a statement of a condition commonly found.’ There is nothing inherently impossible in the assumption that the habit of forming — sive cell plates so that they intersect at right angles has itself been acquired by the cells as individual units. The form of oe shoot may then depend on this power of the cells, assuming always a further regulation of their activities by reciprocal stimuli between the cells themselves as well as by stimuli from their environment. There can be no question that at least the details of cleavage in the sporanges I have studied have been modified in the course of phylogenetic development with especial reference “2 needs of the organisms concerned. I have elsewhere pout i out the correlation between the abbreviated cleavage poe Sporodinia and its more rapid spore development. The gi pation of vacuoles in the formation of the cleavage aa Pilobolus is another example of such modifications. Ps nt cleavage processes are after all so similar in the ae Sporanges studied is doubtless due to the fundamental phys and chemical properties of the protoplasm, but the process 4 ne : Nn. none the less to be conceived of as modified by select? 90] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 245 must also be borne in mind constantly that in such cleavage phenomena as these the cleavage planes are influenced in no way by the need of forming any specific tissue or plant form In the growing point of a metaphyte it may perhaps be difficult to decide whether the planes of division are determined by the cells themselves in accordance with internal conditions, or whether, as Sachs claims, it is the shape and differentiation of the shoot taken as a whole which determines the planes of division. In the division of these spore-forming masses no such question can arise, since no differentiated tissue is to be formed. The problem is simply to divide the large mass into smaller masses more con- venient for distribution. In doing this, as I have shown, entire megularity prevails as to the orientation of the cleavage planes with reference to each other, and with reference to the axes of the mass to be divided. It may be concluded that the proto- Plasm is per se perfectly isotropic so far as cleavage is concerned, and that it is a matter of indifference whether the cleavage planes intersect at right angles. If rectangular intersection is the rule in the higher plants, it might well be argued that this is *Sccondarily acquired condition, assuming with Pfeffer that these multinucleated structures are single cells. ‘ _ can hardly be raised in this connection whether iiss: m : the cell forms the new cells or spores. Still it is. ik § and significant to note, as I have pointed out already / similar 1 toa of cleavage in these sporanges is essentially a. the typical cell division by constriction, eR mperibed as producing the one-nucleated cons at found i, Sa pee in the mildews, and which is also Muu CC t0Rtessive cleavage by suriace TES thon, ae modification of cell division by constric- Foheites eae is simply cell division, though not by in the es : ions of the mother cell, as it commonly occurs ret eit idea of the higher plants. ‘ a illustratin 88 the Multinucleatae cannot then be regarde logically Ais € growth of an organism comparable morpho- a metaphyte but without cell formation. The ( 246 BOTANICAL GAZETTE [OcToRER coenocyte is a cell, and its growth and differentiation is compara- ble to the growth and differentiation of a single cell in the growing point of one of the higher plants. That such a cell may become multinucleated is illustrated by the multinucleated cells in the red alge and the striped muscle fibers of animals Just as in these cases, the coenocyte everywhere is a cell which has beome multinucleated strictly for functional purposes requir ing the distribution of the nuclear material through the enlarged cell body. The more modern theories as to the division of the animal cell all assume a definite correlation between nuclear and cell divi- sion. The mechanism of the one is definitely connected with that of the other. The division of the cell at right angles to the long axis of the karyokinetic spindle is the rule among the higher animals as in the higher plants, and the later theories have assumed this relation as fundamental. Thus, Heidenhain’s (5) theory assumes that division of the cell is a result of tensions in the unequally stretched elements of a ‘system of orgaiie rays” extending from the centrosome to the plasma membrane. Kostanecki (11) attributes cell division to the development of a cell plate which is formed as a result of the migration of the ends of pairs of polar rays, produced by the splitting of parent rays, from the points of their original attachment to the plasma membrane of the mother cell into the equatorial region between the daughter nuclei. Jennings (10) has summarized the theories of cell division and of the orientation of the successive cleavag® planes in animal cells, and I need not enumerate t They are for the most part directed especially to the explana tion of cases where nuclear:and cell division have become d nitely correlated, and in many cases they assume t tion of the plane of cell division by the axis of ™* spindle. Cases of cell division between resting nucle! are found in the cleavage of the insect egg, have re ; attention. Still, in all the sporanges studied it is plain ue all else that nuclear division neither determines 10 Way connected with cell division, and it is thus show f the nucleat n that the : hem here. he determina 190] CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 247 cytoplasm can divide without any reference to the division of the nuclei. The cytoplasm both initiates and completes the process of dividing any particular cell mass, while the nuclei are through- out in the resting condition in Synchitrium; and, on the other hand, in Fuligo it can carry on the cleavage in essentially the same fashion simultaneously with nuclear division, without apparently being influenced in the least degree by the occurrence of the latter. In Fuligo the axes of the nuclear spindles and the planes of the cleavage furrows may be inclined at all imagin- able angles to each other (figs. 5-7, 8,9). It is always to be hoted in these cases _that the cleavage furrow in question is not the one destined to separate the daughter nuclei which are in process of formation at the time when the furrow is forming. The cleavage always lags behind nuclear division, never sep- “ating any two daughter nuclei until they have reached at least the resting condition, or are themselves engaged in the next fol- lowing nuclear division. This latter case, as found in Fuligo, is rernally interesting as showing most clearly that the apparatus of nuclear and cell division is entirely distinct. The mechanism of nuclear division is in full operation while cleavage furrows are ages in adjacent regions of the cytoplasm in entire inde- here oe It is thus shown that no such conditions exist division ig a Osperm formation in the lilies, where, after nuclear com omplete, a set of new connecting spindles are formed in a resting daughter cells. The mechanism of cell divi- a rem is the same as in ordinary cell division, e cs. sie upon nuclear division. It is presumptively ics. i Haclear division which is in OperaHee in this and ieoueatt division, though operating independently of divisions ee: id the completion of a long series of nuclear the nuclei are i osperm formation we find no cases in wee tes are form emselves dividing at the same time that ce also dividing os between them and adjacent nuclei which are tvidence in the “gs ag contrary, | have observed considerable iN cases Where eke of the pollen mother cells of the larch, € first nuclear division is not to be followed by sion whi 248 BOTANICAL GAZETTE {octosER cell division, that the fibers of the connecting spindle are utilized, at least partially, in the formation of the spindles for the second nuclear division. It is by no means to be argued from the fact that cell divi- sion and nuclear division are independent in these simple forms of plant life that the processes are not most intimately connected in the higher plants and animals, nor that the position of the nuclear spindle may not determine absolutely the plane of cleav- age in these latter cases. In the higher plants there can be no doubt that the spindle, persisting after nuclear division, forms 4 cell plate and determines the plane of cleavage. On the other hand, the process of cell division in Fuligo shows very clearly that such a correlation is by no means fundamental or universal. Whether or not the nucleus in any fashion influences the orienta- tion of the cleavage planes in this latter case, it does not do tt by means of the karyokinetic spindle. Wille (25) ina preliminary communication has reported ine discovery of a type of division in multinucleated cells in which the nuclei participate in the formation of new cross walls. Just what the nature of the nuclear activity is in this case is not clear from the brief report referred to. It certainly represents 2 0e¥ and most interesting condition in multinucleated cells. It is quite possible that the irregular cleavage of the eB Sporange indicates a primitive condition when nuclear and ¢ division are entirely independent processes, and that the ae lation of the two has been gradually achieved in the wee of the higher plants and animals. It is plain, therefore, that theories of Heidenhain and Kostanecki (5 and 11) can ai 2 application in the explanation of cleavage in these sporange®s ge it is further plain that no theory which interprets cell divisio® a function of the karyokinetic figure can claim to be of : nie mental value for the explanation of the process. Cell divis may be much more definitely related to purposeful . i i i ; : tissue formation when, as in the higher plants, it is accom! pa by the mechanism of the spindle and cell plate. It 1 si e planes sible that the regular orientation of successive cleavag results 100) CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 249 so as to form tissues,z. ¢., aggregates of cells with specific shapes and dimensions, only became possible after cell division came to bea function of the same mechanism which effects the division of the nucleus, but, as Strasburger (22) long ago pointed out, the process itself is by no means necessarily dependent upon the existence of this particular mechanism or of any correlation whatever with nuclear division. Hofmeister (8) calls attention to the fact that the division of protoplasmic masses for the formation of reproductive cells is quite universally accompanied by loss of water and reduction of volume with increased density of the protoplasm. This phe- tomenon of contraction and loss of water is especially conspicu- ous in the process of spore formation in sporanges, as I have already noted. It seems a fairly natural assumption that the tensions set up in a mass of protoplasm which is contracting as @result of loss of water may be utilized in some fashion to pro- duce the extremely irregular cleavage furrows which we observe in the early stages of spore formation. That these furrows, however, are not purely mechanical in their origin and analogous ‘0 the fissures that appear on the surface of a drying colloidal Mass js shown, as I have noted elsewhere, by the fact that, _ oe they never cut off Sr inne containing no lacks 5 ultimately they produce approximately equal uni- a... Some form of organization must be assumed the cleava 2 in the protoplasm which determines the progress of ~~ ge So as to lead to a constant result. Still, even with fe “sumption, the possibility remains that the source of the Seg eae controlled may be in the tensions produced Nn due to loss of water. Usiversiry op Wisconsin, INDEX OF LITERATURE. Mortierella Van Tieghemi. Beitr. zur Physiol. der : /’. Dr Bary. A i, wiss, Bot. 3 > 279. | der Pil, oy Vergleichende Morphologie, Physiologie und Biologie €, Bacteria und Mycetozoa. London. : * Bacuwany, H- Pilze, Jahrb, 250 2. ~*~ o M3 | M4 ° Leal Leal led bd + > WwW 5. 16, 17. 18. 19. 20, 21 aa. 23. 24. 25 . 26. 27. . Lister, A.: On the division of nuclei in the Mycetozoa. BOTANICAL GAZETTE [ocToBER Hicker, V.: Uber die Selbstandigkeit der vaterlichen u. miitterlichen Kernbestandtheile wahrend der Embryonalentwickelung von Cyclops. Arch. f. mik. Anat. 46 : 579-618. . Harper, R. A.: Sexual reproduction in Pyronema and the morphology of the ascocarp. Annals of Bot. 14:—. 1900. [ined. Harper, R. A.: Cell division in sporangia and asci. Annals of Bot, 13: 46 HEIDENHAIN, M.: Neue Unters. iib. die Centralkérper u. ihre Beziehungen zum Kern u. Zellenprotoplasma. Archiv f. mik. Anat. 43 : 423. 1894. . HERTWIG, O.: Die Zelle und die Gewebe. II. Jena. HorMEISTER, W.: Abhandl. Siichs. Gesell. d. Wissenschaften 642. 1857. HorMEISTER, W-: Die Lehre von der Pflanzenzelle. Leipzig, 1867. JAHN, E.: Der Stand unserer Kenntnisse iiber die Schleimpilze. Rundschau, Oct. 1899. Nat, . JENNINGS, H.: The early development of Asplanchna Herrickii, Bull Mus. Comp. Zool. Harvard 30: No. I . Kostaneckt: Uber die Bedeutung der Polstrahlung wihrend der Mitose und ihr Verhiltniss zur Theilung der Zellleibes. Archiv f, mik. Anat. 49:651. 1897 Linn. Soc. Jour. 29: 529. 1893. . LisTER, A.: Mycetozoa. _ London, 1894. . Massee, G.: A monograph of the Myxogastres. London, 1892. KieEss, G.: Uber die Bildung der Fortpflanzungszellen bei Aydrodico™ utriculatum Roth. Bot. Zeit. 49 : 790-876. 1891. PFEFFER, W.: Pflanzenphysiologie. 2 Aufl. Leipzig, 1897- Rosen, F.: Beitrige zur Kenntniss der Pflanzenzellen I. die Kerne und die Membranbildung bei Myxomyceten Beitr. zur Biol. der Pflanzen 6: 245. d Phylo SACHS, J.: Physiologische Notizen VIII. Mechanomorphosen um : genie. Flora 78:215. 1894. SACHS, J.: Arbeiten des Bot. Instituts zu Wiirzburg 2:4 SACHS, J.: Vorlesungen iib, Pf. Phys. Vorles. 26. 1882. Studien iiber und Pilzea. 6, 1882. . SCHLEIDEN : Grundz. 2: 12. [2 Aufl.] STRASHURGER, E.: Zellbildung und Zelltheilung. 3 Auf. Jen’. STRASBURGER, E.: Zur. Entwickelungsgeschichte der Sporang! Trichia fallax. Bot. Zeit. 42 >305. 1504. t Woods WHITMAN: The inadequacy of the cell theory of development Hole Biol. Lect 1893 . 7 +238: WILLE, N.: Die Zellkerne bei Acrosiphonia, Bot. Centralbls ® | WILSon, E.: The cell in development and inheritance. N. Y. ZopF, W.: Die Pilzthiere oder Schleimpilze. Breslau, 1885- BOTANICAL GAZETTE, XXX HARPER on FULIGO i900} “CELL AND NUCLEAR DIVISION IN FULIGO VARIANS 251 EXPLANATION OF PLATE XIV. All figures were drawn with the aid of the Abbé camera lucida, and all but 7 and 7 with the Zeiss apochr, obj. 2™™. igs. 2-7, and g with oc. 6; fg. 8 with oc. 12; and figs. zo-78 with oc. 18. Fig. 1. Portion of aethalium showing first cleavage furrows and lacunae ___ tfthe protoplasmic reticulum. Xx about 150 ___FiG. 2. Superficial portion of protoplasm showing two cleavage furrows and scattered nuclei. . . Fie. 3. Portion of dividing protoplasm from interior of aethalium and ' : lying between three lacunae marked 7. about 2 50. fa Fie, 4. Several entire segments and portions of others from surface of : lacuna in the interior of an aethalium ; profile and polar views of nuclei in the equatorial plate stage. Fig. 5. Segments from interior of aethalium with cleavage furrows cut- ting into it from various directions ; nuclei in equatorial plate stage. * Fig, 6, Segment cut through by hyaline areas which predetermine the course of the cleavage furrows. _ Me. 7. Later sta ‘Fig. 8. Segment Med protoplasm and i Fie. 9, Binucleat ge in cleavage of a segment like that shown in fig. 6. showing cleavage furrows cutting through undifferenti- others cutting into hyaline areas. Uninucleated segment; nucleus in equatorial plate stage; part of nuclear membrane of parent nucleus still present. hey Present, "2 Metaphase ; part of nuclear membrane of parent nucleus still - Later stage ; connecting fibers straightened. T; Connecting fibers a narrow strand. > Cell division beginnin fated segment; cell division not yet begun. : cleated spore with thin wall; granules of reserve material i h Binucl "6.18. Uninu “Ytoplasm, Sects Encysted segment. DOUBLE FERTILIZATION IN COMPOSITAE. : CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. XXI. W.-). G. LAND. (WITH PLATES XV AND XVI) In August 1898, Nawaschin communicated to the Russian Scientific Congress at Kieff the results of his work on fertiliza- tion in Lilium Martagon and Fritillaria tenella. Guignard, upon learning of the discoveries of Nawaschin, contributed to the Academie des Sciences a short account of his unpublished researches upon fertilization in several species of Lilium. Miss Sargant, from a reexamination of her preparations of Lilium Martagon, fully confirmed the observations of both Nawaschia and Guignard. A recent study of Tudipa sylvestris and i Celst ana by Guignard gives results in strict accord with the earlier observations. These observers find that both male cells upon emerging from the pollen tube are vermiform and twisted on their pa * ; : e suggesting the idea of non-ciliated spermatozoids. One ™ : r atm cell, coming in contact with the egg nucleus, retains fo a its vermiform shape, gradually enlarges until it becomes ; spherical, and finally fuses with the egg nucleus. The — male cell fuses with the upper polar nucleus, and the muclets resulting from this fusion unites with the lower polar a Sometimes the polar nuclei fuse and then unite with the fuse cell, and sometimes the polar nuclei and the male cell aL simultaneously. Preparations of Lilium Philadelphi oe Ze tuigrinum in this laboratory show the last named condition. are _Sargant figures one case in which the ends of the male see : applied to the polar nuclei, uniting them as kes ga : . . + 48 . : 5 Experiments in hybridization by De Vries and Corren roctoxst 252 Miss 1900] DOUBLE FERTILIZATION IN COMPOSITAE 253 indicate that double fertilization may be more frequent than is commonly supposed. So far as recent work indicates, the spermatophytes produce two male cells, The persistent appearance of a second male cell, seemingly as well organized as the one which functions, has found no better explanation than a phylogenetic one, although it would be hard to explain why a cell which has long been abandoned continues to be well organized. We may well inquire whether a simultaneous fertilization of the egg and of the endo sperm nucleus may not be universal in angiosperms. The study of the mature embryo-sac of Erigeron and of Silphium was undertaken for the purpose of determining the fate of the second male cell. The first named genus was chosen because a large number of ovules in different stages of develop- = could be cut at once; the second, because of the differen- tation shown by the disk and ray flowers. MATERIAL AND METHODS. Material was collected in the vicinity of Chicago, from June lg July 20, 1900, Collections were made at all hours of the ¥,and all material was killed in the field. The outer involucral = Erigeron were removed and the heads closely trimmed, y ough of the receptacle being left to hold the ovules SOE A I per cent. aqueous solution of chromo-acetic a killing and fixing agent. Carnoy’s fluid was oe pe unsatisfactory. The material was passed through nto paraffin and cut in serial sections 3.3 # and 6.64 thick. that for aay of treating Silphium was slightly different from ing the oo The ovules, except those intended for trac- issueg z = the pollen tube, were freed from the surrounding ‘emperature ated plunged into chromo-acetic acid at a acid me about 100°, and were allowed to remain in ge hey were ut two hours. After washing and dehydrating, soy Ep “brough xylol into paraffin at a temperature : : Sec i : ° . Tictotome. Hons 2~5 w thick were cut on a Reichert rocking 254 BOTANICAL GAZETTE [ocroser Flemming’s safranin-gentian-violet-orange G and Haiden- hain’s iron-alum-haematoxylin both gave excellent results, but the most satisfactory differentiation in fertilization stages was obtained with cyanin and erythrosin. This last combination, after the sections had been treated with acetic acid and chloro- form, gave details of structure not obtained by other methods. ERIGERON. Two species, E. Philadelphicus L. and E. strigosus Muhl., were examined. The following statements apply to both, as there is no difference in their embryo sacs, except in size. The devel- opment of the embryo sac does not differ essentially from that of other Compositae. The number of antipodal cells varies, and the one nearest the endosperm nucleus is frequently binucleate. The lower polar nucleus moves up and unites with the upper polar nucleus ( fig. r), which remains near the egg. The nucle- oli are very conspicuous, with highly refractive bodies scattered through them. The mature egg is pear shaped, its nucleus being very large (Jig. 2) and showing a fine network of chromatin. A vacuole near the middle of the egg is very conspicuous. The synergids are pear shaped, and the position of their nuclei is variable. The end nearest the egg invariably wets large vacuole, while the smaller end stains deeply and sometimes has a distinct constriction (fig. 2) between the nucleus and tip. | The embryo sacs in the ovules upon the edge of the head are larger than those at the center, several cases being een Which the sacs in the center of the head did not function. ; E. Philadelphicus the usual length of the embryo sac when reac} for fertilization is about 216 pw, the width being about 4° aoe endosperm nucleus at this stage is from 14.4 to 19.8 # 10 idom eter, its nucleolus being usually gu. The egg nucleus S€ varies from 10.8 w, with a nucleolus measuring 5.4 #- The sperm nucleus is usually about 16m distant fro Chamberlain has shown that in Aster Novae-Angliaet F : he nucleols a 1900] DOUBLE FERTILIZATION IN COMPOSITAE 255 of the endosperm nucleus is very constant in size. Many meas- urements of the egg nucleolus of &. Philadelphicus were made, and in only one instance did it vary from 5.4 M. The male cells were not observed in the microspores, and no pollen tubes were found except in the embryo sac. When the pollen tube (¢, fig. 3) enters the sac, the synergids rapidly disintegrate, so that by the time fertilization is effected only fragments of their nuclei are visible. One male cell fuses with the egg nucleus, and at the time of fusion (fg: 3) cannot be distinguished from the egg nucleus. The other male cell fuses with the nucleus which is formed by the union of the polar tuclei, the product of this second fusion being the definitive or endosperm nucleus. No male cells were observed in the pollen tube, or in the embryo sac before fertilization, so that nothing can be said con- os their previous appearance. The pollen tube, after it has discharged the male cells, usually contains two bodies of irregular 7g Vig. 3), which stain intensely with cyanin. They were ae in preparations of Lilium. These bodies may be plete MIGULA: System der Bakterien 2: —. 1900. 4 NEW CHROMOGENIC MICROCOCCUS 267 regard to the two points most evidently characteristic of this form, its pigment coloration, and non-development on potato. Migula describes eleven non-liquefying micrococci as Rot wachsend. Of these, seven may be thrown out, since they are distinctly said to have a seal, or carmine red, or vermilion pig- ment, Another is “gray on agar, red in milk, and does not grow at 37°.” The other three are said to be “flesh red,” the exact color of which is doubtful. Of these: (1) M. carneus Zimmerman is described by Zimmerman him- seli*as having a deep flesh red, almost violet tinge, and growth potato. Lustig describes a form, Coccus ruber,5 found in water by Maschek which is probably identical with this. (2) M. sarcinoides Migula (n. sp.)® looks often like sarcina, Itis also described as a very large coccus, 1.8—2.5 » in diameter. Nothing is said of the potato growth. Gelatine plate colonies feach 5™" in diameter. Color on agar is a warm flesh red, more mense along the middle line. Bouillon has a flocculent whitish srowth and a red sediment. This description, I think, hardly comcides with that of the salmon-pink form, although much Pends upon what is meant by ‘flesh red.” (3) M. rubellus (Migula)s is morphologically scarcely distin- Suishable from the last, and shows the same sarcina grouping. ‘ a are almost opaque, dark gray-brown. The descrip- of . gelatine stick culture has some resemblance to that ate “an form, 2.¢.,a dry surface colony slowly spread- act als of tube and having underneath a deep dry cavity of ot gelatine. But this growth is said to bea “bright red.” All cultures have a trimethylamine odor; on agar 8s are finely dentate, and bouillion has a red slimy ough points of difference to distinguish the forms. class of germs which Migula describes as liquefying Mine i out of the twelve named that three have we 2 Babes). hispiiat (MW. rubens, M. persicus Kern, M. haemat . hes Swine red (M. rubiginosus Kern); one has a vio : akterien unserer Trink- und Nutzwiassser. Chemnitz, 1890. tik der Bakterien des Wassers 40. 1893. $Op. cit. Streak the ed — Sediment, en 0 the Selatine 268 BOTANICAL GAZETTE [ocToser tinge (M. roseo-persicus) ; and one (M. fragilis Dyar) is “brown red on glycerine agar.” These colors are not likely to be mis- taken, and cut the list down to those which are either “rose”’ or “flesh red.” Three of these (1) IZ. roseus Fliigge, (2) M. roseus Eisenberg, and (3) MW. voseus Maggiora are sometimes confused because of the similarity of name. (1) M.roseus Fliigge is only briefly mentioned by Fliigge him- self,’ but is described more at length by Macé® as a form com- mon in air, non-liquefying on a gelatine plate, slowly liquefying ina stab culture. But he says that the liquid portion is tinted red “more vermilion than by prodigiosus.”” Macé also mentions among species closely related to this, MZ. agilis Ali-Cohen, M. rouge cerise of List, isolated from water and producing a cerise color on potato, and Zimmerman’s M. carneus which Mace, a well as Lustig, regards as identical with Maschek’s Coccus ruber. He describes these as all non- motile cocci, about 0.8 in diameter, “rose red or flesh colored,”’ and abundant on potato. (2) M. roseus Eisenberg,’ to which Migula gives the name of M. subroseus, is found in sputum and produces a pigment like that of B. prodigiosus on potato. (3) M.roseus Maggiora is described by Ward ® asa non-lique- fying form 0.6 in diameter, associated in irregular glomeruli, and forming a pale rose pigment. I was unable to gain access ws further description of this micrococcus. As to the remainder of Migula’s list there is left: (4) M. cumulatus Kern, a too definitely liquefying for identified with the salmon-pink micrococcus. Otherwise, excePt ing the fact that it is a facultative anaerobe, the descriptions a somewhat similar. ae (5) MW. subcarneus Migula shows in a gelatine stab pee after one day a small light red colony at point of aa After three weeks a sirupy liquefaction begins in stocking-sh@ m to be 7 Die Mikroorganismen 185 [3d ed]. 8 Traité Practique de bactériologie 432. 1891. ° Bakteriologische Diagnostik 408. 1891. “Annals of Botany 12: 309. 1898. 1900] A NEW CHROMOGENIC MICROCOCCUS 269 form, with the colony on the surface and a thick, flocculent, rose- colored sediment. On agar, ‘‘rose”’ or “flesh colored.” (6) M. rosaceus (Frankland) is found in air. The entire description is as follows :** Cocci which are very variable in size, the largest 2.5 in diameter. On gelatine forms a smooth pink expansion on the surface, while needle track below remains almost undeveloped. In older cultures the margin assumes a tadiated appearance, while still older cultures frequently exhibit slight liquefaction. On agar, smooth bright pink surface expan- sion, devoid of any further character. Broth after nine days is clear, free from pellicle, and has a pink deposit. On plate, the colonies appear to the naked eye as pin-head dots on surface, and are bright pink in color. Under low power ( X 100) are seen to be of distinctly reddish tint, the edge irregular but smooth. As the colonies approach the surface, the irregularity diminishes. Certain points in this description agree somewhat with that of the Mississippi river form, 2. ¢., the gelatine culture and plate colonies, but the size of the coccus, the agar and broth culture seem to differ, and the description is too imperfect to make the eterison at all complete. The same may be said of Frank- land's description of his (7) M. carnicolor, the only stated points ° difference from (6) being (a) amore rapid growth, (6) fainter fa of pigment, and (c) slightly different appearance of colo- the (more circular and brownish-pink). In both these forms ala cultures are said to be clear after nine days, while ci cultures of the salmon-pink form have remained cloudy una g oneludes the list of red growing cocci which I wee ; Scribed, except that mention might be made of Bumns ar oo oseus and Sarcina rosea (Schroeter), both Bei git will ap s : aa Sarcina mobilis (Maurea),” which pire © | already 2 “0p on potato, but has a brick-red pigment. I have nee ntioned J. agilis in the body of the description. — Roy. See. London 178 : 269. 1887. "STERNDERG . large . » Manual of Bacteriology 720. 1893. 270 BOTANICAL GAZETTE [ocroser It is evident that the color of the pigment produced by these germs, otherwise so similar, is of some importance in an effort to distinguish them. The descriptions are, for the most part, very meager in actual definite points, and it is doubtful if even the twenty-six standard tests given by Fuller™ as showing a high degree of constancy would serve to differentiate these forms, so general in character are the indicated reactions of the table. In regard to the chromogenesis, it would be of no advantage to multiply the list of adjectives, for few bacteriologists could describe in this way the exact tints of “pink,” “reddish” “rose,” oF “flesh color” with sufficient accuracy to make them unmistaka- ble. An article by E. B. Shuttleworth,*3 on “ Nomenclature of colors for bacteriologists’’ is one of the most recent attempts to systematize colors for bacteriology, This author’s classifica- tion gives under yellowish-pink three subdivisions: salmon, salmon-buff, and flesh color. I was unable, however, to obtain the plates or illustrations to which his numbers evidently referred, and without them the words do not give a very distinct visual image. . A simple and more exact method of determining the pig: ment color of germs is that of the use of a color wheel. Alittle contrivance of the kind often used in morphology and easily obtained is a wooden top with the various small color disks. A better determination, because of the more constant rate of revo- lution of the disks, can be made with a Bradley color whee standard Maxwell disks. By this means I obtained the following result in the case of the salmon-pink germ. An agar culture begins in a cream-pink grow ie determination is not necessary. After about 5-7 days it Ge to acquire the characteristic color, the composition of the ©? wheel for this age being near: red, 70 per cent.; white, - He cent.; yellow, Io per cent.. Later the color deepens with addition of a more orange tone, and remains almost cue: ) after two weeks for all culture media —agar (different reactions) “Jour. Expt. Med. 4: 609. 1899. “SJour, Amer. Pub. Herath. Assoc. 1895 : 406. | and th for which the PA Ee, ls lena oO ee eee ~ Bists or 1900] A NEW CHROMOGENIC MICROCOCCUS 271 gelatine, on which the orange tint is slightly more marked, broth sediment, milk ring. Several trials to match this with the color wheel resulted in a range of: (z) (2) (3) Me - - - = + 765 Jae yee White - - - = 10 14 13 Yellow - . > - oe 9 9 Orange - - « “ 6 ; 6 100 100 100 Of these, (3) was almost the exact tint of my cultures, with the one difference of a slightly duller tone than that shown by the actual growth, particularly of a rather bright gelatine culture. But any one of these combinations gives a good idea as to the characteristic tint. By this method of description of colors there would be less possibility of confusion. The great advantage would lie in the ‘ystematization of those chromogenic germs whose color is tirly constant on ordinary media for certain ages. For instance, the whole prodigiosus group produces a color in liquefied gela- = which may be variously described in the vocabulary of sche bacteriologists ; but the fact that it closely approxi- ay a color combination on the wheel of red 80 per cent., aaa per cent., is sufficiently definite. B. prodigiosus and B. tend alticus both give the above color to liquefied ‘gelatine in a ay culture. In addition to this B. ruber balticus produces a "Y Surface membrane of an entirely different shade, i. @., red ag cent., Orange 20 per cent. a. a difficulty pid this accurate determination of pigment duced Course, the liability of variation in the pigment pro- ee same germ on media made by different bacteriolo- It has on under different circumstances. aghie physical aig that if this organism is brought ees ‘ *cording ‘ soa ny a Series of broth and gelatine cu . ’ Media are not — suggestion,” such variations in stan “Jour. E We range. “xP. Med, 4 : 6009. 1889. 272 BOTANICAL GAZETTE [ocToBER Because of the peculiar salmon or yellowish-pink color of the pigment produced by the organism described in this paper, and because, although it is evidently closely allied to some of the pink micrococci previously described, it is not identical with any of them, I suggest for it the name Micrococcus roseus flavus. In conclusion, I wish to express my sincere thanks to Pro- fessor Edwin O. Jordan for many valuable criticisms and sug: gestions during the progress of this work. BACTERIOLOGICAL LABORATORY, UNIVERSITY OF CHICAGO. Rees Pe ee BRIEFER ARTICLES A NEW SPECIES OF NEOVOSSIA. (WITH ONE FIGURE) Durinc September 1899, while collecting near Colo, Iowa, I found that the ovaries of Phragmites communis were affected by a peculiar smut. The specimen proved so interesting that Dr. Pammel, to whom | referred the matter, sent material to Dr. Farlow, who replied: “Your spores are perhaps a trifle smaller and have a slightly more marked epispore than in Meovossia Moliniae. Still the differences are very slight, and without further study I should not know whether to call them Specifically different or not. You may have the material to “ermine this point.” Accordingly Professor Hume, who is working = the Ustilaginez of Iowa, after a thorough examination and com- Parison of the specimens, decided that it was sufficiently distinct for a neW species, ge genus was founded and distributed by von Thiimen,’ who tag Vossia. As this name had been given to a genus of grasses, Kor- ca changed it to Neovossia. Saccardo,? in his treatment of the oda refers this fungus to Zi//etia Moliniae (Thiim.) Wint., ‘mg o Vinter Dietel,* in describing the Ustilaginex and Tille- “tage Neovossia of Kérnicke. Massee,° in his ropes of rity ae Tilletia, refers it to Neovossia, saying : “This species differs return oe . - mode of spore germination, and must consequently Sia, and ¢ Aggie Quite recently, Magnus’ has discussed Neovos- Brefelq? ena it separated from Tilletia by the fact, discovered by _. ? that in germination the conidia do not copulate. He also Yon Thiimen has well founded the genus upon the fact t M : 2 Yeotheca universalis 1216, 1878; Oester. Bot. Zeitschr. 29 : 18. 1879- Syl * Bot. Zeitschr. 29: 217. 1879 Sap Fung. 7; 486. 1888, 4Rabenh. Krypt. Fl. Pilze 1+ 109- 1884. egg Nat. Pflanzfam. x : (Lief. 160). 1897. “oy. Gard. Kew 153. 1 8 ther. & dex 53, 154. 1899. t. bot. Gesells, 18: 73. Ig00. 8 Unters. Mykol. 12: 210. 273 274 BOTANICAL GAZETTE [OcToBER that the ripe spores, with the upper end of the sterigmata, break off from the full grown sterigmata. There seems to be sufficient reason, therefore, for establishing the genus Neovossia. Neovossia Iowensis Hume and Hodson, n. sp.— Spore mass filling the ovary, black; spores globose, subglobose, or ovate, brownish-black, opaque, 16 X 20 to 24X28 p, enclosed in a hyaline capsule; appendage slender, hyaline, two or three times the length of the spore ; epispore appar- ently pitted. A careful comparison with specimen no. 1216 of von Thiimen’s AMycotheca univer- salis leads to the belief that the Iowa specimens are specifi Fic. 1.—NEovossiA IowENsIs ; a, spike- cally distinct. The spores let of Phragmites communis; 6, affected ovary ; differ from those of Neovossia ste Moliniae (Thiim.) Korn. in being darker in color, broader and shorter, and generally blunter at the end opposite the appendage. The markings of the spore, also, are somewhat coarser. Ten spores of von Thiimen’s specimen, selected at random, gave an average of 27.7X17p, while the spores take? from the material collected at Colo, Iowa, gave 24.8X18.9 B— E. R. Hopson, Ames, Jowa. NOTE ON THE ORIGIN OF TANNIN IN GALLS. THE origin of the different plant constituents is as much a mystery as their functions, and neither of these questions can be settled of more observations have been made. In considering the Bone” tannin in galls the writer limits his observations for the present !0 © examinations of the common “ink-ball’’ or “ink-gall,” — Te : % duced on Quercus coccinea Wang., probably by Cynips aciculata The same kind of gall is produced on other oaks, as Mr. ee ol : the Biltmore Herbarium, has sent me specimens which were Pr on Quercus imbricaria Michx. ee *Presented at the New York Meeting of the American Association : Advancement of Science, June 1900. 1900] BRIEFER ARTICLES 275 I. These galls are produced during the summer months on the young branches and sometimes on acorns. When mature they fall from the trees and are nearly globular in shape, varying from 20-30" indiameter. They are solid throughout and of the consistency of the pulp ofa green apple. Externally they are smooth, and in color are a mottled green, yellow, and brownish-red. At this stage they are made up of three distinct zones: (1) a central zone, made up of nearly isodiametric parenchyma cells packed with numerous small somewhat spherical or irregular starch grains which are colored blue with iodine; (2) the middle zone, composed of radially elongated parenchyma cells, possessing thick cellulose walls with prominent simple pores and containing a mass of protoplasm lying on the sides of the walls and afew starch grains (with the development of the egg of the insect there also appear in the cells of the middle zone numerous starch grains closely resembling those found in the central zone); (3) an ‘ternal layer made up of irregular parenchyma cells somewhat collen- chymatic in character, with the protoplasm as in the middle zone. Il. Decided changes take place soon after the galls fall from the ane (1) A larva develops in the central zone and there are signs of aavity in the protoplasm of the cells of this zone. A large nucleus = nucleolus lies centrally in the protoplasm and in some cases yel- Wish globular vesicles are apparent. These latter are fixed in the - Specimens by means of copper acetate (7 per cent. solution), age treatment they become more yellowish in color an i eS ” chloral, glycerin, potassium hydrate, or which, «c ) doubt in the nature of tannin vacuoles. (2) In he galls ntain a larva, and have been allowed to remain im Cope? Vig Solution for several weeks or months, there separate in the par- line oes a the middle layer or zone yellowish crystals or ine i 8, which are insoluble in water, chloral, glycerin, or alcohol. fan. a. and crystalline bodies are lens-shaped, star-shaped, “4 din, inulin, and chee resemble the different carbohydrates, as se nes are ie which separate in certain plant cells when the spe : in ay aced 7” alcohol. These crystals, however, do not Hepae '€ material, and are to be found only in galls which have ag popper acetate solution. Their appearance, ae that ¢, eon with copper gallate crystals lead to the conclus : *y are identical in composition with the latter salt. (3) in Me layer or zone of specimens which are at this stage of maturity, 276 BOTANICAL GAZETTE [octoper and have been treated with copper acetate, reddish-brown, amor- phous, or somewhat crystalline masses are found adhering to the walls of the cells. These masses when amorphous are made up entirely of tannin and when somewhat crystalline contain an admixture of tannin and gallic acid. III. When the winged insect has developed, (1) only a few layers of the cells of the central zone remain, and these contain a number of tannin vacuoles. Surrounding the latter are several (as many as 12) rows of prominent lignified cells. (2) The cells of the middle layer in specimens which are of this age and have been treated with coppét acetate, contain numerous brownish-red tannin masses to which may be adhering some yellowish-brown crystals of gallic acid. But the tannin is in by far the greatest quantity in the cells of this layer and at this age of the galls. (3) The cells of the external layer also contain tannin masses. Conclusion.—(a) It is well known that gallic acid occurs naturally in the nut galls (the product of Cynips gallae tinctoriae Olivier on Quercus Lusitanica Lamck.) ; the leaves of Arctostaphylos Uva-urst (L.) Spreng. Thea Chinensis Sims, and of various species of Riws; the fruit of Laiaas nalia chebula Retzius(Myrobalans), and Caesalpinia coriaria Willd. (Divi- Divi); the acorn cups of Quercus A¢gilops L. (Valonia); and may be obtained by extraction with water in the form of silky needles and asymmetric prisms. With the alkalies, alkaline earths, lead and copper salts, it forms crystalline compounds. (4) Tannic acid, on the other hand, is an amorphous substance and does not produce cry" line compounds with the salts mentioned. (¢) Therefore, the cys line compound found in the galls examined by the author is 12 probability gallic acid. This appears to be formed at the mee of the starch during the chrysalis stage of the insect. Mage maturing of the winged insect this is changed to tannic acid. ; transformation of gallic acid into tannin appears to be one of pee condensation of two molecules of the former with the loss 4 molecule of water, as follows: 2C,H,O, (gallic acid) =“ pe (tannic acid) + H,O.— Henry Kraemer, Philadelphia College of Macy. . : i. CURRENT LITERATURE, BOOK REVIEWS. The Cyclopedia of American Horticulture. Tue second volume of Bailey’s Cyc/ofedia* has just been issued, including the letters E-M. The general design and scope of this important work of reference has been set forth already in these pages.? To this nothing need be added beyond the statement that the second volume is even better than the first, a natural improvement as the editors attained greater familiarity with their work and materials. Aside from the articles treating large and horticulturally important genera like Gladiolus, Iris, Mamillaria, etc., there are some notable articles on gen- tral subjects, which serve to illustrate well the wise plan of the book and the thorough treatment of topics. Forcing is discussed in its general aspects by tssor Bailey ; forcing vegetables, by C. E. Hunn; forcing fruits, by William Turner; and forcing hardy plants, by B. M. Watson. Ferns are teated botanically by L. M. Underwood ; growing hardy ferns, by Edward Gillett and F. W. Barclay; growing tender ferns, by N. N. Bruckner. Again, in the article Grafe, the editor writes of the general and historical aspects, while specialists in the various grape-growing regions of the north, the South, and the Pacific slope write of the practical phases of grape culture, and Sethe of grape growing under glass. In the same way the horticultural and historical features of the Greenhouse are treated by the editor; Lord ox tg am write of its structural details; glass for it is discussed by J. C. Blair; the methods of heating by L. R. Taft. ee Among the biographical articles, the one on Asa Gray, by the editor, sat a of what Such writing should be, for here Professor Bailey we = bie and with clear insight and appreciation. Morphological articel oo Wanting ; but perhaps these ought not to be expected. . ever, that fruits might have been discussed from this point of /nsects are well treated, and in part morphologically wis RK. B, Bay age estions for cult; eal L. H.: Cyclopedia of American Horticulture, comprising sugs' vation 9 er has been, MY, Slingerland,—— ‘ its, vegetabl a f horticultural plants, descriptions of the species of fruits, ¥ sr Btogra s *mamental plants sold in the United States and Canada, toge Ae Ses. 5 oe biographical sketches. Vol. II. E-M. 4to., PP: xiv-+ 5 e 94-1453, pls. 1o~7 ‘ 9 ew York: The Macmillan Company. 1900- $5- Bor. Gaz, "500] 29: 282. 1900. 277 278 BOTANICAL GAZETTE [ocTonER Tobacco. : DER TaBak, by C. J. Koning, treats of the commerce, manuring, cul- ture, anatomy, diseases, and fermentation of tobacco. Well-known facts are here presented in a popular manner to those interested in the culture and trade of tobacco. Only the contents of the chapter on the so-called fermenta- tion of tobacco are essentially new. This process the author ascribes to the action of bacteria, in accordance with the hypothesis of Suchsland, “The aroma of the tobacco is caused by facultative anaerobes in so far as we cat speak of aroma in our Dutch tobacco.” In this remark, from page 23, the bacteria are called “facultative anaerobes,” while on page 53 we find the statement that the supposed principal generator of the aroma is an obligate aerobe. The reviewer is sorry to differ from the views of Koning, yet he has examined very carefully, with very high magnifying powers, wrapper leaves withdrawn directly from the interior of the fermenting heaps @ Florida, without discovering any colonies or any coating of bacteria. The few isolated rods and cocci found on some square centimeters of leaf cannot possibly have any significance. The water content (18-25 percent.) of fer. menting wrapper leaves prevents not only the development of fungi, but still more, that of bacteria. Filler leaves are more heavily moistened, and ngs contain 35 per cent. of water, but even these do not show bacterial colonies when heated in closed vessels to 55° C., a temperature reached often by the heaps of fermenting tobacco leaves. Even 60° C. is often reached, and nevertheless the heaps will heat up again when taken apart and rebul afresh. This would be impossible if the Bac. Tadaci really generated the aroma, since this dies at 50° in thirty minutes, and at 60° in five pa according to Koning’s own statement. At the same temperature also : Diplococcus Tobaci Hollandicus, which Koning claims improves the com: bustibility of tobacco, also succumbs. f the n not a single instance has the author stated the water content ee tobacco when he started his bacteriological investigations. Tobacco ao to 60 per cent. water will no doubt readily develop bacteria, especially hers. : Proteus group which Koning found ; cocci also will thrive in great num a Should this rotting be interrupted at the right time, a change of the might have occurred, which may be very desirable-with Dutch ore a such rotting is carefully avoided by the progressive American peer ufacturers, by keeping the water content so low as to avoid the action ‘al teria. Under these conditions alone the oxydizing enzyms of the py - will develop their activity, and to this the changes of odor and aroma a teristic of superior tobacco have to be ascribed.4— OSCAR LOEW. , sual $KoninG, C.J.: Der Tabak, Studien iiber seine Kultur und pegs 4to, pp. 86. Amsterdam: J. H. & G. Van Heteren. Leipzig: Wilhelm : oo 1900. M 4 (unbound), * Compare Reports No. 59 and 65 of the U. S. Dept. of Agriculture. =e Weis he NN eRe ie ire Tene aaah peak taree ee ar WE feo (Mita Gel p = Titerrg Pyne Ss Se ease a : aR ey edie sat moe rua Caldiera me Rane we) Sent 2 1900] CURRENT LITERATURE "279 MINOR NOTICES. THE FIRST SUPPLEMENT to Paris’s /ndex Bryologicus has been issued by the house of Georg & Cie, Geneva, as one of the Mémoires de 1’ Herbier Boissier, It contains 234 pages, with innumerable entries correcting and adding to the original 7ndex.-—C. R. B. Paes THE FOURTH PART of the new edition of Weisner’s Die Rohstoffe des __ Phanzenreiches completes the account of plant fats (42 pp.); Dr. K. Mikosch _ treats vegetable wax (21 pp.); Dr. A. E. von Vogl writes the tenth section on camphor (6 pp.); Dr. S. Zeisel contributes the chapter on starch (80 pp-); and Dr, Lafar’s section on yeast is begun (11 pp.)—C. R. B. ‘THE SECOND FASCICLE of the list of the genera of seed plants, according _ tothe system of Engler, has appeared with remarkable promptness.° In the notice of the first part in this journal’ the general character of the work was stated. In the present signatures 1220 genera are listed, bringing the number _ §p to 2490, the list beginning with A/opAza (Iridacez) and ending with Silene iS (Caryophyllaceae),— a MC, RECENT NUMBERS of Engler’s Die natirlichen Pflanzenfamilien are as llows: Number 198 contains a continuation of Muscé, by Carl Miller and W. Rubland, and deals as yet with the general morphological characters of the group. Number 199 contains the conclusion of the Marattiacee by G. ane : d Ophioglossaceze by the same author, and a general discussion of : ales by H. Potonié. Number 200 and 201 (a double number) con- : oe ricton of the Hyphomycetes, by G. Lindau, and with it bees ace n of the part of the first volume which deals with the Fungl.— & parenchyma cells, or of other cells differing but little from es. It never occurs in cells containing oxalic acid (as oxalate). whi 3 % dissolved in the cell sap in young organs, but is an amorphous * the parts. The alkaloid is formed in the leaves and transported to ; where it is stored either in its original form or, after transformation Gaz, - se 1640, figs. 89-122, Leipzig: Wilhelm Engelmann. M5. See BoT- * rer, 4900 ; ‘Da ‘ 2 | Rgleri at TORRE, C. G. De, and Harms, H.: Genera Siphonogamarum ad edi Mo Lei zi eg Fasiculus secundus (signatura 11-20). Small 4to, PP- ‘ Oa ig: L Im Engelmann 1900. : SS Gaz, 30: 67. 1900. eti ’ ° m de l'Institut botanique de Buitenzorg. III. 8vo, pP- 43: ~ 280 BOTANICAL GAZETTE [OCTOBER to another alkaloid. It is considered not at all impossible that the alkaloid is formed by direct synthesis, and not as a decomposition product of pro- teids.—C. R. B. A SMALL PAMPHLET by W. Johannsen presents an account in popular form of a process which has already attained considerable use in forcing: houses. The method, which extraordinarily hastens the development of shoots and flowers, consists in exposing dormant plants in a suitable chamber twice to ether vapor for 24-72 (mostly 48) hours, with 48 hours interval, the time depending on the temperature and phase of the resting period. first section, “Zur Orientirung iiber die Ruheperiod,’”’ has con- siderable theoretical interest. The author defines clearly the expression “resting period,” and shows the erroneousness of the common idea t the riper the wood or seed of a plant is, the easier will be the budding * sprouting. He points out that the resting period has no sharp limits, but is a passage from diminished power of growth through complete rest to increased power of growth again. In the second section detailed directions are given for the practice, is especially applicable to syringas, azaleas, snowball, spireas, deutzia, the valley, and tulips.—C. R. B which lily of THE FIELD CoLUMBIAN Museum has recently come into possession of a set of plants collected by Don José Blain on the Isle of Pines, Cuba, some time in the middle of the sixties. The list, including 185 numbers, — which are four new species (Polyga/a, Salacia, Spigelia, and Heliotropium) has appeared as one of the publications of the museum (1:425~439 1900) under the title “Plantae Insulae Ananasensis,” by Charles F. Millspaugh, title “Plantae Utowanae’(Field Columb. Mus. Bot. a: 113-135- 1900) which he reconsiders the Cyperacee and Cakile of the former ne GAZ. 29:360. 1900). The present paper takes up the two groups | fe form to be used in the proposed Yucatan Flora, in which all of the spe a descriptions are to be based upon the characters of the fruits, and — a by text cuts illustrating these characters. Dr. Millspaugh differs ON Clarke in regarding Mariscus and Torulinium as but sections of aie q Cyperus (Mariscus) Caymanensis is described as a new species. In | ten species are recognized, two of which are new; also two n described. The author makes the very interesting observation th nee has laid special stress upon the development of the fruit for disse ae in the q and that the “evolution for floatage seems to have reached its height "" new species growing upon the Alacran shoals.’’ — J. M. C. — 9JOHANNSEN, W.: Das Aether-Verfahren beim Frithtreiben me eee ’ ote cartailagad der Fliedertreiberei. 8vo., pp. 28, #5: # jena : : 900. fy, ; 1900] CURRENT LITERATURE 281 STIGMONOSE is the title of a bulletin by Albert F. Woods® in which he s fully his studies of the disease of carnations and other pinks, formerly called bacteriosis by Arthur and Bolley, and ascribed by them to the action of Bacterium dianthi. The preliminary statement by Mr. Woods ato the cause of the disease and the tone of his criticisms on Arthur and Bolley's work * were criticised by this journal *. Mr. Woods has now presented the evidence on which his conclusions rest, and it entirely justifies the sub- stance of his criticism. Moreover, the account of Arthur and Bolley’s work in the bulletin is full, and the defects in it are pointed out in a way to which no exception can be taken. Woods shows that neither fungi nor bacteria are present in the earlier siages of the disease, and though they may appear later, their presence is not shown repeatedly by colonizing aseptically these insects on carnations. As “bacteriosis” is misleading, stigmonose is suggested to replace it. Mr. Woods believes “that the insect injects some irritating substance of an acid a enzymic nature into the wound; that this substance causes the increase of ouidizing enzymes in the cells which it reaches, and that these enzymes interfere with the nutrition of the cell by destroying the chlorophyll and set- — ling up other changes which finally result in death.”—C. R. B. va FRENCH TEXT on the anatomy and physiology of plants by Er. Bel- 8 . St gric., Diy, 00 nu A.S., Toronto meeting ; see Bot. GAZ. 24: 200-205. 1897. a Gaz, 95: 129-130. 1898, MUNG, ER.: Anatomie et physiologie végétales, al’usage des étudiants ¢m frien des universités, des éléves a J’institut agronomique, in 6 a , ete. 8v0, pp. iv-+ 1320, figs. 1699. Paris: Felix Alean. 1900 282 BOTANICAL GAZETTE [octosEr announced purpose. . For example, the ‘“‘circulation”’ of the sap is repeat- edly described and impressed by a diagram with arrows showing the diree- tion of the “ascending sap’’ and the ‘‘descending sap.” The “osmotic force’ is presented as “une nouvelle force,” residing in the protoplasm ia virtue of which it exercises “une puissante attraction.’’ Many other similar cases might be cited from all sections. The best thing about the book is the illustrations, most of which are excellent. But as a whole it can hardly be commended.— o- A NEw Part (second series, Part IV) of the A/iumesota Botanical Studies has appeared, bearing the date August 15, Igoo0. It contains seven papers of varying length, and is altogether a worthy member of the series. “A col tribution to the knowledge of the flora of southeastern Minnesota,” by W.A. Wheeler, is in the nature of a report of the work of the State Botanical Survey during the summer of 1899, and the results are presented with a well- -organized ecological background, accompanied by seven excellent plates from photographs showing cl teristic vegetation features. ‘“ The seed and seet- ling of the western larkspur (Delphinium occidentale), by Francis Ramaley, is a brief morphological and histological study, illustrated by a plate. “A ‘preliminary list of Minnesota Erysiphez,” by E. M. Freeman, catalogues nineteen species, with their hosts. K.C. Davis publishes three important -revisions which have been developed in connection with the work on Pro fessor L. H. Bailey's Cyclopedia of American Horticulture. They ate fol lows : “ Native and garden Delphiniums of North America,” 52 species being described, one of which is new ; “ Native and cultivated Ranunculi of North America and segregated genera,” the genera being Batrachium &. F. 5 (5 spp.), Ranunculus L. (96 spp., two new), Kum/ienia Greene (1 sp.), e Huds. (1 sp.), Cyrtorhyncha Nutt. (1 sp.), Arcteranthis Greene (I pe a ve the conclusions reached the following is of general interest: young mycelial threads very good evidence of the occurrence 0 previous to, or in intimate connection with, the formation of the sp —jJ.M.C, NOTES FOR STUDENTS. NAWASCHIN* has recently made a cytological study of brassicae Woron. Plasmodiophora is a parasitic myxomycete various deformities and distortions in the roots of its host. Plasmodiopho? which caus brassicae ® ™Beobachtungen iiber den feineren Bau und Umwandlung vom # 427 plete Brassicae Woron, im Laufe ihres intracellularen Lebens. Fora 86 + 404 = 1899. ‘ 1900] CURRENT LITERATURE 283 the cause of “club root” in cabbages and turnips. Woronin, who studied this form more than twenty years ago, found that it does not form sporangia, but that the spore masses lie free in the cells of the host. Eycleshymer, in Jour. Mycol. 7: 79-88. 1892, gives a clear account of its life history and distribution in the United States. The present paper is concerned with the more minute details, Infected roots were cut into very small pieces and were treated with Flemming’s stronger solution for twenty-four hours. The author thinks that the advantages of a more prolonged treatment are entirely imaginary. Sections were cut 2-3 # in thickness, and were fastened to the slide with distilled water without any further fixative. The Flemming triple Stain gave the best results. Dilute Delafield’s haematoxylin, followed by eosin in clove oil, and also the gentian-violet method according to Gram, save good results. The peculiar method of nuclear division in the vegeta- le amoebae is worthy of special mention. In the resting condition this mucleus has a membrane, a nucleolus, and an extremely delicate chromatin network, As division begins, clearly differentiated chromatin granules appear inthe place of the network, the granules having no genetic connection with the nucleolus. A plate evidently derived from the chromatin granules is then we hear the nucleolus. At this stage there is a sort of one-sided “ achro- matic figure” with its base resting upon the chromatin plate and its apex at the nuclear membrane, but the figure afterward acquires a symmetrical “spect, the drawings in the plate bearing considerable resemblance to bipolar tg The nucleolus then divides transversely, and the two resulting . hia Positions on opposite sides of the chromatin plate, Bydeoee » now Panetta disk with chromatin granules imbedded in its Pee ie plate then splits, and the two parts, each accompanied by a » Move toward the poles of the spindle, and two daughter nuclei are _ This method of division is of greater interest because the nuclear type, in the plasmodium is simultaneous and of the usual bipolar mitotic § a author's summary of the entire paper is about as follows — of infected cells arises by the repeated division of a primary division ea : : : S a the growth of the infected cells numerous multinucleate 3. fa *ppear, which multiply without fusing to form a plasmodium. ir lag Condition the amoebae of Plasmodiophora are remarkable for 4. Sagat and especially for their unique mode of nuclear division. in oth © mode of nutrition of the amoebae seems to be different from that she aeomycetes. Place in og coos fuse into a plasmodium, characteristic changes take ‘4 The § Structure of the body and in the nuclei. oe *xhausted, ormation of a plasmodium takes place only after the host cell 1S ‘ 284 BOTANICAL GAZETTE [ocrosEr _ 7. Inthe mature plasmodium spore formation is preceded by repeated \ nuclear division of the typical mitotic sort. 8. In the first period of its development the parasite does not kill the host cell, but merely causes it to hypertrophy.— CHARLES J, CHAMBERLAIN. HERBERT J. WEBBER, who has long been investigating the subject of plant-breeding for the Division of Vegetable Physiology and Pathology, U. 5. Department of Agriculture, has published his results upon xenia in maize, which means the immediate effect of pollen upon structures outside of the embryo. His experiments are by no means complete, but they have already yielded suggestive results, and the author is justified in their publication by the fact that the subject has been brought under discussion by the discovery the explanation of xenia. Mr. Webber had already reached the same conclusion independently, and was collecting a large amount of experimental data to substantiate his more leisurely developing paper. It seems that the name “‘xenia” was applied to this phenomenon by Focke.* While it is claimed to be a somewhat common phenomenon in many plants there are very few cases on record that are not open to doubt, and in no plant is its occurrence so well substantiated as in maize. The experimen's of Mr. Webber have been conducted since 1898, in Washington and at the ‘Nebraska Agricultural Experiment Station. The greatest care was used to obtain pure races and to prevent the access of foreign pollen. Full details are given of about thirty experiments which yielded pertinent results, and the paper closes with their discussion. : The author abundantly confirmed Kérnicke’s statement that xe shown only in the endosperm. Color in the endosperm is frequently ap mitted by the pollinating parent, but color in the pericarp 1s ame chemical composition of the endosperm is also greatly affected by ue pollin- | ating parent, sweet corn crossed with dent or flint races producing re grains with starchy endosperm, and vice versa. All of the experiments ae the theory that xenia in maize is caused by the fertilization of the gene ’ nucleus by one of the male cells. All of the grains showing xenla — be true hybrids. In the case of spotted grains the author proposes an ap ; ing hypothesis by way of explanation. He suggests that the male nu may fail to fuse with the polar nuclei, and in such a case may a spindle and divide separately. In this event.two races of ie two be formed, and when the parietal placing and tissue formation beg age 4 races might become intermixed. A second hypothesis éxplaining pats me homenon suggests that the male cell fuses with but one of the i ee nia is *5 Xenia, or the immediate effect of pollen, in maize, Bulletin 27, — (one colored). Issued September 12, 1900.‘ : **Die Pflanzen-Mischlinge 511. 1881, 1900] CURRENT LITERATURE 285 the other polar nucleus dividing independently. - This also would result intwo races of nuclei which might become more or less intermixed before fixation in a tissue. We await with expectation the minute investigation of the structures con- cemed, which should settle the question of double fertilization for maize.— J. M.-C. THE POWER of the infusoria to adjust themselves to certain changes in their nutrient medium is the subject of a paper by Atsushi Yasuda.” From anutrient fluid in which they were growing normally, the infusoria were trans- ferred to solutions of the following substances: saccharose, glucose, lactose, glycerin, KNO,, NaNO,, MgSO,, KCI, NaCl, and NH,Cl, in varying con- centration, The organisms used for the experiments were Euglena viridis, Chilomonas paramecium, Mallomonas Plasslii, Colpidium colpoda, and Para- mecium candatum. In general, excepting Euglena, these can withstand about 6 per cent. of glucose, while Euglena withstands 1] per cent. With instead of 4°, as above. In the case of the electrolytes it is still greater, usually however less than unity. Aspergillus ntger withstands over 50 per “ent. of glucose and 21 per cent of NaNO, (Eschenhagen), and Zygnema Put has also deprived it of its proper food supply. This, it — an “ osmotic chan More conc 'sm, and its outline becomes uneven.— BURTON EDWARD Liv- Studien ; i oo _~i@leniber die A Wee i ee : n concentrirte Los- : Oe Joi, Coll. § npassungsfahigkeit einiger Infusorien an cones” ™ = ci. Imp. Univ. Tokyd 13: 101-140. pls. 20-72. ‘1900 the other sugars the ratio is nearly the same, but with glycerin it is about $ * 286 BOTANICAL GAZETTE [OCTOBER Dr. TH. Bokorny discusses ® the various modes of storage of proteids and their microchemical relations. Proteids soluble in 5~10 per cent. salt solution (globulin) are stored in the proteid grains and crystals (“aleurone” and “‘crystalloids’’) of seeds. Proteids insoluble in NaCl solution have not been observed in proteid grains. Neither “active proteid”’ nor fat could be detected in the proteid grain itself; the fat is associated with the general proteids of the seed, probably with the plasmatic proteids. The plant caseins seem not to occur in the proteid grains, for these dissolve completely in NaCl solution, whereas the caseins are not soluble therein. The glutens of the cereals constitute a special case; they dissolve in 70-80 per cent. alcohol, @ fluid which usually serves to precipitate proteids. Peptone was not recog: nizable in resting seeds. It and peptonizing enzymes occur in plants only exceptionally, as in fungi and carnivorous plants. Simple amides (aspara- gin, tyrosin, leucin, etc.) are well known in seeds and are widely distributed in vegetative parts. They appear to be the first decomposition product as well as the first formative stage of the proteids.— C. R. B L. Kny has been unable to find any traces of the intercellular living pro toplasm said by Baranetzki to be observable in the intercellular spaces of young stems of Myriophyllum spicatum and Ceratophyllum demersum, an intercellular protoplasm in roots of Matas major. Kny examined a number of water plants and says: ‘In no case was I successful in obse protoplasm (whether with or without nuclei or chromatophores) as the young or full grown air spaces except when its origin from the ing cells was in the highest degree probable. Even in the most advan geous covered preparations, in which the protoplasm within the cells adjoining the air spaces proved motile for several (in extreme cases fo sign of self-movement in the periphery of the air spaces was to ae existence of a living extracellular plasma in the large air spaces of sal plants must remain improbable until proof more convincing than at Pt is forthcoming.” —C, R. B a lining in surround: ‘ ‘ freien HERMANN FIscuEr concludes his paper on “ Der Pericykel in den Stengelorganen” ® as follows : res 1. In about 32 per cent. of the dicotyledons investigated a more °°” perfect endodermis may be recognized marking the distinctio tex and central cylinder. The so-called pericycle, dy tts fos’ limit of the cortex and the ring of vascular bundles is allied witht pine bium of the root. Considered histologically, genetically, and 3 ee : ’ ‘ an be region, no characters common to pericycle and pericambium can be P cated. hiladelphia: 994 Chestnut © t tra eg 74 State St. ————— Mem Fevers frequently follow the Fall house- opening, due to foul gases and disease-breeding matter developed during Summer, Platt’s Chlorides poured into waste pipes, sinks and closets, also sprinkled about the cellar and suspected places insures in- stant disinfection. Platts Chlorides, The Household Disinfec tant. An odorless, colorless liquid ; powerful, safe and cheap; endorsed by over 23,000 physicians; sold in quart bottles only, by druggists and high-class grocers everywhere. Prepared only by Henry B. Piatt, Platt St., New York $* gogo? 0 000@0e AND OCORZ x2 CHOCOLATE: SOLD AT OUR STORES Ano BY GROCERS EVER “IT’S ALL IN THE LENS” A Beautiful Picture Is Always Made With a Korona Camera and OUR FAMOUS LENSES Send for Catalogue GUNDLACH OPTICAL CO, 700 Clinton Ave., South, ROCH ESTER, N.Y. i il i al a a at ere nominal prices. Good durable pan as low as $100, warranted as yi pe Square pianos $20 and upwart from$z0o. Send for comp the makersare: Decker B Knabe, Steinway, Weber, ee : : a are interested ina plano ving exe ed at He as represented may expense. Address LYON & 100 HEALY d ’ ‘ ica8?” Adams st. on we rrr? en a + =) Town £ A ‘7 “MERICAN C FOR BRAIN AnD EK ADV'G AG'Cy 66 Staying Qualities.” , '“0ple won't be humbugged forever. It is getting to be pretty well that Quaker Oats is better than meat to build up the athlete s g to understan e wo is c . Guaee >and delicious makes ts, besides this daily _ 0 Mnovations an EREAL Co., fl sS ay it | keep Air spaces woven in the woo o pace in, the cold out, the skin dry rheumatism vent be llores and and sage bbe rie itacl Bee WRIGH 8) HEAL LTH UNDERWEAR (i FRANKLIN 5S Big Four Routt THE ELITE LINE A Le | pypranapouls ] CINCINNATI || LOUISVILLE | and | All Points The ** UNIVERSITY ” is a“ name of this, one of our latest eee Chairs. It’s a brain worker's chair. The back is . SOUTH and adjustable. The arms lift up a ond turn i forming ook, for 1 ie —— books, etc. Our Adjustable Rea ing Desk, is wanted, may be attache ed. SOUTHEAS We als he s of Reclining Chairs, 7.c., Rogsat’ Columbine, Siesta, ee and Orien te 1, ering id for either necessity , eg ury. Our new isles Cc Gust out) ceecthes th hem a SARGENT’S ECONOMIC c SYSTEM oF ‘DEVICES FOR AIN ORKE is also something worth — about. It embraces all the sT REE! ati als of the fitvest, includ erren Be ha Sena one t 234 CLAR RK * ook Cases, epeara-velioa, styles of Reading Stands at 2 tionary, Atlas and Folio Holder “4 mb area Reading Desks, TICKETS - _ a r sere attachable to chairs, etc, Catalo ogu lgtng i ed ae 0. F. SA ARGENT COMPANY, and 6 ‘gad, “goth, 534, 280 N. Fourth Ave. » Next 23d St., NEW YORK. A A re ee ee ee ee vy 66 ope The Death Certificate should have stated that he died of flannels.’—Dr. J. L. Milton, Senior Surgeon St. = & John’s Hospital, London. Woolen underwear is more deadly 3 in winter than in summer. Read the > proof at your leisure from our pam- y phiet. ' DR. DEIMEL'S. is the only Winter Underwear: @i which will free you from fear of 7 draughts and render you immune > from colds, catarrh, bronchitis, la > _-grippe, pneumonia, rheumatism, irri- ‘ ) tation and eczema, which woolen © underwear has subjected you to, and ® you will be supremely comfortable > too. Try it this winter. Ds Js SJ As AS Sa nl Ss Y Y ty As Ip ia A The Standard for Gentlemen ALWAYS EASY The Name “‘ BOSTON “weg CARTER *” is stamped on every loop. Samples and a common-sense pamphlet about it free. +> $ if the goods bearing this trade mark are not for sale in your community, we will aS Be f hs @& i: e24)} fF) 2 . ‘ =4 The fy ° (mis inform you where % hk : —_ they can be bought. - CUSHION . ‘ sitaghte © We also manufacture the finest dress ~ «_CLASP > Shield in existence. Can be washed, are i ; ir. Lies flat to the leg—never }| $ odorless. A guarantee with every Pa Slips, Tears nor Unfastens. é The Deime! Linen - Mesh System Co. @™ SOLD EVERYWHERE ‘ i | Zam, Semple pair Silk 50¢, Cotton 2c 4 491 Broadway, New ai é | Mailed iptof $ * itreal, : end > San Francisco, Cal. : therine Street © GEO. FROST co. ‘ 202 St. Ca | EVERY P Boston, Mass. rw : iil Montgomery _ ’ London, E. C. AIR WARRANTED-2a CEO 8 Washington, D. C. ® 728 15th Street, N. W- ei 6 é 9854405450 The Grand Prix (The Diploma of Honor-Highest Possible Award) itt wes Oe SS Che Smith Premier Cypewriter | AT THE PARIS EXHIBITION HIS award, made by an In- ternational Jury of twenty- | five members, was at a rating, on points, Higher than any of | the other twenty Typewriters | in competition, and, quoting from the jury's report, was | | Its Interchangeable Type given “for general superiority | Makes it the Most Useful of construction and efficiency | Teachers Clergymen. at the highest rating.” Send for new Catalogue, and inclose 45 | year for a aoe Map 0 of the Wor | tne the Back The Smith Premier Typewriter Co., | Ye a yreser tbe just fied ts the Hammon SYRACUSE, N. Y., U.S. A. THE HAMMOND TYPEWRITER CO) 537-553 E. 69th St» New Yo Ee | Dixon’s | ¢ American Graphite Pencils F ‘ 1 . L L ’ Made i { conceivable tae to ware mt style for x every y be p & penc ve : : 4 a pte your deal 4 ealer doesn’t keep th, ‘ 6c. for Samples worth double a —. Cruel J rey €) »N. J. i i a. + Y | S wwwy ESTERBROOK’S || “oth =! Qua, vers Service ui, RELIEF PEN No. 314. tase in Writing Unsurpassed 20 other varieties of stub pens. . 15 0 styles fine, medium and blunt points. . . ASK YOUR STATIONER FOR THEM. emington THE Standard Typewriter = Rega STEEL PEN Co, defies competition ” #00n St, New York, rks, Cam I Works, Camden, N. J. WYCKOFF, SEAMANS & BENEDICT 327 Broadway, New York, N. Y. ARS ARE ae THE BALL BEARINGS OF THE DENSMORE FOTECTION PENCILS] Located AT THE WEARING FOINTS; ON THE enn OF WHICH IN ANY MACHINE, CONTINUOUSL CHIEFLY DEPENDS. » 123 W, ithe STREET Main Office, 309 Broadway, satiad! les AF be | a” Kon LNOOR” Fp ie and ne lea Dene ce oe Srachomet C xX "THE train leaving Chicago a | 6.30 p.m. daily was spokes | of by the late Geo. M. Pullmas | as the “ Finest Train in the | a Reclining Chair Car and 3 Dining Car, as me hice trains are Pe fe) mate as City, Om aha and De ars. plan ere Car service is a speci feature of excellence on this i Delicate chtieg roses, rete neh perfect ventilation and strictly 5 class cooking. St a P a U : “Colorado Outings” “California” BEST LINE CHICAGO OR ST. LOUIS TO Are the titles of descriptive : ~ e booklets ets wat t charge upon applicate * Inneapolis = «<8: se ae Cc. B&Q. R.R., CHIC. ALTON RAILROAD. CHICAGO mc KANSAS CITY, CHICAGO «xo ST. LOUIS, CHICAGO 4x PEORIA, _ST. LOUIS «»oKANSAS CITY. man service between Chicago and HOT TOPRINGS, Ark., gris TEXAS. FLOR RIDA,U fac CALIFORNIA AND OREGON. ou are contemplating a fies any portion of sah a Lad md tis nde sn cago & aoe it will ets, rates, time-tables et en ee — GEo., P AUC bi te 6 re) rs "Suaties LTON, Rey Vere Paar Dee, Preset General P Passenger and Ticket i, | are & Ww, H. Mc cDo CHICAGO, ILLINO as. H. ickguand “Tr. Mer JISURE 1 the | RAVELERS, of Hartford, Conn. Life; 23 Oldest, Rost Endowment, an Best ana ACCident a _. . Insurance OF ALL FORMS. ‘Health Policies. Indemnity for Disability caused by Sickness. Liability Insurance. Manufacturers and Mechanics, Contractors, and Owners of uildings, Horses, and Vehicles, can all be protected by policies in THE TRAVELERS INSURANCE COMPANY. : iti-up Cash Capital, . . $1,000,000.00 ont ... 27,700,511-50 matics, | |. , . 23,730:027-0! MESS, 3% per cent basis, 4,020,083-95 7 - Insurance in force... $100,3341554:00 | “turned to Policy holders, . _ 39»734»920.59 | & ‘ TONHAM, Vice Prosi, J GS BATTERSON, President. ae oe 4 E resident, H. 3 MESSEN ana? ie MORRIS, Secretary. E, V, PRESTON, Supt of Agencis® al ADAPTABLE TO ANY STYLE OF RECORD OR ACCOUNTS spon WE VISIT EVERY TOWN IN THE UNITED STATES SALES OFFICES IN ALL THE PRINCIPAL CITIES BAKER-VAWTER COMPANY LOOSE LEAF SYSTEMS ATCHISON, KAN.-CHICAGO—HOLYOKE, MASS. Originators and Manufacturers of the Perpetual Ledger and other Labor Saving Systems IF YOU WANT THE MOST MODERN METHODS OF ACGOUNTING, LET US SYSTEMIZE YOUR OFFICE Leis amenvaew CHES) | USED IN ALL DEPARTMENTS OF UNIVERSITY OF CHICAGO FIVE OF'EM] §, FINE ‘UNS,TOO. (| Run Daily In EACH DIRECTION -setweenNEW YORK “ | Waukesha Hygeil iG v Mineral Springs ; b Water # # 4) av BUFFALO. 4 r 3 OF ‘EM & FROM IT 18 MADE Boro-Lithia Water, Gin- } ger Ale, and ST.LOUIS 47> Wild Cherry : hate ag, WANSAS CIT, poorer Dre s/ Vee. eK K AH Geaganna aitoad Ve LaCKdwald naliroad. a Vestibulefirains, CHICAGO, I | Comfortable Coaches, : Luxurious Sleeping Cars, = Fine Café Cars, Fare oomy P . Lackawanna}. Seomy Parlor Cars 4 vert THE Ww Water Go ailroad EG.RUSSELL, T.W.tEE. 8.0.CALOWELL, | aukesha ater i i CEN’. SUP y GENL PAS ACN TRAPIIC MANAGE ee ——— R. R. Donnelley & Sons The Lakeside Press , PRINTERS AND BINDERS [Sex ae, : ee) ee bed Me ee ep pie OF BOOKS DEMANDING TASTE AND CAREFUL ees eke ATE EDI seb aN Fete Beli ade TIONS, COLLEGE CATALOGUES AN OUSES bA a PRINT MORE FINE KS THAN ALL OTHER H WEST OF NEW U pena The ty) 18 THE MOST ADVANCED PRINTING PLANT IN THE CO O. | THE LAKESIDE PRESS BLDG., PLYMOUTH PLACE, COR. POLK, CHICAG i a <2; best Soa 2 . oF Stor folks who want their @ money’s worth of soap = &@L Don't buy soap and premium knick-knacks J Ugether, } wre likely to pay too much for your pre- Yi “tums and much too much for your soap. __habhitt’s Best Soap for nearly half a century fi lead all laundry soaps in purity and economy, ¥% “Lit still leads, and always will lead. _ \ Tour full money’s worth of soap-quality and {{S "whsting economy in every cake. . 0 Premiums %% and no need of them — the best sells on its merits. Made by B. T. Babbitt, New York WHAT IS POND’S=== =—EXTRAGT? A Family Remedy which, for over 50 years, has Pp awee a test of time. INVALUABLE FOR ALL AOHES, PAINS, INFLAMMATIONS, OATARRHAL TROUBLE AND sie GET THE GENUINE. AVOID IMITATIONS. POND'S EXTRACT CO., New York and London. LIEBI | 6 d é rf i 1 A y COMPAN?> EXTRACT si P Sa = = == SSS ——— SS 4 - Y | ~~ TH A” ay i aa anne lien “THE OR S eT Aone | Fegstered by Ui, Patent Office 2 ) Siz re q | HIA WATER OF VIRGINIA, Springs Nos. 1 and 2, IN COMPOSITION APPROXIMATES THE Blood Serum. A BLOOD FOOD AND NUTRIENT. John V.S nV. Shoem Medica Pi pene r; M.D. Et D., Professor of Materia delphia, etc, New Y; ot an the Medico-Chirurgical College of Phila- . An additional ads ork Medical Journal, July 22, 1899 (extract). : PECULIAR EFFICAC vantage and extremely important reason f be that its compositi Y of the BUFFALO LITHIA WATER ies in the Metefore it is Aeon approximates that of the Serum of the Blood; 6 iate Meoiporatio fitted for absorption into the blood current and me somes tk éace 3 sate ie the watery portion of the nutrient fluid. m7 ich far surpas em with the BLOOD SERUM. These are quali- , Sites chemical ite ose possessed by any extemporaneous sat Water for i paration, as when a lithia tablet, ©. g-. 5 dis- ® It is of a ‘ae spies administration. When we speak of y altogether relative, and what the physician i This we | “Sphatically dec; ave in the desires in a dose is therapeutic efficiency an a vi arefully noted its ef- d from amounts 50 the accompanying tiess to “Those who. ‘FALO WATER > mall ve often been a ade use of this water and c : Wiens me aoe at the results obtaine “WS: The explanation of this extreor Lithia and Bo in the conditi of this extraordinary activity is dou mete of these = pac just adduced.” ; is Onic, si Pa powerful Nerve Tonics, and No. 11s also a - 9Mptons erty or Defi is especially indicated in all cases where » No. 2 is mor iciency of Blood. In the absence of — e especially indicated. Sstimonials, which d is for sale by Grocers and Druggists generally. ! efy all imputation or questions, sent to any address. | ET __~ NIETOR, BUFFALO LITHIA SPRINGS, VIRGINIA h a ty are Prings are is i Oo le engl pen for guests from June 15th to f the Southern Railway. fom all di ir i éctions over the Danville Division 0 APAPAPYAYEPAPAPAPAP KY KR HKY ial 2 > 2 $ tal bd NOVEMBER, 1900 No. 5 THE on EDITORS A HN M. COULTER anp CHARLES R. BARNES, WIT H OTHER MEMBERS OF THE me STAFF. OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS FRITZ NOLL Cae Univ versity of Bonn VOLNEY - Se ROLAND nA§ WILLIAM panne gel Missour? Botan: H. MARSHALL WARD. University of t a EUGEN. ve URA fi University, Toby VEIT va sen CHICAGO, ILLINOIS. 9. Wblished by the Gniversity of ehicage Che Aniversity of Eicage Pree PYRI 1GHT 3906. BY THE UNIVERSITY © oF ii i HN a AMEE iat ah Botanical Gazette ly Journal Embracing all Departments of mp at oes Science » $4.00 Single Numbers, 40 Cents ption price must be paid in advance. No numbers are sent after a expiration f the time paid for. No reduction is made to dealers or agents. FOREIGN AGENTS: — Wa. WESLEY & SON, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin 18 Shillings. SW. 46, Schénebergerstr. 17a. 18 Marks Issued November 35, 1900 CONTENTS JRE OF THE ook koe WHICH CAUSES THE aig OF FORM ORPHIC GREEN A CONTRIBUTIONS FROM THE HU 1. Bapanaeay va I (WIrH PLA aid XVII AND xvi). Burton dward Lonisuen 289 ON LESSONIA (wirH PLATES XIx-xx1). Conway MacMillan - - 318 TAGUS. Il. C.D. Beadle - . - : - 5 ER Day (with portrait). John F. Cow ; i" NS ON THE Roor SysTEM OF CERTAIN Cacrac Carteon Be Preston PROPAGATION OF OPUNTIA. Cane: E. Pree ht oe URopsis. 7. D. A. Cockerell : ere Ss ‘ae ERATURE ss : 3 : 352 b TROPICAL NATURE. 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Chicago Press, Chica _ M/ntered at the Post Office at Chicago, Ill., as second-class mail matter.] Laird & Lee’s New Publicatio For Home, “hd ae and Birthday Gifts yh Sage eps vol Alt Sak 5 aa desi iceiee Duce ind h page a i i ec hi aiden ‘fair, t the ‘little darkey, the mo kits, pe dozens ea be chs this brilliant little drama. A be autitul holiday gift ete 3% cover in four colors. Twelve colors used through the book. Fam rapper. In a box. $1.25. “Among childrehs books, stands first.” Times. Fireside Battles. By ANNIE Ma ita wn. A delightful story for girls Te to life, full of : elnanelpy wit, and action. Exquisitely illustrated by J. © Lam ER, the eminent artist. ae on oF Luxe. 8vo. papeiee oe In a box. $1.25. c Will rapidly rank among classics.” Milwaukee Worll. Size, 11 X 9% inch ze The Heart of a Boy. Zdition de Luxe. From the 224th ener es a D800 AMICIS. i ogee 26 — etchings and 33 full-page, half-tone engravings. in steht langu Fine os Bas per. 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Prizes For the best memoir presented a prize of sixty dollars may be awarded ; if, however, the memoir be one of marked merit, the amount may be increased to one hundred dollars, at the discretion of the committee. F For the next best memoir, a price not exceeding fifty dollars may be : awarded. Satural Prizes will not be awarded unless the memoirs presented are of ade- 3 quate merit. : The competition for these prizes ts not restricted, but ts open to all. : Attention is especially called to the following points : . In all cases the memoirs are to be based on a considerable body of original and unpublished work, accompanied by a general review of the made directly in competition for the prize. al Each memoir must be accompanied by a sealed envelope enclosing the author's name Lega with a motto corresponding to one borne by the manuscript, and must be in Se of the Secretary on or before April 1 of the year for which the prize Is offered. Dike a0 FOR 1901: Monograph on any problem connected with or any group belonging * North American fauna or flora. eta 1892: (1) Nuclear fusions in plants. (2) The fate of specific areas of the as determined by local destruction. The reactions of organisms to solu- om the standpoint of the chemical theory of dissociation. Theos C. F. BACHELDER, Secretary. ON Soctety or NATURAL HIstTory, Boston, Mass., U.S. A. : of OOD paper and printing, North Ameri G printing on one side of ie can page only, larger tyP® | Plants, ee J os genera arranged and numbered if according to the Engler & and exten- are fea- Prantl sequence, sive use of synonyms, tures of this edition. SECOND EDITION tte Price $1.00 per COPY paper covers, or $1.25 bound in cloth. ad | a | =, HELLER “ : wha Walnut street Sample pages may Lanca ; STER, , U.S. A: upon application. be had - 2 aaa 3 Send 12 cents, U. S. stamps, for : f Wm. Wesley & Son’s New Botanical Catalogs} No. 127-128 of Bibliography History Biographies and Portraits } f f Herbals Early Botanical Science . THE Linnaeus Handbooks _ Dictionaries } f NATURAL 1 Microscopy Morphology aad Physiology | f HISTORY and f Botanical classification Nomenclature SCIENTIFIC ime iaile Botany Encyclopaedic works i | a a Cryptog: Phanerogams ; Natural gear eo of Plants (Floras) N f CIRCULAR ; Fossil Plants § Transactions of Scientific Societies | Periodicals Applied Botany Botanical Gardens J BOTANY { Agriculture and Horticulture to the end of the J8th century ; Upwards of 3000 works, Gardening Ornamental Plants Food Plants classified under 50 i Medical Botany "Tobacco Gums, Caoutchouc, elt f headings Fibres Forestry Agriculture ( | j 92 pages Diseases of Plants ‘ £ William Wesley & Son, Booksellers, f f 28 Essex Street, Strand, - - London, England. Be preomeine Botanische Zeitechag Die “Allgemeine botanische Litsch hrift” Yai ies vor allem Abhandlungen tuber scl ~~ phos ruppe: n, Diag gnosen kritischer Arten Formen und rar elo Schilderungen hse fai tos geographisch tnteressanter Gebiete, botanische Reiseberichte, Referate wber systemaltse ae wd To? Phanzengeographische Arbeiten, Berichte iiber die T. hitig heit einige Institute, Verel etc., und deren Schriften und Ka laloge, biographische Not he MT h tne besondere Sorgfalt wird gts den Re feraten tiber iessccatnsuere, bo botanise: sie oe Rear phen sugewendet, deren Ziel u. a. die Her ausgabe von Ex cuinagper mplett vorliegende itis ang 1899 wurde unter Mitwirkung von 43 Botan enthalt je ae sg oe a ii Pp 4 r rn he ce tet aber 7 Se » 31 Referate, Inhaltsangaben von 20 b Zeacheten, bre ee ereinen, ‘Anstalten etc., tiber 32 Tauschvereine und Exsiccaten aa und pan samtliche zur Kenntnis der Redaktion gelangende Personalnachric aller E Erdteil A Aligemel eine botanis ” m 15. jeden Monats Umschla versehen in min sos pene a be: Fpro. Quarta ri Mk. und yer ome Abonnenten portofrei unter Kreuzband zugesandt. 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Home Office: aaa N. - ME XXX | NUMBER 5 MPANICAL GAZETTE tb NOVEMBER; 19000 ; HE NATURE OF THE STIMULUS WHICH CAUSES es 1E CHANGE OF FORM IN POLYMORPHICGREEN GA. IBUTIONS FROM THE HULL BOTANICAL LABORATORY : (WITH PLATES XVII AND XVIII) BuRTON EDWARD LIVINGSTON. orabiga A.: Die anorganische Salze als Hilfsmittel zum Studium ‘der is ig chlorophyllhaltigen Organismen. Mélanges biol. St, Peters- onli tiicwe. ee » = DWaKy, a I. Zur Morphologie der Ulothricheen. Mélanges biol. 76. Ueber Palmell i Sti ium, Bot, Zeit. 3417 ¥, Fs ellenzustand bei Stigeoclonium. ot. Bed Para nezherches sur le développement et la classification de quelques a — Die Bedingungen der Fortpflanzung bei einigen Algen ae ae 289 290 BOTANICAL GAZETTE [ NOVEMBER Descriptive. A species of Stigeoclonium (perhaps a form of S.. tenue), found growing with Pleurococcus, etc., on moist bark at Ann Arbor, was chosen for experimentation. It shows two very distinct and easily distinguishable forms according to the culture medium in which it is grown. The normal form is that of Palmella. A culture of this form shows the surface of the fluid covered with : spherical cells 12 to 15m in diameter. The cells multiply by division in planes generally vertical and at right angles to each other, and the daughter cells separate more or less completely after division is accomplished (figs. 7, 6, 72, 27). The walls are quite thick and somewhat gelatinous on the exterior, though not by any means so markedly as in the form studied by Cienkow- sky (/.c., 2),and the protoplasm is quite coarsely granular. The chloroplast has the form of a hollow spherical shell with aa opening in one side, which has a diameter about equal ad the sphere’s radius, varying somewhat in different cells. Sometimes the opening is nearly a great circle of the sphere (fig: 4% i) This opening in the chloroplast is probably the “bright spat” described by Cienkowsky. There is always a pyrenoid present : in each cell, and sometimes two are found. This body has # : diameter of about 3m, and lies in a thickened region in ie | chloroplast. When the cell is about to divide the chloroplast separates into two parts, each part taking a half of the orig!” | pyrenoid, and a wall forms between the two portions (fig. 7 ¢)- : The plane of division always intersects the plane of the - e in the chloroplast. Then the two hemispherical cells: Decor more and more spherical, splitting the wall between its middle lamella, until finally, if nothing prevents, to lie as two separate spheres side by side (fig: shes “fg and often division proceeds more rapidly than this rounding - with : there results a group of four or more cells making cone : each other in plane surfaces (fig. 6). Often thi tinued without any parting till there results a spread out over the surface of the nutrient medium. 5 of the. too, the planes of division lie parallel to the surface 1900] CHANGE OF FORM IN GREEN ALGH 291 medium as well as perpendicular to it, and a parenchyma-like mass of polygonal cells is formed, projecting down into the fluid and up into the air (figs. 6, 2g). After division the chloroplast enlarges until it occupies about as much of the daughter cell asthe mother chloroplast did of the mother cell. The form just described will be termed throughout this paper the palmella form In the other form we have a very different mode of growth. The cells are not spherical, but cylindrical; they divide, except- ing at the origin of a branch, only by transverse planes, and show no tendency to break apart, but remain closely connected to form branching filaments (jigs. 2,8,18). This will be termed in the following discussion the filamentous form. The cells of the filaments are 4.5 to 10m in diameter and from two to four times as long. The longest filaments are 175 to 200p long and their branches extend in all directions. No true hairs have been Seen, but often the tips of the filaments are very narrow. The protoplasm is not granular, as in the palmella form, but usu- ‘lly contains two to seven refractive bodies smaller than the Pyrenoid, but looking much like it. They are probably of an eS tatare. They have a diameter of about 154, that of the Pyrenoid being, as in the other form, about 34. The chloroplast elongated to suit the cell and becomes somewhat trough- shaped ; lying along one side of the cylindrical wall it reaches eee vee half way around the cell and usually partially covers nd, sometimes both. Its opening is thus longer than in the eer. The pyrenoid lies somewhere in a thickened part - Of the chloroplast. Reproduction in both forms is accomplished by comparatively of ts “xual biciliate zoospores. These are the nice chlor, “bs. I have observed no conjugating ZOOspores. ; Splast of the mother cell divides into four to eight parts ee “unously parallel planes running nearly at right its ag of the opening (fig. 7). Each of these portions, _ : oe ot the protoplasm, becomes a zoospore. Afier division is, ae © °mplete and the zoospores are nearly formed, the ne 292 BOTANICAL GAZETTE [NOVEMBER chloroplasts often lose much of their bright green color and become pale. Palmelloid cells usually produce seven or eight zoospores; the number produced by a filament cell depends upon its size. The zoospores are ellipsoidal to spherical when discharged into the water by the rupture of the parent wall, and immediately begin to move about, swimming by means of two long cilia (fig. 3). Each contains a pyrenoid, and often several smaller granules; the chloroplast can be seen lying against the wall opposite the origin of the cilia and extending in its usual cup-shaped form somewhat more than half way around the cell. The dimensions of motile zoospores are from 3X6 to 6X9#. After swimming freely for several hours, the motion of the cilia becomes sluggish and finally ceases altogether; the zoospore comes to rest and assumes the spherical form. The diameter 's now not far from 64. The chloroplast then regains its original bright color, the cell enlarges, and, if conditions are favorable, may go over imperceptibly into the palmella form, The cell thus produced may dividé a number of times, as described above, and then produce more zoospores; or it may produce ps immediately upon the attainment of its full size, or even earlier) or it may become a resting cell and remain quiescent indefi- nitely.? However, a zoospore does not usually show this mode of growth. It generally elongates as it lies on the ue sometimes at the bottom) of the fluid, and becomes 4 cylin = with rounded ends. As it grows longer it bends so nag : = becomes somewhat crescent-shaped, semicircular, or even horse S shoe shaped, and soon divides into two cells by @ — . wall near its middle (figs. 8, 32). Thus a filament 1s “— - which soon branches. Its cells produce zoospores sooner 2 later, according to conditions. Methods. Pure cultures were first obtained by growing in _ . : % : : € tion in suspended drops; then the material was trans : *KLEBs, G.: Loc. cit. tssch de on PRINGSHEIM, N.: Ueber die Dauerschwarmer des Wassernetzes- M Berliner Akad, 1860, or rred 10 1900] CHANGE OF FORM 1N GREEN ALG# 293 small, loosely covered glass dishes - containing from 5° to 10° of solution. The cultures stood on glass shelves close against the panes of an east or west window, and were always shaded from the direct rays of the sun by a curtain of white muslin, which was left constantly in position during the high tempera- tures of the summer months. Examination of cultures was made from time to time by transferring the dish with its con- tents directly to the stage of the microscope and using an objective of medium low power. The culture media used were modifications of the well-known fluid of Knop,3 consisting of the following salts: calcium nitrate, four parts ; magnesium sulfate, one part; potassium nitrate, one part; di-potassium acid phosphate, one part; iron, a trace. Owing to the extreme weakness of the solutions to be used, it was deemed advisable to secure at the outset, as nearly as might be, the exact proportions of the constituent salts here given, and to this end the following method was adopted. By reference to tables of the physical properties of solutions,* the corresponding Specific gravity and gram-molecular strength of solutions of the first three of the above salts were found. By the use of these fata a stock solution of each of these compounds was made up, “ga contained a specific number of gram-molerwies ea ae Sa specific gravity bottle was used in these pn Surth we was always taken into account. For the in the list data could not be found; therefore it was ‘ae in solution from normal solutions of phosphoric acid and oY droxid. The physical constants for these ae . a. eae of this method for producing K, re pe Retite f * ‘umetric analysis and found to is ae t a . oe of a normal solution of this salt so made es throughout € 1.13207 at 15°C. Redistilled water was us all of the work. he stock solutions thus prepared were kept in flasks tightly with rubber; their specific gravity was taken from 8,G.: op. cit., p. 8 tE. g. ; ; bridge, . given by WHeTHaM, W.C.D., Solution and Electrolysis, 215. Cam- 294 BOTANICAL GAZETTE [ NOVEMBER time to time, and water was added as it was lost by evapora- tion. Knowing the gram-molecular, and hence the percentage, strength of these stock solutions, any strength of Knop’s solu- tion may readily be made up from them; and solutions so made up are much more accurate in their proportions than those made in the ordinary way; for in dealing with salts such as mag- nesium sulfate and calcium nitrate the amount of water con- tained in the compound as taken from the laboratory bottle is always an uncertain quantity. All troubles arising from erystal- line and amorphous, as well as deliquescing, salts may thus be readily avoided. The exact amount of ferric salts in my solt- tion was not determined, The transfers of material from an old culture to a new were made, after a pure culture was obtained, by means of a needle. This was first heated in a flame and then cooled by plunging in the new culture fluid. Bits of steel always scaled off in this operation, and these furnished sufficient iron for the plants, In making up culture media from the stock solutions . constituent salts, it is necessary to dilute as far as possible before bringing the K, HPO, and Ca(NO,), together. As was remarked by Klebs, this avoids, in a great measure, the sepat- ating out of quantities of CaHPO,. This method is not com- pletely satisfactory, however, and it would be better 5° to | modify the proportions as to avoid the white precipitate entirely: I have planned to do this in future work. A solution made up of the quantities of the salts given by Knop dissolved ar parts of water is, of course, a 7 per cent. solution (since — iron is of such small amount it may be disregarded). aoe further diluted to 1 per cent., 1.5 per cent., and 2 pe sacs growing the stock material. : of the Investigation. la At the outset it was found that if material in the pale! form were transferred trom a 1 per cent., 1.5 pet crs i ae cent. solution where it had been growing for some me solution of less than 0.5 per cent., the plants responded : 1900] CHANGE OF FORM IN GREEN ALG& : 295 change in solution strength, and sent out long filaments from the original palmella masses ( figs. 7, 75; 16). Also the palmella cells produced numerous zoospores. These germinated, as pre- viously described, to produce, not the parent form, but the widely different filamentous one. Conversely, it was found that if filaments, produced as above, and growing healthily in a 0.2 per cent. or 0.25 per cent. nutrient solution, were changed to one of I per cent., 1.5 per cent., or 2 per cent. strength, the tesponse was scarcely less well marked than in the other case, the cells of the filaments soon became spherical, and changed to the palmella form by dividing in both longitudinal and transverse planes (jigs. 4, 5). In the strong solutions few or no zoospores were produced. Where a few were produced they changed, either directly or after the first or second division, to the palmella With these two facts in view the attempt was made to deter- mine where, in this change of solution strength, lies the neces- sary stimulus for the production of the corresponding change of fom. A stimulus arising from a change of solution strength (always retaining the same salts in the same proportion) may be of any one of three different natures. (1) It may be of a chemical nature : z. @., it may be due to a change in the amount of salts with which the organism is supplied. This implies three possibilities : (2) the plant may be affected by the increase jaa in all the inorganic salts. (4) Again when such aia is made it may be the change in amount of a single salt, rad Pedetum nitrate, to which the plant responds. There are Ur possibilities under this head, one for each of the four salts *¢. (c) Again, it may be possible that neither of the pre- Cedi me f the to Ry Suppositions is true, but that the response 1S due, not | k the changes together, nor to any one alone, but to @ come. aah or three of them. For example, it pers — i. € amount of potassium phosphate together wit! Le — €xce ti Tutrate, which acts as a stimulus. We know 7 ._ bit in very weak solutions, these two salts react chem- Y "pon each other, so that it is at least conceivable that am 296 BOTANICAL GAZETTE [NOVEMBER increase or decrease in both of them together might influence the plant differently from a corresponding change in only one. Under this head there are no less than ten possibilities, six taking the salts by twos, and four by threes. (2) Further, the stimulus may be of a physical nature and due, not to the change of salts at all, but to a change of osmotic pressure upon the living cell. This osmotic change may be effective in several ways. (a) It may be that the response is due to a change in the osmotic pressure of the solution in general. When the saline constituents of the solution are increased or decreased as 4 whole there is a corresponding change in the so-called osmotic strength. (4) However, it may be, as suggested by Copeland's recent work,’ that the plant is more influenced by osmotic pres- sure when this is derived from one salt than by the same pressure _ derived from another. And if this be the case, then this influ- ence is as complicated a one as that of the change in the com ditions enumerated under (1} above. (3) Finally, the response in the plant may be due both to the stimuli from chemical com- position and to those from osmotic pressure, a combination of (1) and (2) above. My experiments were devised to determine primarily whee the stimulus is a chemical or a physical one. For this it § necessary to have solutions in which the relative and absolute amounts of saline constituents can be varied without changing the osmotic pressure of the salt content as a whole, and seed must be brought about, as far as may be, without the saclay tion of any new conditions. It is necessary first to know Fe Osmotic pressure of the complete Knop’s solution. Now P : is € ual, pressure of any weak solution of several constituents 6 2 . essures 0! 2 , we salts the constituent salts, as these pressures would exist if eee were ; i lution whose V Separately put into a simple so makes it # equaled that of the complex one. This principle simple matter to calculate the pressure of complex Gad. 24: 39% SCoPELAND, E. B.: The relation of nutrient salts to turgor. BOT 76 1900] CHANGE OF FORM IN GREEN ALGAE 297 non-electrolytes. But the pressure of a simple solution of an electrolyte is not so easily calculated. On account of dissoci- ation, such pressure is much greater than that of a solution ofa non-electrolyte of the same gram-molecular concentration. It was this fact with which DeVries® was dealing when he derived his so-called isotonic coefficients. But since his range of concen- tration was very limited, he was unable to get at the truth of the matter; and the result is, that, though his coefficients may be, and doubtless are, true for a certain strength of solution, yet, Since dissociation itself varies with the concentration, they are fot true in general. However, in weak solutions dissociation becomes nearly complete, and if we assume that it is complete Weshall probably be able to approach the truth much more nearly than by following the method of DeVries. My solutions are all very weak, and I have no doubt that my approximations by this method are,as a rule, quite nearly true. By assuming complete dissociation, the calculated osmotic pressure for any solution becomes that of an equimolecular solution of a non-electrolyte, multiplied by the number of ions derived from a molecule of me salt under consideration. This makes the calculation very ‘imple. Taking the pressure of one gram-molecule per liter of oo undissociated substance as a unit, W, table I gives the oo of the complete solution which was used a ee . his is practically the same as a so-called 2 per cent. : at as usually made up: In the pressure Yee Method - 4 assuming ionization complete; & by ee ae which om Z Bives: the actual pressures for a sa ee “ee : dissociation could be found. The geo ts te 8 6 lies almost wholly the Ca(NOs) a dia Su: € a greater error in using DeVries coefficien a spe complete dissociation. Column gives the num zi sce tived from a molecule, 7 is the ratio between the eS : : Pressure and that of an equimolecular solution 9 “DeVairs, H Uco: Do Jahrbiicher fiir wiss. Bot. 14: 427. 1884. N PERS and HAMBURGER: Zeit. {. Physikal. Chemie 6: 319- 1890 298 BOTANICAL GAZETTE [NOVEMBER non-electrolyte. MV at o°C., is 1686 of mercury, or 22.18 atmospheres. TABLE I. Partial pressures in terms of V Salt ae $ Ps P . Ca(NO,), - - .07143 3 .21429 .28571 .175004 [2.45 KNO,- - - 02857 2 05714 08571 055712 [1-9 MgSO, - - - .02381 2 .04762 04762 035712. [Ee MghtPO, + - .01664 4 06655 .06736 | “ing uaue ee Total pressure - - - - - 38559 . 48641 Drea This 2 per cent. strength is the most concentrated solution | have used. Anything stronger kills the alga almost immedi- ately. As we decrease the concentration below this standard 2 per cent. strength the dissociation becomes more and nee complete. Thus as the concentration approaches zero as a limit the value of i approaches that of z. The value of ¢ for the dif- ferent concentrations used, as far as they can be calculated from the physical data at hand, are given in table II. Values for : H,HPO, are inserted for comparison, in place of those for y K,HPO,. Table II also shows the total pressures of the sac : ous percentages in terms of V. The pressures in column gor - be used throughout this paper, but instead of 0.3856 N 1 a write 3856 VX 10+, etc. ce TABLE IL. ed T. faces § a - OSE N. et DY Der cent,’’ Values of 7 calculated from conductivity shee eee ee Pee a HPO. % a! é Ca(NO,),| KNO, MgSO, | KsHPOs | a 2. ; 38559 | .486405| 2 2.45 1.9 1.50 "heme -289193 | .364804 | 1.5 2.5 (?) as 1.56 3.02 192795 | .243203 I 2.55 ks ae ce -096398 | .121601 a 2.66 £98 os eo -048199 | .060801 ae a vei 1.68 ae oe totes .024320 S 2.9 BA re Pie +00064 -013160 .0 2 09 7 The meaning of a and 4 is the same as in table I. 1900] CHANGE OF FORM IN GREEN ALG# 299 Having shown thus the partial and total pressures of the nor- mal solution, I shall now describe the modifications of this solu- tion which I have used. Four solutions were made up according to the formula given above, excepting that each was deficient in one of the four constituent salts. All of one salt, it was thought, _ should not be omitted, since then, as in higher plants, patho- + logical responses would probably ensue. In these solutions the deficient salt was reduced to one tenth its normal quantity. The decrease in osmotic pressure, caused in each instance by this diminution of one salt, was calculated by the two methods just spoken of, and of each of the three other salts a sufficiency si added to increase the pressure by an amount equal to one third of the calculated decrease. Two full sets of these solu- tions were made up and used, one by assuming dissociation com- plete, the other by DeVries’ method. These two solutions were identical in their effects upon the plant. Thus finally we have four solutions in each of which a deficiency of one salt is obtained, and the osmotic pressure of the solution is kept up to formal by an increase in the amounts of the other three salts. 4 amounts are not equally increased by weight, but accord- . om pressure which they produce when in solution. I INo . these solutions as follows: A, delicies in Ca a ; B, deficient in KNO,; C, deficient in MgSO, ; D, given b 2 K,HPO,. K will denote the normal solution 3 | a nop. The pressure will be expressed in terms of ad part of my work was done in a ae bike : upon the plants of the four solutions just describe is thy — one of Knop. They were all diluted, gee ae me to the various strengths given in table = : a 4. ‘ tests were made with solutions of ie). pe 2 simple soy salts with which we have been dealing. ne Pressure as .. were made up so as to have the same — . oar € complex ones just described. They will be des as follows: E, a solution of Ca (NO,) 2: F of KNOs; Wit Q inated of : | | Made 9 oe H, of K,HPOQ,. Calculations for these were nly by De Vries’ coefficients. rx, 300 BOTANICAL GAZETTE [NOVEMBER In all, 278 cultures were made and record kept of each for five to fifty days. They were made at all seasons of the year _and in two different years separated by a period of nine months, during which the material was kept alive and changes made from time to time but no records kept. It will be seen that there are two possibly very different questions to be answered in the prob- lem before us. First, what are the conditions that bring abouta - 4 change from'the palmella form to the filamentous form; second, what are the conditions that bring about the reverse change? bs shall divide the experimental data which are to follow into two groups corresponding to these two questions. Details of typical responses are given in the explanation of plates. A. RESPONSES OF THE PALMELLA FORM, In table III are given the results of fifty-five transfers of palmelloid cells into the different solutions. For convenience, all cultures having the same osmotic pressure are brougiit together. In the left hand column the letters denote the chem ical content of the fluid, as just described. The number of eae tures made is given in the second column. In the third 18 recorded the number of cultures in which the cells multiplied but remained of the palmella form after twenty to twenty-five ‘ days. Columns four and five (Filaments) give the eee” a which filamentous branches were produced from the orig! : masses in five to twenty-five days. Those in column four 2 duced few filaments, but continued mostly as palmella ; pee o column five produced many filaments. In columns six and ghee (Zoospores) are the numbers which produced few or Fe Spores within fifteen to twenty days. Column eight oo” ts those which produced no zoospores for twenty-five ye pee The last two columns record cultures on which definite ner | tians are lacking, the first with regard to the proces ee at spores, the second with regard to the production of flea the edges. | CHANGE OF FORM IN GREEN ALGH : 301 TABLE HE E Pressure 96 V X 104 or 16.19 cm. Hg. Pressure 1928 V X 10 4 or 323.71 cm. Hg. a Filam’ts Zoospores Filam’ts Zoospores ae er wean wa GRE NE §\8/4 Sileil¢i#te = 5 = ial > co) ral ret 5 = b> b 5 eizleleleiele|2(2i4il 2 |elaleleieielelg Sleigislselsizlalzell 3 |sle&lelslslelaia K 12} 1 2 10 ie os Pee 2 > ae " 5 5) K 15 21} a4478 Bi5|-.}2/3| 3/2 A |4|1)3 i Patt... 515 B | 4 pee 4 Disi2|..13| 5 C Lal.) 2 bee oe : Bisi..| 2 31711 3 Def 41 2 248s 13 a : ef ef es | 2 a Eee aie a 0 ei 24 Pps) 2 Pressure 2892 V X 10 or 487.59 cm. Hg. z H = . I I 2 mt | fore : P K 343 25 3 t= Re Se fe A X10 8 32.37 cm. Hg. : 5 $122 ewe : petit i 1 Cc : “ : 1 ; a I I a S15 y it DS Be es ee ; 2 E vais Nae Rae Ih das Sc. ed be Brae Bee F 240 BPG fa es ese Se PS ee fae G 212 ae ee ee rot Hoe 2 eee a Pressure 482 V X 10 or 81.27 cm. Hg. Li Ne K | 2 | Pressure 3856 V X 10 or 647.42cm. Hg. terete. | 2 a eens eae a ee a ee | |*)- || P, Ke} 2:7:2 Hobe Ts A ip 964 VX 10 or 161.86 cm. Hg. A} 2i2 I oy Lee kK aT Gre B 2 2 Fa es we Pio 1% Clete bass a1 5 ere th ap ee ve Eas B eA ES a c mee ii 3) 5)..]. a Bees itis ce ae eee F 2 I I i : Gia : I = ve a I here ee be General Remarks on Table III. of aaa lived indefinitely in solutions K, A, B, ©, and D, 2G: a oo” and also in the lower concentrations of B, single Salts - But the higher pressures, when caused by 4 pressure , usually resulted in more or less speedy death. sexe ‘the cell é ies N X to, it was quite a common thing to set = ‘dead within a week. Sometimes they were perceptibly 302 BOTANICAL GAZETTE [ NOVEMBER plasmolyzed before death ensued, sometimes not. Of the four simple solutions one appeared to affect the plant as much as another. The table shows the general result of the experiments almost graphically. In the higher concentrations the larger numbers appear mainly in the middle of the series, in the columns headed Many, while the third and eighth columns are nearly vacant. Passing to higher concentrations this gradually changes, until, in the highest, the reverse is true. The following conclu- sions may be drawn from these data. 1. Nature of the stimulus—The effect of a solution of any given osmotic pressure upon the plant is both qualitatively and quantitatively the same, no matter what the chemical nature of the solution may be. If this effect were due to change in the amount of any one salt, then this fact would have appeared in the course of the experimentation, for my various solutions were devised to test this point. So we have eliminated, as far as the palmella form is concerned, the possibilities designated under I, a,b,c, and 2,6, page 295. The stimulus cannot be of a chemical nature, nor can it be physical and depend more upon tag aah ence of one salt than upon another. Also, we have eliminat 3 from among the possible primary stimuli. Consider : facts shown, it is impossible that the external stimulus S208 : iment of certain be either physical, with the necessary accompan a chemical factors, or yet chemical, with the necessary acreye 2 ment of certain physical ones. It is, however, nae : but probable that the external and primary physical “43 may cause an internal and secondary chemical one. a a lies beyond the scope of the present research. Thus ene 5 driven to accept 2, a as the truth in the matter, and to that the response of the organism is determine and never, as far as my experiments have gone stimulus. The effect of osmotic pressure upon t upod 2 form is a double one. I shall consider, first, the eich vegetative growth, and second, that upon reproduct! 2. Response in vegetative growth.—In my leat ae solution (pressure 96 V X 10") the mode of cell divisiOv” on. . oncentratee 1900] CHANGE OF FORM IN GREEN ALGH 393 invariably changed. In forty-six cultures only three are excep- tions to this. The cells at the edge of the original mass elongate and become ellipsoidal or even cylindrical, and when division occurs it is by means of a wall lying transverse to the long axis of the cell (figs. 7, 75, 76). Thus filaments grow out around the periphery of the original group. The interior cells do not change their form and do not divide further; at length they form the center of a mass of long radiating filaments. Branching of a cell. occurs by means of a papilla-like lateral outgrowth, which Scut off by a wall transverse to its long axis. The walls between the filament cells thus formed do not split ; thus there is no bulging of the end walls and no separating of the cells. The older cells of a filament (for division occurs only at the tip) bulge at the sides and become barrel-shaped, but the end walls hold them fast together and prevent them forming true spheres (ag. 25). It is apparently this coherence of the walls and the ‘nentation of the walls with reference to the form of the cell Which constitutes the filamentous habit. ‘s the osmotic pressure of the fluid is increased there is a “ntinual lessening tendency on the part of the plant to produce filaments, and there arise fewer and fewer filaments at the edge %: the cluster, Also these filaments tend to go back, by a round- Si breaking apart of their cells, into the original form. -The see ee of the filaments do this, while the younger ones corsant li, © advance by cylindrical cells (figs. 4, 51 10 25: a 34). 2 y most concentrated solution (pressure 3856 WV x 10 +) there ome still five cultures out of ten wherein filaments were formed, Fi s but one of these the filaments were only two or — a, aaah Back of these cells a thick strand of palmelloi : od ed the path of the advancing filament tip (Jigs. am 30). »~Hange of the cylindrical cell to a spherical -— aed aay * it is, by a corresponding change in the direction of in filam 8, will be discussed when we consider the eeee — For the present it is enough to note - sme Mri pressure the tendency to produce peripheral fila 2 8, both in the number and in the length they “_ 304 BOTANICAL GAZETTE — [NOVEMBER before reverting to the palmella form. In my weakest solution they often attained a length of twenty-five to thirty cells (but the older cells here are destroyed in the act of producing zoospores, which will presently be discussed), while in my strongest solution the few filaments produced seldom attained a length of over two cells. The tips of filaments are quite sharply pointed in weak solutions, but blunt in strong ones. A maximum pressure beyond which filament growth cannot take place lies somewhere between 1928 x 1oand 2892 X_ 10*. No limit was found beyond which no filaments at all could be formed. 3. Response in reproduction.—The most marked response of this form to the change in osmotic pressure conditions is the sudden production of enormous numbers of zoospores. In some of my cultures almost every original cell has thus emptied out its con tents within ten or twelve days of the date of transference. In — general, within a fortnight and often within five days these very weak solutions become literally swarming with zoospores, and their production continues as long as the pressure remains lof: Zoospores come both from original spherical or parenchyma-like cells and from the older cells of filaments radiating from the palmella mass. This method of producing zoospores bee often a been resorted to by various writers, and Klebs especially hss : found it successful in a great variety of forms. But, as ge I know, previous writers have not tried to determine the re Nature of this weak solution stimulus. As would be expected from the response i growth, when zoospores come to rest in these we : and begin to enlarge, they do so by elongating into 4 ches. which soon divides transversely and later produces eee { figs. 8, 14, 78, 32). Thus the response to low osmotic a 7 of a zoospore which has come to rest is the same 4° a young cell of the palmella form. Zoospores often grow 4 ae ends, however, which I have not observed in case of oe thus The two cells formed by the first division of the 2008 sometimes become a base from which reach out two long bs | ing filaments (figs. 2, 78). n vegetative a) CHANGE OF FORM IN GREEN ALGE 305 In solutions of greater pressure zoospores are produced more and more tardily and in fewer and fewer numbers. The limit to the healthy production of zoospores lies at a pressure of about 964 VX 10%. Their production in a higher pressure is exceptional and | have never seen them produced at all in my strongest solution. With a pressure of 964.V x 10% all zoospores germinate very Soon in the manner already described ; some of them go directly into the palmella form while most of them germinate to produce flaments of two to five cells, and then these pass into the pal- _mella condition by rounding and breaking up (figs. 17, 79): Thus it often ¢omes about that cultures with this pressure show both forms growing together, short young filaments and free round cells as well as parenchyma-like masses. With Pressures above the observed limit the spores, when they are produced, usually grow directly into the palmella form, and : then continue to grow slowly, or go into the resting stage. I have Ziven little attention to the fate of these resting spores. They retain their green color and remain indefinitely at the bottom of the culture dish. Klebs has described these and a that they may be made to germinate after complete drying out. B. RESPONSES OF THE FILAMENTOUS FORM. : Experimental data from 169 transfers of filaments into the : Solutions are presented in table IV, which is constructed Mater} a plan as table III. Whether or not the original oa changed into the palmella form is shown in the third, a ie columns. Cultures in which all or nearly all VS are r oe * SS alae the other form within a period of thirty Dut not eae in the third column; those in which some, » Went over, and long filaments persisted after twenty are indicated in the fourth column. The fifth © Fesults T none aS to the production of zoospores, whether many, few, / © tabulated under those respective heads. Where 306 BOTANICAL GAZETTE [Novemner zoospores were produced, but many or all of them did not germinate, the numbers are placed in the column headed Ungerm. TABLE IV. Pressure 96 V X 104 or 16.19 cm. Hg. Pressure 964 V X 10% or 161.86 cm. Hg. Palmella Spores Palmella Spores eeu trae Coe —_—| » | —— Blzi_lselelFlsl|*|elsil 3 | 2l. alel es ; H/Ol<|[Hlel/zAlel[ol42lal] & |O|< |e |e) a] mela) K | 6 STG K 17/3/14) & 6 G16). A 16/1/41) Oe Bes oe a es B. | 614|..) 33 Oe Ce ON es ee es I C |} .6:) 3.03 }s. pa ee D | 6 erat 514s 1|| D|6|2)3]1) ees nee E Ly Peet 5 12 ee be oa Pa 4| 2} 2) 3¢8 om! Et ee I ceferl Ort “Eve es tis os ee is ay he ae H|1 P45 Pressure 193 V X 10 * or 32.37 cm. Hg. Pressure 1928 V Se 10 ¢ or ets ws Ae Be ae Oe a Be K | 6131 2)34 3 Cae oe ae ee A 18134 £12 ; oe --|3| 3 3 B pets 2 lB 3 | 3 3 C |5|5 |=) 4a. ee D 13]: a) 3 . D {51316 Sy oe e132 2102 EPs} ga Pressure 482 V x 10 + or 81.27 cm. Hg. Pressure pesos NX 304 OF 487-508% = BAP 3 .. 3 Kobi 1. £133 hs 3 A l3)a4 : f By3| 3 3 3 Bi313 ae ee Be ee ee 3 C13) 3p ea ne fo eS RS eg ea 3 D132) 2 : E | 3 ately 2 BE 16i44= Sa 2 i eee I : G fa : jo ee Be General Remarks on Table IV. The only solutions in which death regularly me more concentrated ones made from single salts; ? eB i id But in almost every case where the original filaments © evinced a great tendency to change into the ara were the 1900] CHANGE OF FORM IN GREEN ALG 307 before death ensued. Often they did not die until they had actually taken the palmella form. Where a definite response was not observed before death, the cultures are recorded in the column headed Died. The graphic arrangement of the numbers in this table is even more striking than that in table III. The : conclusions to be drawn from the above set of results will be 4 given in the same order as were those derived from the experi- tents upon the palmella form. \. Nature of the stimulus.—The stimulus with which we have to deal is always of an osmotic nature. Indeed, this generaliza- tion would have been expected from the nature of the stimulus affecting palmelloid cells, and from the data which we are now considering the same facts can be cited in its support. 2. Response in vegetative growth.— Filaments placed in the Weakest solution used (pressure 96 V X 107) continued indef- mitely to grow without change in form. No going over to Palmella was observed until time enough had elapsed for great *oncentration of the solution through evaporation. The rule is, if filaments are left in a dish without renewal of fluid, they change to the other form at the end of a period of thirty to fifty ay By this time the culture is often nearly dry. Also, by Addition of water from time to time filaments may be kept sowing in a dish as long as one chooses. So there is little room °F question as to what the nature of the stimulus producing this ey response really is. With higher osmotic pressures “pas = the alga to continue as filaments rapidly eee: ay dication of such an effect being the blunting of the : mal cells, A limiting pressure for free filament growth lies “ge somewhere in the region between the pressure of | 10 ont fo and 482 V x 10+. With a pressure of 964 = “id i. out of forty cultures showed good hielo lost aly a et thirty days, while many of these oe a. ie. of them within that time. Nota _ e wae ‘typical gi cultures having a pressure of 2892 NX 10% 5 ang Only f aments. at the end of such a period, and there _* Strwhich showed filaments at all. As in the other form, x ie a if Ate g 308 BOTANICAL GAZETTE [ Novemper an absolute limit to the production of filaments was not found. A comparison of the limit found here with that found in the case of palmella material will be made later. Filaments floating on a strong solution change to the palmella form much more slowly than those at the bottom of the dish. The morphological details of the response just discussed are as follows. If a filament be placed in a strong solution the oldest cells are the first to be visibly affected. They continue rounding up beyond the barrel-shaped stage and become nearly or quite spherical. This process is accompanied necessarily by a splitting of the common walls from their margins inward, until, when the cells have attained the spherical form, they are practi- cally free from one another (figs. 4,5). They may not break entirely apart, however, but may remain clinging together in loose strands. The limitation upon the direction of new walls (cf. discussion of the responses of palmella form, /. 300) is nie apparently removed, and the cells proceed to divide in all direc- tions. The new cells may separate entirely (jig. ro), may remain loosely attached to one another (jig. 34), or may form an irregular parenchyma-like colony (jigs. 6, rr). Inthe latter case the rounding is of course incomplete, but such cells are always nearly isodiametric. Cell division goes on much more slowly in the palmella form than in the filamentous. y : ; : e hee. 3. Response in reproduction.—The response is the sam a as in the other forms and is equally marked. The somewhat above 964 NV x 10%. The limit to their into long filaments is below 482 V X 10%. duced here in the way already described. filament, even in a weak solution, is limited by oldest cells are continually becoming zoosporang!@ - losing their contents (fig. 16). d_ thus General considerations. - ertaken to ansW je The question which this research was und has been answered very clearly as far as the a 1900] CHANGE OF FORM IN GREEN ALG& 309 stimulus to which the alga responds by a change of form is a change in the osmotic pressure of the surrounding fluid. This must be accompanied by a corresponding change in the relation of water to the protoplasm. Whether or not extraction of water brought about by solutions of non-electrolytes or by evaporation from a moist substratum will be accompanied by the same response as that brought about by a solution of electrolytes, my experimentation has not yet gone far enough to show. It is worthy of remark, however, that when found wild this alga was in the palmella form, and was growing in air on moist bark. Loeb® has recently brought about artificial parthenogenesis, both in the eggs of echinoderms and annelids by placing them in a solution which would extract water from them osmotically. The fesult was the same whether electrolytes or non-electrolytes Were used. Thus in these eggs the phenomena of segmentation and growth are profoundly influenced by osmotic pressure. Whether or not his results have any connection with those given in the present paper is an open question. The recent work upon Various Stains and forage plants by Buffum and Slosson? fur- hishes data which may have a somewhat closer connection with Nyown. Absorption by seeds, their germination, and the growth of the plant were all greatly retarded in a strong solution and *celerated ina weak one. And this was true without regard to : €chemical nature of the dissolved substance ; a number of -Clectrolytes were used and also non-electrolytes. On the other of Sal €xperiments have not been at all in accord with those Peland (doc, cit.) when he found, with peas, maize, and F atten J.: Further experiments on artificial parthenogenesis and the nature of as fertilization, Amer. Jour. Physiol. 4: 178. 1900. 170. 1900, * Attificial parthenogenesis in annelids (Chztopterus). Science N.S. 12: 9 . = hie “) SLosson, E. E., and Burrum, B. C.: Alkali studies I, Bulletin 39 Wyo : ( Exp. Sta, 189 : 2 * ‘ Bap uteM B. Cu: Alkali studies III, Ninth Ann. Report, Wyoming Agri. XP. Sta, 1899, . (3) SLosson, a) By E.E.: Alkali studies IV, same. 1899. ee and SLosson, E. E.: Alkali studies V, Tenth Ann. Report, Joming Agric, Exp. Sta. 1900 310 BOTANICAL GAZETTE . [NOVEMBER buckwheat, that K-ions are osmotically more potent than any other ions in his nutrient solutions. Some erratic cultures in my series seem to support his statements, but no generalization can be made from them. Of interest, also, in this connection are the recent researches of Yasuda.*? Working with infusoria, he finds that these organisms are able to adjust themselves to solu- tions of quite high concentration, and that, in general, the limit to their power of adjustment seems to be osmotically about the same, no matter what salts are used. In other words, the limit is apparently one of osmotic pressure. It is probable that many of the so-called chemical or nutritive effects of dissolved sub- stances upon the plant organism may turn out, upon further inves tigation, to be wholly or in part osmotic. In a weak solution vegetative growth is very much more rapid than in a strong one This may be due to the fact that in a strong solution the water content of the protoplasm is reduced in amount below the limit for optimum lability. When the plant grows fastest and best, it is in the filamentous form. In weak solution, where activity seems to be ata maximum, the ions of the electrolytes, which are essential for metabolism, are not plentiful. This may suggest how the cylindrical form of cell with its increased surface™* may be advantageous. A : “ rate, we may be sure that the greater surface of the cylinder puts the plant into better condition for exchange of mate with its surrounding medium. On the other hand, the ca concentrated solution not only withholds water from the a but presents a demand upon them for water. The cell me . ‘“ s the this in part by offering as small a surface as possible yee solution. In this case, although the requisite ions may be p sf water in the” ent, and even in the right number, the scarcity of fount «picket einiger Z * YASUDA, ATSUSHI: Studien iiber die Anpassungsfahigkeit reread 1900. an concentrirte Lésungen. Jour. Coll. Sci. Imp. Univ. Tokyo 13 101-14 so sae *In a cylinder, the lateral surface is greater than that of a yee 2.727 volume, as long as the ratio of the length to the diameter equals . ait js often 4 In typical filament cells of this alga the ratio of the diameters 1S 3, gre filament cell and even greater. It is seldom less than 2.8. Thus it is shown _ alls alone, tha! offers more surface to the surrounding medium through its lateral w does the palmella cell of equal volume. 1900] CHANGE OF FORM IN GREEN ALG 311 protoplasm may so decrease the lability that rapid growth is impossible. What may be the mechanics of this rounding up of cylin- drical cells when placed ina concentrated solution is one of the most important problems suggested by. the present research. The fact that the dead cellulose membrane is almost entirely reshaped during this process, without being dissolved, renders it probable that the change in form is directly caused by some turgor change within the cell. In arounding cell the membrane moves and changes its form and, since it is entirely inert, the source of this motion must be either in the activity of the proto- plasmic body itself, or it must be in the turgor pressure of the mass of liquid within. But since protoplasm and cellulose wall ra be parted so readily during plasmolysis, the first alternative iswell-nigh untenable. If the wall be forced into the spherical shape by a change in the pressure from within, this must be brought about by a change in the mass of the contained liquids. Now, this slight change in mass which might produce a change in the turgor of the cell is most probably due to an alteration in the amount of cell sap within the vacuole. When the surround- ™g medium suffers change in concentration, a change in the Yolume of the vacuole may come about through the protoplasmic “, either secreting liquid or acting merely as a semipermeable Membrane, Fa ; i ‘t a i When filaments are placed in a concentrated solution their behavior Suggests at once partial plasmolysis. Water may be “tracted, the turgor pressure on the walls may be decreased, - by the forces of surface tension and cohesion the proto- : ag May tend to round itself up into a sphere. If this be set . oo a explanation of the lateral bulging which accompa — longitudinal shrinking of the cellulose envelope. If wei Protoplasm tended to assume a spherical form within the cylin- _ | Mall, the pressure upon this would be decreased first at the pe At the same time it would be relatively increased sail . ral walls near their middle. Thus would come — Sing of the lateral walls outward, and hence a shortening f 312 BOTANICAL GAZETTE [NOVEMBER the cell and a drawing of the end walls towards each other, But the internal pressure is to be counted as almost nothing at the angles, while it is still considerable in the middle of each end wall. So the margins of the end walls would approach the mid- dle of the cell more rapidly than do their central portions, and splitting of the common membrane of two adjacent cells would — necessarily ensue. Several facts were observed in the cultures ~ which seem to support some such hypothesis as the one just a stated. I have placed filaments in a solution where they were completely plasmolyzed and killed without any change in form. In solutions a little less concentrated they are not plasmolyzed but round up rapidly and soon die, often in the palmella condi- tion. With a still lower pressure the filament cells round up more slowly and live. Another fact suggesting this idea is that floating filaments can resist a stronger solution, and can resist it longer, than sunken ones. The former are to some extent i contact with the air, and thus present less surface than the latter : to the liquid. Still another observation bearing upo? ae hypothesis of partial plasmolysis is that cylindrical cells are : the only ones which are able to change their form after ses ' have become mature. A spherical cell must remain so till : divides, even if it be in a solution of very low pressure. Only a two other observations with which I am acquainted bear Bie this question: Yasuda (Joc. cit.) says infusoria in pee solutions tend to approach the spherical form, and Rlets a cit.) notes that the form of Stigeoclonium tenue with which he wa working had a tendency to produce round cells ina oe solution. The very marked response in the orientation ee planes in segmentation may be traceable to the change “ cells of the cell, orit may not. The observed fact is that vegetative je under low osmotic pressure divide only across their peer (across the axis of a lateral outgrowth in the case of a ear while under the influence of high osmotic presi ss ulus divide in all directions. It is remarkable, too, that hs ee determining the position of walls in vegetative wie brings about the extreme segmentation which occurs 1900] CHANGE OF FORM IN GREEN ALG 313 mwosporangium. Perhaps this fact is attributable to the increased general activity in weak solutions. It has no relation to the form of the cell, since zoospores are produced from both spherical and cylindrical cells, as well as from those of inter- mediate shape. My approximations to the pressure limits for the different fesponses are given in table V. The numbers are all coefficients of VX 10-4, TABLE V. Maximum | Maximum | Minimum | Minimum | Maximum | Maximum 8 limit to free limit to limit to free limit to limit to free limit to any SS ee production production | production | production | production ing 9-1 ge : of filaments of any long | of palmella | of any pal- o to} filaments orm melloid cells) zoospores | zoospores | zoospores PPM ee Maxim limit to — Tom ig s ae - ic : surface the diametet ing from 6-30 in diameter. Towards the suri io of the cells lying in the radius of the cylinder 1s : there is a general superficial cambial area extending af | 1900] OBSERVATIONS ON LESSONIA 325 boldfast and continuous over its tip immediately under the epi- dermis, The epidermal cells and the layer underneath are the only ones that take a protoplasmic stain with avidity, but the layers within these, to the number of five or six, contain numer- ous sharply staining chlorophyll bodies. It is apparent, there- fore, that the growth in thickness of the holdfast ts essentially 4 superficial, and its whole tissue is produced from the cambial aeas somewhat as is phelloderm from the cork cambium of higher plants. The epidermal cells proper are much the : smallest of all those that make up the holdfast area, averaging from $-lov in height. There is no differentiated medullary area in the organ. The longitudinal section of a haptere shows the customary Structure, and serves to make clear that the growth in length takes place in the same way as does the growth in thickness. Toward the center of the section the cells are much elongated, funning in some instances as high as one tenth of a millimeter mlength. This length diminishes toward the tip of the organ, and directly under the tip a vertical section does not differ “sentially in appearance from a transection. The dichotomous branching of the holdfast is due, as in Nereocystis and generally ® Laminariaceze, to the appearance of adjacent circular fields . cambium which undergo division more rapidly than the Mervening and surrounding areas, so that the centers of these a protruded as the tips of the two new hapteri¢ ches, — eae hapteres which have become matted together ~ e e al holdfast mass, the walls are somewhat thickened in 4 gg Many of the cells are collapsed and i ss engulfed ah whole hapteric branches seem 1 yee eg n the general tissue, reminding one of the me of the Otder, .. each other that often occur in other eS ‘aie thickenin 0€s not seem probable that there 1s ee ay Stipe of 4 of holdfasts or hapteres, such as takes piac embiing - these of 4 €ssonia, and produces the curious ings res N €xogenous tree. 326 BOTANICAL GAZETTE [Novewsen The stipe—Very young stipe areas in Lessonia are much flat- tened, being morphologically nothing more than proximal areas of the lamina. As the plant grows, however, the region of the — stipe undergoes a kind of secondary thickening, resulting in a ringed structure; and, by the greater thickness of the ring on one or both sides of the original flattened organ than over the — edges, the whole stem changes from a flattened to a cylindrical : body. The pith in all such cylindrical stems, even when they : are very large, persists in its original thin and flattened structure, — and does not appear in the form of cylindrical medulla as it does — in Nereocystis. The pith is very often excentrically placed, and the number of rings of growth on one side may apparently be . twice as many as on the other. I have not found anywhere in the literature an adequate account of the anatomical basis of the phenomena which have been known and commented upon since the first discovery of ? Lessoniez ; and in the plates accompanying this paper an ©P© cial effort has been made to indicate the occasion for the ringed appearance so characteristic of the Lessonia stipe. It has long : been suspected that the rings are in no sense annual ring has been believed that they are connected with the developmes : of the tufts of leaves in such a way that for each cycle of — - there should be an additional ring of tissue developed ag : older portions of the stipe. No doubt this view i from the exceedingly rapid growth of these gigantic alge altogether reasonable to suppose that several rings ™g™ produced in a single season. tings of growth were apparent on one side © three upon the other. The pith itself looks almost , same in vertical section as in cross section, anid 15 ™ pedde an anastomosing web of loosely interlaced filaments pi : in gelatin and filled with reserve food materials. The pi 1900] OBSERVATIONS ON LESSONIA 327 not stop abruptly where the inner ring of cortical tissue abuts upon it, but occasional medullary filaments run out between the thicker walled and more compactly placed filaments of the cor- tex. Trumpet hyphz, abundantly represented in younger pith, are not so conspicuous in material of this age. The character- istic sieve-tubes of Nereocystis and Macrocystis, as described by Parker, Wille, Oliver, and others, have not been seen in Lessonia. The tissue immediately outside of the pith has in cross-section much the look of sclerenchyma. The cells in longitudinal section are not conspicuously armed, nor, as one looks at the cross-section under the microscope, do they seem to lie in radial rows. The inner ring on both sides of the pith principally composed of such cells as have been described. The transition to the next outer ring is more clearly marked in the gross structure than under the microscope, and at first I was much puzzled because the thin sections which showed the rings "ety conspicuously when held up to the light did not show them clearly when placed under an objective. The occasion for this seems to be that cells of the new growth are at first somewhat ne armed, and in places stand in rows extending in a radial direction and at the same level. The walls, except over the tads of the arms, are very thick. This has been shown both in diagram and in camera-lucida sections in figs. 8-76. Cross see tions taken through a region where the armed-cell tissue con- -_ Such radial rows at the same level will here and there cut “ight Or more cells in such a way that the whole row comes into ey looking somewhat like a medullary ray. Since eae . eae rows are not all cut through the central ese Res. ee will preserye its tee ae poe .. e ‘medullary rays, s0-Calle pees i te pith rays), come out as streaks of lighter app vies ee “ross section. As the new ring of growth grows older . “segs their arms and become more condensed in see oe : his shortening of the arms, together with a less a : | hae Giren makes the light streaks shorter, sag ees Ch Yindistinguishable from the structure around them 1 328 BOTANICAL GAZETTE [NovEMnER section. The difference then between the inner and outer face of a ring of growth lies in the more armed shape of the inner — cells, and the less armed shape of the outer, and the optical characters of the more-armed cells in the tissue are rendered | peculiar by their tendency to stand in radial rows for some dit tance at the same level. In general the outer rings show longer rays and a generally coarser structure than do the inner, while the innermost ring of all around the pith shows the rays bet sparingly, and in cross section is almost everywhere strongly sclerenchymatous in aspect. In the outer zones I have seen 2 many as nineteen cells in one of the rays, showing that this number of armed cells stood at exactly the same level ina radial plane and were all cut directly through the middle when the section was taken. A comparison of the cross and long sectioms as presented in the plate should make this matter clear. None _of the zones of growth, except the one immediately around * ; pith, are free over any considerable area from the rays resulting in the manner described. Lying outside of the outermost - will be found the immediate products of the superficial cambium, not yet modified into the characteristic armed-cells, but present: ing a more isodiametrical character. The actual cambial pete seems to lie immediately under the epidermis. Th cortical region and extending down to the region of armed § renchymatous cells chloroplastids are abundant, and the ep mal cells themselves, with the layers immedia dense protoplasmic stains. There is probably . going on in several of the layers under the epidermis, would seem to be more active close to the epidermis pears deeper layers. Three or four layers of cells immediately . the epidermis are of the same prismatic shape as hee cells proper, and almost as broad as high. Deepet -_ oe elongated in a direction perpendicular to the pie their walls become thinner. In vertical section they are te lie in definite rows. The distinction made by wes hold between outer and inner cortex in Lessonta ovala seems 10 good also for Lessonia littoralis, but there is a gradual 1900] OBSERVATIONS ON LESSONIA 329 from one area to the other, and the most abrupt demarcation is that between the inner cortex and the pith. Indeed cells which begin life as components of the outer cortex may later become modified into components of the inner cortex, as the stem thickens. Some peculiarities of structure were noticed, especially gobose thickenings where the arms of cells came together in the ray-like rows of inner cortex cells. These bodies of cell- wall substance in their position and form remind one quite exactly of similar structures common in Rhodophycee and recently figured for Gigartina by Miss Olson.’ The © Hohl- tiume” of Grabendorfer I have not encountered in any of the sections under observation, but it is not improbable that they might exist in this species in older stipe areas. Granular cell contents are pretty abundant throughout the plant. The pith is particularly well supplied and the sclerenchymatous tissue of - inner cortex contains an abundance of granules. In general it may be said of the older stipe of Lessonia alis that it consists of a strongly flattened pith surrounded by an inner cortex in which several zones of growth may be Mesent, and an outer cortex consisting of a generally cambial Soup of cells arranged in from ten to twenty layers. Secondary : thickening originates in the superficial cambial region, and the ‘mate appearance of a mature stem is due to the rhythmical _ Production of more-armed and less-armed cells; but the differ- “nce between these is so slight that the optical distinction se pad be accounted for without taking into consideration pe Sete. _ aes of cells, especially in later zones, ! “eae -- same level. and the str a t ¢ damnina.—~ The stipe beconie tionarea oe oe hal Soong prs ee: Lael tee Weanica ¢ the longitudinal rifts originate by ee ieee - Sctioning th sats each other, the structirs can : kas early Seat petiolate bases of laminae. These ar sink. yindrical, but generally more or less oval in cross § *Olson ; . ison : Observations on Gigartina. Minn. Bot. Stud. 2: 154- 1899- much flattened At the transi- hich the laminae 330 BOTANICAL GAZETTE [NOVEMBER In this region the pith is large, composed of slender interlacing hyphe, the majority of which seem to run transversely. Mingled with the ordinary hyphe of the pith web are numerous trumpet hyphe with the well-known sieve-plates, where the flared-out ends of the cells come in contact with each other. Very many ot the trumpet hyphz run transversely in the pith web, and they are perhaps rather less abundant in the perimedullary portion where they take principally a longitudinal direction. The inter- stices between the hypheare filled with gelatin, and immediately outside of the general mass of interlocking and interlacing fila- ments is a sheath of sclerenchyma, between the longitudinally extended cells of which occasional pith hyphe extend trails versely. The pith in this region occupies about one third of the diameter of the cross section. The remaining area is composed of three well-marked layers. Immediately outside of the pith is the narrow band of sclerenchyma just described, an area om staining deeply with aniline-water-safranin. Outside a this sclerenchyma lies a band of fundamental tissue having the appearance of parenchyma in cross section. The content cells are not strongly granular. Outside of the last mention region is chlorenchymatous tissue, in which the cells eg! marked tendency to develop with their long axes perpendi : se to the epidermis. Surrounding the whole is @ eee epidermal layer of low prismatic cells. The longitudinal se" through this area does not show a development of the of : cells so abundant in older stipe and characteristic of gee of secondary thickening, but the layer directly under the ¢ | chyma is composed of almost isodiametrical cel!s " om the ing in form to elongated-prosenchymatous in passing ond Oe periphery towards the pith. The cells immediately ne pith are very long, some of them measuring 1™° fon es not while the lumen as revealed in cross sections is sone over 24 in diameter. This condition reminds one 5 ee = shaped sieve-tubes of Nereocystis, which occur in the Bos callus ing region of young stipe and lamina, but no end see in othe has been detected in these cells of Lessonia, whic = Is rapidly chang” 1900] OBSERVATIONS ON LESSONTIA 331 respects resemble the true sieve-tubes in form, There seems to be the same evidence of passive elongation and occasional obliteration that was noted in Nereocystis, and the same frag- mentation of nuclei no doubt initiates the process of elongation. Upon the comparison of Lessonia with Nereocystis, it would seem reasonable to describe the “true sieve-tubes”’ of Wille and Oliver as modified sclerenchymatous elements of the perimedul- lary cortex, The splitting of the lamina has not been definitely studied in this material, but so far as can be judged from the observations that have been made it takes place in the same manner as pre- viously described for Nereocystis. The lamina.— The essential structural basis of the lamina and me arrangement of the tissues indicate its complete morpholog- al equivalence to the stipe. The pith is of course greatly extended into a narrow ribbon, the cells of which while retaining the web structure are not conspicuous for granular contents, indicating that the stipe serves in some sense as a reserve organ, Maile the leaf, essentially photosynthetic, uses its pith more Particularly as strengthening tissue and a conduction path. On ag of the ribbon-shaped pith parenchymal’ tissue - “Yeloped outside of a narrow intervening layer of sclerenchyma. pil the epidermis chlorenchyma cells extend, ares of the S layers. The epidermal cells are not so low as a “a stipe, but generally show a long diameter perye ag ing - the surface of the lamina. A well marked cuticle a0 - ee 1s uniformly present. The edges of the epee to the aad ae (belonging in part to the pith a a oe give stre ex) with small diameter and thick we s, ete ‘ ves it is probable tha ae gth to the margin. In younglea ance of b — is not immediately differentiated. bei noe ae . midribis caused simply by the greater hi . ed the central portion of the lamina and its . i th Pr thinning out at the sides. I was unable to dieteg™ : : ing tract any distinction between that portion which lay ™ midrib and the portions on either side, nor do the cortical 332 BOTANICAL GAZETTE [NOVEMBER cells differ in character in midrib and wings. The midrib may therefore be described in Lessonia as a central longitudinal hypertrophic cortex-area of the lamina. The structure and development of the sorus.—The development of the sorus in Lessonia littoralis does not differ in any important respects from that described by many observers for other genera of the Laminariacee. The epidermal cells elongate into para- physes, from the bases of which the sporangia arise. The spo- rangia themselves in some instances become almost as long as the paraphyses, reaching a length of 45, but this is unusual. Commonly they are from one half to two thirds as long as the slender paraphysal filaments, and of an elongated ellipsoid or club-like form. The zoospores are about 4m in diameter, and have been observed in different stages of formation. A peculiar condition which I have not seen in other Laminariacee exists in the exfoliated cuticle. This during the extension of the epl- dermal cells into paraphyses has become greatly thickened, and may separate eventually after the manner described long ago by Thuret.? It does not always, however, in Lessonia separate as # continuous membrane, but is often broken up into pieces corre- sponding to the original epidermal cells and retaining a genet ally prismatic outline. Each paraphysis over a large area may carry on its end such a little cuticular cap. : oe The drawings of sections have been made by Miss Josephine 4 E. Tilden, to whom I must express my thanks not only for | . c assistance, but for the specimen upon which my observations - have been based. THE UNIVERSITY OF MINNESOTA. EXPLANATION OF PLATES XIX~KRE . drawn with ra one ball All sections not described as diagrammatic were and reduced camera lucida under a magnification of 600 diameters, in reproduction. PLATE XIX. -Lessonia littoralis, from photograph. 7THURET: Ann. Sci. Nat. Bot. 1850. 1900] OBSERVATIONS ON LESSONIA 333 PLATE XX, Fig. 1. Diagram of end of branch bearing two pairs of leaves, the outer pair without midribs and bearing sori, the inner with midribs and without sori. - Fig. 2. Cross section of haptere showing epidermis, superficial cambium, and ground tissue. : Fic. 3. Longitudinal section of haptere through superficial area. Fic. 4. Longitudinal section through central portion of haptere showing elongated parenchymatous cells. 1G. 5. Section through older area of holdfast showing irregular and dis- torted structure due to appression and coalescence of original haptere ranches, Fic. 6. Diagram of cross section through stipe 12° above holdfast show- ing flat pith and zones of secondary thickening. Fig. 7. Cross section through outer cortical area of stipe 1% in diameter tg epidermis, cambial tissue, chlorenchyma, and transition to scleren- chyma, . Fig. 8. Cross section through zone of secondary thickening showing por- tion of a “ray’’ caused by the section striking the middle of adjacent cells in atadial row ; between two of the cells button-shaped masses of cell-wall sub- Stance have been developed. Fig. 9. Section through same area as last showing two “rays ee slightly different levels so that the shapes of the component cells are differ- ent: the majority of the cells have been cut across their arms giving a sclerenchymatous appearance. ts Fic. Io. Cross section through the zone of tissue nearest to the wing the absence of arms and consequently of “ rays,”’ and illustrating the a sclerenchymatous appearance of this region. : : u, Cross section through the pith of same stipe showing the ne web of hyphz with cells packed with reserve materials. : i. fais Longitudinal section through outer cortex and epidermis ; transi- armed to outer cells. pith PLATE AX, 13. Longitudinal section through secondary growth zone of stipe ement side by Fic, maintained by the dinary sclerenchy- “ rays” as in = i . re armed oN Diagrammatic representation of the position Matous “"S) Cross sections at point a would show the or Ag. 9 “tucture, while those made at point 4 would show the a wing on the left Fig, cS ay a Longitudinal section of stipe near the pith s a © the pith web, and on the right the sclerenchymatous cells in ner Stowth zone, : 334 BOTANICAL GAZETTE Fic. 16. Diagram of cross and longitudinal sections th 0 late base of the lamina; a, outer cortex and chlorenchyma; 4, pz tous tissue; c, sclerenchymatous layer around the pith ; d, pith. G. 17. Detail of cross section through same area as last cal chlorenchymatous, and parenchymatous tissue. _ Fie. 18. Cross section sane area as last and oe t out from the central pith. Fig. 19. Longitudinal section through the region of be _ Fic. 20. Longitudinal section through the region of fig. 18 1G. 21. Cross section of lamina through the wing.” Fig.-22.. Section through a sorus showing sporangia with BOTANICAL GAZETTE, XXX PLATE XIX MACMILLAN on LESSONIA ICAL GAZETTE, XXX MACMILLAN on LESSONIA PLATE XXI "BOTANICAL GAZETTE, XXX => = = Z ° nN WY ea) a | 3 Z. < oo = = oO < = STUDIES IN CRATAGUS. II! C. D. BEADLE. Crategus aprica, n. sp.—A large branching shrub, with one _ several stems, 3-5™ tall, or occasionally arborescent and . attaining a height of 6-7" under favorable conditions: bark of | he trunk and larger stems dark gray or nearly black, often con- §picuously furrowed, the surface being broken into small, irregu- lar, persistent, plate-like scales: branches ascending, armed with - Stout, either straight or slightly curved spines 2-6™ long, which : ae frequently branched and of greater size on the trunk and larger branches; branchlets at first villose-pubescent, but ulti- _-‘Mately glabrous, marked by numerous small, pale lenticels, the : bark reddish-brown, after the first winter changing to gray, with ‘ges of red or brown, the whole forming a compact, oval, or fund head: winter buds globose, bright reddish-brown, the _ Miter scales of the terminal ones thick and pointed: leaves thin ‘: ‘9 subcoriaceous in texture, obovate, rhombic-ovate, or orbicular outline, T.5-7™ long including the petiole, 1-5™ wide, the a ‘ders dentate or crenate-dentate and conspicuously glandular, ae or less lobed near the acute apex, or on vigorous shoots. Sa lobed, especially below the middle of the blade, i“ Y Narrowed but sometimes rounded at the base and pro- . ae amargined petiole 7™™—2™ long, which, like the base : ea bears numerous black colored glands ; stipules linear, -tinate} neeolate, or on strong shoots foliaceous and lunate, Pee a ¥-glandular or glandular-serrate, caducous: flowers, which “They a = vicinity of Biltmore, North Carolina (type seer ia . ata are nearly grown, borne in 5-6 flow es : egg Peecent, bracteate corymbs ; pedicels 1-2 pt deciduous ay bearing one or two small pectinately-glandular, ao tactlets: calyx obconic, pubescent, at least near the “ag from Bor. Gaz. : 28: 417. 1899. 335 336 BOTANICAL GAZETTE [NOVEMBER — base, the divisions 3-5™™ long, glandular-serrate, or pectinately- glandular: petals rather broader than long, g-13"™ by 8-12", with a short broad claw at the base: stamens 10, 5—8™™ long, the anthers light yellow: styles 3-5, surrounded at the base with pale hairs: fruit globose, 9-14™™ in diameter, red or orange-red, ripening and falling after the middle of September, the flesh thick, yellowish, pleasant to the taste: nutlets 3-5, hard and bony, 6-8™™ long, 3-4m™m measured dorso-ventrally, the back ridged and grooved and the lateral faces nearly plane, a volume of 125° containing about 1598 clean and dry seeds. Crategus aprica has been confounded with C. fava Ait. from which it differs in the shape and color of the fruit. The new species is abundantly represented in the mountainous region of North Carolina, and has been found 4 in similar situations in Tennessee, Alabama, and Georgia, inhabiting sunny exposures in dry, rocky, or clayey soils. The type material is preserved in the Biltmore herbarium. Crateegus sororia, n. sp.—A tree 5—7™ tall, with a trunk I-15" in diameter, dividing two or three meters above ground into several stout, ascending or spreading branches, which form an oval or round head; or usually smaller, 3—4™ in height, forming a large shrub with one or more stems: bark gray, tinged with brown or nearly black, furrowed and broken on the surface into small, persistent scales: branchlets armed with gray oF chestaut- brown spines 1.5—3.5°™ long : buds globose, bright reddish-brow?: leaves 2-6™ long, including the petiole, I-3™ broad, or on Vif" orous shoots sometimes 6™ broad, obovate, round-ovate, S ‘ . | all nearly orbicular in outline, or on the shoots even pete at long, with a truncate or subcordate base, acute OF dae e the apex, either gradually narrowed or abruptly contract base and prolonged into a margined, glandular petiole 5~ isel¥ long, the borders sharply and irregularly serrate and Bee lobed, especially above the middle, the serratures es yx the apiculate ; sparingly pubescent when young (at eae ni ith petiole, midrib, and principal veins), becoming glabrous: ing the a few hairs in the axils of the prominent veins and border! * Hort. Kew 2: 169. 17809. mo_1 5% 4900] STUDIES IN CRATA:GUS 337 petiole, bright green on the upper surface, paler below, fading inthe autumn to tones of yellow and brown, or with occasional _ dashes of red: flowers, which appear in the vicinity of Rome, _ Georgia (type locality), during the last of April or first of May, aid when the leaves are nearly grown, borne in pubescent, @landular-bracteate 3—6-flowered corymbs; pedicels 5-15™™ long, sparsely pubescent, bearing one or more pectinately- _ §andular, caducous bractlets: calyx obconic, usually with a few - soft hairs, the divisions 6-8™™" long, glandular-serrate: stamens tormally 20 : Styles 2-5, commonly 3, surrounded at the base with pale hairs: fruit large, globose, 12-18™" in diameter, red, - Ted and yellow, or yellowish-red, ripening and falling after the middle of September, the flesh thick, soft, and pleasant to the ; laste. nutlets usually 3, hard and bony, 7-g™ long, 4-5" thick, measured from the back to the inner angle, the lateral faces nearly plane and the back ridged and grooved. ‘i Crategus sororia is related to C, africa above proposed and to C, flava 0, 4.¢. From the former it may be separated by the more numerous er fruit and calyx segments, and coarser seeds; while from the pecies it differs from the accepted figures and descriptions which tawn from specimens in cultivation in Europe, in the shape of the he pubescent corymbs and petioles. The proposed species is represented on wooded hills, slopes, and rocky exposures, and in rom northwestern Georgia and adjacent Alabama southward to hast tamed s have been q it and ¢ abundantly ‘ld fields f ~ Plotida. pte material is preserved in the Biltmore Herbarium. : wi tet Alleghaniensis, n. sp—A large shrub 2-4" tall, or ti uty a small tree 5™ in height: bark gray, sometimes ay brown or much blackened: branches ascending, he slender, gray or reddish-brown spines 1.5-4™ long, — Sason 4 forming an oval or round head; the growth of or - fale le othed with reddish-brown bark and marked by smal ; - * cels: leaves ovate, oval, or nearly orbicular in outline, including the petiole, 1.5—6™ wide, very sharply and serrate and incisely lobed, acute at the apex, either “"rowed or rounded at the base, or on vigorous shoots * and prolonged into a margined, glandular petiole * 2 long, . gu arly bruptly n -Sibcordat paye 338 BOTANICAL GAZETTE [ NOVEMBER 5™™—2°™ long, bright green on the upper surface, slightly paler below and with 3—4 prominent pairs of veins, sparsely pubescent when young, especially on the upper surface, soon becoming glabrous: flowers, which open in the vicinity of Valley Head, Alabama (type locality), the first of May, disposed in simple, 3-6-flowered corymbs ; pedicels 1-2 long, bearing one or more small, linear, or lanceolate, pectinately-glandular caducous bractlets: calyx obconic, glabrous, the divisions 4—-6™ long, glandular-serrate: petals 7-9™ broad, g—12™™ long, the claw at the base narrow: stamens normally 10, 4-6™™ long, the anthers purple: styles 2-5, mostly 3-4, surrounded at the base with pale hairs: fruit, which ripens after the middle of September, globular-pyriform, red, 9-14™™ long, 8—12™™ broad: nutlets 2-5, usually 3—4, hard and bony, 5—7™™ long, about 3” thick meas- ured from the back to inner angle, the lateral faces nearly plane and the back grooved and ridged. Crataegus Alleghaniensis is abundant on Lookout mountain above Valley Head, Alabama, growing on rocky exposures or in the shade of oaks and pines. It is related to C. africa above proposed, from which it may be dis- tinguished by the less glandular foliage and inflorescence, the form and darker color of the fruit, purple anthers, and the sharply serrate borders of the leaf-blades. The type material is preserved in the Biltmore Herbarium. Crategus venusta n. sp.—A tree seldom more than 8” tall, or frequently a large branching shrub growing on rocky a or occasionally on the banks of small streams: flowers, W Alabama open about April 20, in the vicinity of Birmingham, ee. (type locality), and when the leaves are half grown, 2-2.5 ei occasionally more in diameter, disposed in 3-6- flowered ae : pedicels 1.5-2™ long, glabrous, bearing two or three pectina | glandular, deciduous bractlets which vary from 5" ' 1.5 ae length and from 2-4™" in width: calyx obconic, — 3 segments acute, 4-6™ long, 1.5-3™™ wide, glandile . often pectinately-glandular below the middle: petal 2 an orbicular, rather broader than long, g-12™™ wide, 8-10 with a short and broad claw at the base: sta 4-8™™ long, the anthers yellow : pistils 3-5, surrounde 1990] STUDIES IN CRATAGUS 339 base with pale hairs: fruit which ripens and falls after the first _ of October, globose or slightly oval, g-13™" wide, 9-15™™ long, dull red to greenish-red when fully ripe, sometimes, when more _ exposed, brighter, and frequently presenting surfaces of russet- ted; the cavity 3-5™" broad and nearly as deep, surrounded by the remnants of the stamens: nutlets 3-5, hard and bony, _ &9"" long, 3.5-5™™ measured dorso-ventrally with the lateral faces nearly plane and the back grooved and ridged: leaves thin _ ‘subcoriaceous, sparsely pubescent when young, scon smooth, . bright green on the upper surface, paler below, showing 4-7 | pairs of prominent veins, from obovate to ovate in outline, occa- Sionally on strong shoots round-ovate, 2.5-12™ long, including the petiole, 1-6" wide, acute at the apex, rounded or narrowed = the base into a narrowly winged and remotely glandular ; Fnic 7™"-4™ long ; the borders irregularly or doubly serrate 4 = icisely lobed, with ‘minutely glandular-tipped serratures: Stipules linear or linear-lanceolate, pectinately-glandular, cadu- | “us: bark of the trunk varying from ashy-gray to light-brown, ‘ veitly fissured: branches spreading or ascending, bearing ) aa stout, dark chestnut-brown, or gray spines 3-7" long, 3 rthe older branches and frequently the trunk armed with a ring much branched spines of greater size; the bark reddish- _ ™1,marked by small, pale lenticels: buds globular, bright dish-brown, j of "eda venusta is abundantly represented in the Red Mountain region | of the . : was apparently first discovered by Mr. C. gr 3 ld oii and later by Professor C. 5. Be Se Bead - The new species is closely related to C. £4 3 | oa ee oc... nly in the more numerous-flowered ede ae | ind the oo” anthers, narrower and pine: me The ae a obovate and elongated outline : 8 Crate-eus i: es Is preserved in the Biltmore ns oe i ae More than ae Shei, n. sp.— A tree seldom attaining a mee a Note stems - ‘2 commonly a large branching po” ee Nidish brown ark of the trunk and older branches ps et see Bor, Gay _ Smooth on the smaller plants, fssur arg i+ 1899. le3 differing mai 340 BOTANICAL GAZETTE [ NOVEMBER slightly scaly on the trunk of larger individuals: branches ascending, armed with stout, simple or branched, gray or chest- nut-brown spines 2-5°" long, forming a pyramidal or oval head; . branchlets at first pubescent, becoming smooth, the bark reddish- brown or gray, sprinkled with small, pale lenticels: buds globu- lar, bright reddish-brown, the terminal on strong shoots with thick, acute, slightly spreading scales: leaves, which are about half grown at flowering time, ovate, round-ovate, or occasionally obovate, 3-9™ long including the petiole, 2-6™ wide, or occ sionally larger on vigorous shoots, rounded or acute at the apex, abruptly contracted or wedge-shaped at the base and pro- longed into a margined, pubescent petiole 5""-2™ long, which, as well as the base of the blade, bears several sessile or stalked dark colored glands, the borders sharply and often doubly set rate, the serratures tipped with minute black glands, the Tr surface pubescent, becoming nearly smooth with age, bright green and lustrous, pale green on the lower surface, sa densely and permanently pubescent, especially along the midrib and principal veins, which are displayed in 5-7 pairs, aasentt olate, stra pectinately-glandular or glandular-serrate: expand in the vicinity of Montgomery, Alabama early in May, produced in simple or branched Ee ter: glandular-bracteate, pubescent corymbs, 1.5-2.5°% ine calyx obconic, pilose or pubescent, the divisions 7-1o™™ long, 3-5™™" wide, smooth or nearly so tt face, pubescent on the inner, pectinately-glandular ms ee glandular-serrate, reflexed after anthesis: petals slightly bros than long, 10-14™" by g-12™™: stamens normally ste ier long, the anthers yellow: styles 3-5, surrounded at t sk with pale hairs: pedicels 1.5-3% long, pubescent pie bearing a linear or lanceolate, pectinately-glandulat fig fruit, which ripens and falls the last of September ve in diame” October, red, more or less pubescent, globose, pele: “roundel ter or occasionally slightly oval; cavity 4-68 - : 1900] STUDIES IN CRATAGUS 341 _ bythe persistent calyx lobes and remnants of the stamens: nut- -Iets 3-5, hard and bony, displaying a prominent ridge on the back, or conspicuously grooved and ridged, 6-g™ long, 4-6™" _ measured from the back to inner angle, the lateral faces nearly ~ plane, Crategus Ashei has been found in the abandoned fields and woodlands, _ ‘Snerally in clayey soil, of Montgomery county, Alabama. The species is : tlted to C. Harbison? Beadle,* from which it may be distinguished by the 3 comparatively simple and less floriferous corymbs, more lucid and less ; pubescent foliage, and by the attenuated calyx lobes. I take pleasure in | *ociating with this species the name of Mr. William Willard Ashe, forester _ ofthe geological survey of North Carolina. The type material is preserved in the Biltmore Herbarium. Crategus senta, n. sp.—A small tree 5—6™ high, or more { frequently a large shrub with one or more stems, bark of the | “unk rough, dark gray, usually much blackened near the base: } branches spreading, slightly pendulous or recurved, zig-zag, : clothed with smooth, dark or brownish-gray bark and armed Mth stout, gray or chestnut-brown spines 1.5-6™ long: leaves Sbovate, obovate-cuneiform, or on vigorous shoots round-ovate oly orbicular, 2-7 long including the petiole, 7™"-5™ Wide tart Carolina (type locality), and when the leaves are nearly 342 BOTANICAL GAZETTE [NOVEMBER broad: stamens normally 20, 5-7™™ long: styles 3-5, surrounded at the base with pale hairs: fruit globose, 1o-14™™ in diameter, red, ripening and falling the last of September and beginning of October: nutlets 3-5, hard and bony, 7-9™™ long, 4-5 meas- ured from back to inner angle, the back shallowly grooved and the lateral faces nearly plane. Crategus senta is abundantly represented in abandoned fields and in open woods near Biltmore, North Carolina, and was referred to by me in an article published in the BorantcaL GAZETTES under the name C. elliptica. The new species is related to C. Michauxi Pers.® (C. glandulosa Michx.,’ not Aiton or Wildenow), from which it may be recognized by the longer petioles and pedicels, the sharply cut and nearly smooth leaves and less glandular characters of foliage and inflorescence. The type material is preserved in the Biltmore Herbarium. Crategus Alabamensis, n. sp.—A tree 4-6™ tall, or more commonly a large branching shrub with one or more stems, and spreading, often pendulous branches: bark rough, gray tinged with brown, or much blackened near the base : branchlets at first villose-pubescent, often puberulous in their first winter, but ultimately glabrous, gray tinged with reddish-brown, 0 ee : brighter on the younger wood, slender, zig-zag, armed with dark gray or chestnut-brown spines 1.5—4™ long, or sometimes longer and branched on the older branches and trunk: winter buds globose, bright reddish-brown: leaves obovate or Oba form, rounded at the apex and often with a short point 6 eS end of the midrib or occasionally very abruptly contracted 1" an acute tip, gradually narrowed or cuneiform at the base x prolonged into a margined, pubescent, glandular pare? 2 2.5™ long, 2-7™ long including the petiole, 1-3.5™ pene even larger on the shoots, pubescent at the time of er ole especially along the principal veins, and at maturity ee ceous or thinner, bright green and lustrous above an ie on the lower surface, the borders crenate-dentate = ge the especially above the middle, glandular-serrate OF entire ne4 base; stipules linear or linear-oblong, on the strong § 5 Bot. Gaz. 25: 447. 1898. 6 Syn. Plant. 2:38. 1807. 7 Flora Bor.-Am. 1? 288. 1803- 1900] STUDIES 1N CRATAGUS 343 lunate or variously lobed, pectinately-glandular or glandular-ser- rate, caducous: flowers, which appear when the leaves are almost fully grown, borne in simple or branched, 3-9-flowered, densely-pubescent corymbs, and open in the vicinity of Mont- gomery, Alabama, (type locality), early in April; pedicels 1-2.5™ long, densely pubescent, bearing one or more small, glandular, caducous bractlets: calyx obconic, pubescent, the divisions 6—-8™™ long, glandular-serrate, reflexed after anthesis: petals orbicular or longer than broad, about 1™ in diameter, with ashort and broad claw at the base: stamens normally 20, 5-7™™ long, the anthers yellow: styles 2-5, usually 3, surrounded at the base with pale hairs: fruit large, elongated, 1.5-2™ long, E15 wide, red, ripening early in August: nutlets usually 2-3 hard and bony, 8—10™™ long, 3-4™™ measured from the back to the inner angle, the lateral faces nearly plane and the back Stooved and ridged. Crategus Alabamensis is abundant in dry, clayey soil near Montgomery, Alabama, the fruit, less glandular foliage, and attenuated calyx segments. The type material is preserved in the Biltmore Herbarium. a Crategus Pinetorum, n. sp.—A shrub 1-5™tall growing in _* tocky woods where the prevailing forest growth consists : : pines, oaks, and hickories: stems one or more, clothed with eee roughened, dark gray bark, which is frequently Natkened near the base: branches slender, armed with slender, ; —" patel dark gray or chestnut-brown spines 1-5 a. ‘Sa % . of the season covered with smooth, reddish- inthe * which is marked by small pale lenticels, becoming °F obovate ay pubescent on the midrib and veins on the upper sur- soon glabrous, acute at the apex, sharply and and incisely lobed, the serratures minutely ed int , Narrowed or rounded at the base and pro- © 4 Margined, sparsely-glandular petiole 1-2.5°™ long, 344 BOTANICAL GAZETTE [NOVEMBER thin in texture, bright green on the upper surface, below paler and displaying 3-5 pairs of ascending, prominent veins; stipules linear or linear lanceolate, glandular, caducous: flowers, which open in the vicinity of Albertville, Alabama (type locality), about the first of May, and when the leaves are nearly grown; produced in simple, glandular-bracteate 3—6-flowered corymbs ; pedicels 1-2 long, bearing one or more narrow, pectinately-glan- dular, caducous bractlets; calyx obconic glabrous, the divisions 3-5™™ long, glandular-serrate : petals orbicular, or a little broader than long, about 8—ro™™ in diameter : stamens normally 20, 4-6" long: styles 3-5, surrounded at the base with pale hairs: fruit, which ripens in September, subglobose, 7-10" in diametet, changing from tones of green and yellow to light red when fully ripe, the flesh thin: nutlets 3-5, hard and bony, 5-7" long, 3-4™™ measured from the back to the inner angle, the lateral faces nearly plane and the back ridged and grooved. Crategus pinetorum is probably related to and easily cont Boyntoni Beadle®, from which it may distinguished by the sma more numerous stamens. The type material is preserved in the Biltmore Herbarium. rasted with C. ler fruit and A shrub 1-4™ tall, growing in uch branched, clothed hly fissured Crategus rubella, n. sp. upland woods: stems one or several, m _ with gray or reddish-brown bark, either smooth or slighl'y and scaly: branchlets numerous, armed with slender, straight oF slightly curved gray or chestnut-brown spines 1.5-4™ long; "3 on the older plants nearly destitute of spines back to - — branches: leaves oval, ovate or obovate, 3-9™ long including the petiole, 1.5-4.5™ broad, thin at first, becoming ne we ure, sparingly pubescent when young especially on ee surface, soon glabrous, sharply and doubly serrate to near base, and incisely lobed above the middle of th the apex, narrowed at the base and prolonged into @ sparsely-glandular petiole 1-2.5°™ long, reddish-green Of purp ler at the time of unfolding, becoming bright green eT a | below and fading with decided tones of yellow 5 stipules ae ® Bor. GAz. 28: 409. 1899. : 900] STUDIES IN CRATAGUS 345 or linear-lanceolate, pectinately-glandular, early deciduous: } flowers, which appear when the leaves are nearly grown, borne | insimple, 3-6-flowered glandular-bracteate corymbs, and open- ing in the vicinity of Valley Head, Alabama (type locality), _ about the first of May ; pedicels 1-2°™ long, bearing one to three pectinately-glandular, deciduous bractlets: calyx obconic, the ; segments 4-6™" long, glandular-serrate: petals rather broader than long, 8-12™™" wide, 7-10™™ long, with a short, broad claw atthe base: stamens normally 10, sometimes united in pairs and appearing to be fewer, 5-7™™ long, the anthers light purple: Styles 2-4,rarely 5, surrounded at the base with pale hairs : fruit red, pyriform or oval, 12-15™™ long, 10-12™ wide, ripening aiter the middle of September: nutlets 2-3, rarely 4-5, hard and | bony, 6—7™™ long, 3-4™" measured dorso-ventrally, the lateral faces nearly plane and the back ridged and grooved. Crategus rubella is abundant on Lookout mountain above Valley Head, Alabama, growing in the shade of oaks and pines, and has been collected in _ “ilar situations in eastern Tennessee and western North Carolina. It has oan customary to refer this species to C. flava, C. rotundifolia, and C. ¢coc- 3 “nea, but Iam inclined to place it near and compare it with C. Boyntont "oe l.c., from which it differs conspicuously in the outline of the leaves, cee color of the fruit, and the purple color of the anthers. € type material is preserved in the Biltmore Herbarium. Crataegus straminea, n. sp.—A low shrub, about 1™ in height, ' oy growing in large patches in upland woods, or occasion- a pe oiling larger proportions, 2—3™ tall, and developing from AL ah stems a coarse shrub with loose or straggling ee a = gray, tinged with brown or reddish-brown, arme ' ae ner, curved or straight spines 1.5-6™ long which oe E a. = chestnut-brown in color: leaves oval, ovate © : Me in o d or a the and 346 BOTANICAL GAZETTE [NOVEMBER pubescent when young, becoming glabrous with age, or with a few hairs along the midrib and principal veins which are disposed in 3-5 pairs; petioles winged, 7™™—3.5°™ long, bearing several or many stalked glands ; stipules linear or linear-oblong, pectinately- glandular, caducous: flowers, which appear in the vicinity of Valley Head, Alabama (type locality), and when the leaves are nearly grown, disposed in glandular-bracteate, 3-6-flowered corymbs; pedicels 7™™—2.5°™ long, bearing one or two, pectin- ately-glandular, caducous bractlets :. calyx obconic, the divisions 4-6™™" long, glandular-serrate and with a few stalked glands below the middle: petals nearly orbicular, 6-10™™ in diameter, with a short, broad claw at the base: stamens normally 10, oy long, the anthers purplish: styles 3-5, surrounded at the base with pale hairs: fruit subglobose or pyriform, 10-13™ high, g-11™™" wide, yellow or greenish-yellow, ripening after the — of September: nutlets 3-5, hard and bony, 7-8™™ long, 3-4 measured from the back to the inner angle, the lateral faces nearly plane and the back grooved and ridged. ya Crategus straminea frequently covers large areas on the top of Loe mountain above Valley Head, Alabama, growing in the shade of oak; - and hickory trees, and will probably be found to extend into the adjace® regions of Tennessee and Georgia. It is related to C. Boyntoni Beadle, /. < from which it differs in habit of growth, form and color of fruit, color of the anthers, slender spines, and leaves with more sharply cut bare he type material is preserved in the Biltmore Herbarium. BILTMORE HERBARIUM, Biltmore, N. C. BRICFER ARTICLES. DAVID FISHER DAY (WITH PORTRAIT) Jupce Davip F. Day, who died on August 21, 1900, was born in _ huffalo, N. Y., in 1829, and his whole life was spent in that city. For more than fifty years he was engaged in the practice of the law, in which profession he held high rank. His love of nature, which was that of an enthusiast, early led him tothe study of the natural sciences, im which he became most profi- Gent. As a field botanist he was ticelled by few. Gifted with a *markable memory and a particu- ‘tly clear conception of the rela- lionships of families and genera, 3 he was able to place a new plant 4 with a facility that I have seldom wn equaled. His methods of fact that h © Was not a professional s Seems to have deterred him from publishing many of his Pee ee deductions, which were of great interest and pet ay with Judge Clinton he made a thorough study of a Butial Ry ne result appeared in “A catalogue of the plants . plants “icinity,” published in 1883. Later he prepared a list 0 q ‘lished gg on the reservation at Niagara Falls, which was pub- i as the state, He was one of the founders of the Buffalo Society 347 348 BOTANICAL GAZETTE [NOVEMBER of Natural Sciences, and a life-long member of it. To him alsoisdue _ the credit of establishing the Buffalo Botanic Garden, in which he was particularly interested. His library was a notable one, as he was a collector of rare dis- _ crimination, and the works on botany were many and valuable. In addition to the more pretentious volumes, a large collection of pamphlets was accumulated, among which appear a very large number of local catalogues. These books of botanical interest, as well as his col- lections of living plants and herbarium specimens, he was preparing to transfer to the Botanic Garden when he was stricken down. Mr. Day had for years been a member of the Park Commission of his city, and drew the act which created the Park department. In this act, drawn thirty years ago, he made provision for both botanical and zoological collections, both of which, after years of waiting, he saw established. His loss will be mourned by the many botanists of his acquaintance, as well as by his fellow-citizens, by whom he was held in high esteem. —Joun F. CoweE.., Buffalo Botanic Garden. OBSERVATIONS ON THE ROOT SYSTEM OF CERTAIN CACTACE:. , l Tuose who make botanical trips into arid regions do s0 generally with the one idea of collecting material to be worked up at leisure laboratories or herbaria, or to be deposited in botanic gardens. Theit stay, as a rule, is closely limited, and the necessity of covering ae amount of ground brings with it a tendency to pass over those de which take considerable time. The root system of desert pl as regards structure and distribution of roots, is one 0 which has perforce been to a great extent neglected, our generally speaking, being confined to the examination of th of underground growth collected with herbarium specimens. fore it seemed to me worth while during my stay in TucsoM, i ani in the midst of the great cactus plains, to make a rather carefu : of the roots of certain large forms which could scarcely be pres am toto, The root systems of Cactaceze, in general, are so! than would be expected. The distribution, however, 1 way to make good the deficiency in size and length. e amount 1900] BRIEFER ARTICLES 349 _ amples here cited may be fairly taken, I think, as types, as without - exception they agree in general characters. Echinocactus Wislizent Engelm.— Plants about 7.5% high and 3.5 tog in diameter gave the following results. Three or four main Mots, 10 to 12" thick, branch out horizontally from beneath the plant, taper but slightly, are sinuous and much branched at the tips, in such 3 amanner as pretty thoroughly to cover the ground about the plant | Nithina radius of 2.4". The fine rootlets are very numerous, gener- _ alyturning upward. In no place does this set of roots run much more _ than to™ below the surface. Just beneath the plant are a few small “ati passing almost directly. downward, tapering very rapidly, and vith numerous branchings penetrating to the depth of scarcely 30™. Opuntia fulgida Engelm.—An arborescent form. A specimen - tbout 15% high, considerably branched, showed, as in the above case, | “veal long slender roots with fine branches, running horizontally _ $0 10 below the surface, and covering an area with a radius of 30 . about the plant. In a plant of O. fulgida mamillata (Schott) Coulter, } Yhich was washed by a small torrent, these roots appeared above ‘ Jobat in many places. In both of these specimens a few quickly 7 '’apering roots were demonstrated at the base of the plant, passing - diteetly downward to the depth of about a foot. Opuntia Whipplei Engelm. showed a similar state of affairs. A Platopuntia (O. angustata Engelm.?) possessed much the same Cters. out roots and alee by innovation * but the chief roots are to be found attached 5 oldest living joint. At this point a double system can be seen, the surf a couple of large roots, horizontal and but slightly age “and one or two much smaller roots passing downward with «“Mpid branching, : ole - iil Engelm.—This plant thrives on x0 Lee a system is modified somewhat by conditions of soil an dons In the specimen examined the roots which passed directly ard came very quickly i ‘th a flaky bedrock, into the y in contact with a 7 ae oe Which they sent their branches, sometimes causing eae - beet do — Although these roots seemed to pass ——. ge re directed hee hold was exceedingly firm. The aggre € Mlace, by: —" entirely up the slope, sometimes gaseneee sae ‘Tov often descending to some little depth to avo | *Y,J.W.: Vegetal dissemination in Opuntia. Bor. Gaz. 20: 356. 1895: 350 BOTANICAL GAZETTE [ NOVEMBER bowlders. The area which they covered extended about 45°" away from the plant, which was between 12 and 15” high. Cereus enneacanthus Engelm.—This rather ascending, cespitose form showed, as in the other cases, the two sets of roots; horizontal, long and sinuous; and vertical, abruptly tapering and much-branched. Several specimens were examined. ConcLusions.—In the majority of the larger Cactacee there are two distinct root systems—vne horizontal, for absorptive purposes ; the other passing downward, for anchorage. The depth of the hori- zontal system varies with the degree of penetration of surface water, which, in turn, is to some extent dependent on the character of the soil. One of the specimens of Echinocactus was examined the day fol- lowing a gentle rain which had continued intermittently for two days, thus giving the water time to soak in as it fell. The soil was dampened to a distance of about four inches below the surface, just the depth of the absorptive root system. Observations upon young plants of Opuntia fulgida give the follow- ing as the sequence of root formation. The first to be developed by joint or seedling are the vertical roots, which for a time act both for ; ee d absorption and anchorage. As the plant increases In size the groun and in order developed onally appearing in part as horizontal branches of vertical roots. zontal system covers an area far greater than that sheltered by the parts above ground. As the soil is generally firm, and the horizon i give some support, there is little need for any roots to descend pa : great depth. The small amount of surface exposed to winds considered another reason for this. is Correlated with the difference in distribution of the two a eps a difference in structure, as may be proved bya most superficial se nation. The horizontal roots are in all cases extremely brittl¢ ter vertical much more elastic and capable of withstandin; gree j tension. The woody cylinder in large absorptive roots p his differ- relatively than that of the anchoring roots. Microscopically t ani ence may be most easily seen in the secondary xylem. For ae were tion sections of roots of Opuntia fulgida and Echinocactus Wist oe Ff used. The xylem of the absorptive reot is composed te of the ducts, that of the anchoring roots, which, after the app tal roots 900} BRIEFER ARTICLES 351 iorizontal system, seem to have developed for this purpose entirely, _masists almost wholly of wood cells. Owing probably to the general senness of climate the annual rings are not easily demonstrated.— _Carteron E. Preston, Harvard University. NON-SEXUAL PROPAGATION OF OPUNTIA. ProressoR Toumey, in an article in the BoTANicaL GAZETTE -0:356. 1895), speaks in general terms of the use of spines as aids in -Gsemination of opuntias which are dispersed by the breaking off of | lie separate joints. A short note may be added as to the function of these spines, especially in such long-spined species as O. fudgida : Engelm. A joint falling upon the sand very often rebounds from the tusticity of the spines, and by this impetus is carried some distance fom the parent plant. The greatest aid, however, is in the placing the joint. Joints destined for such dissemination are as a rule » ovate, the best developed areolae with the longest spines being situ- *sdon the distal end, those of the proximal end being scarcely at all Xt. The result of this is that the joint upon falling almost invari- Uy lights with its base downward, in the best possible position for i iy eee The distal parts are kept off the ground in all cases by : Rg spines.— CarLeton E. Preston, Harvard University. a GAURELLA =GAUROPSIS. I : uy “AVE to propose the restoration of the name of Gauropsis Torrey ; Fremont : Tor. Bot, ¢ | Bectea, 7 Pag Pres}? tees The type species of Gauropsis, Gaurella guttulata Set) Small ie ' Gaur ha sl a fy CURRENT LITERATUR. BOOK REVIEWS. Fossil plants. MorPuovoeists have made but little use of the work of paleobotanists, both on account of the nature of the material and because paleobotanists too often have not been trained morphologists. The time for this feeling, how- ever, has passed, and some of the most striking extensions of our morpho- logical horizon have come from the work of paleobotanists. What was needed more than any thing else was the critical sifting of the vast accumu- lation of paleobotanical data from the standpoint of modern botany. a 1891, Count Solms-Laubach published his admirable Fossé Botany, which has ever since remained a standard work for botanists, although more destructive of previous claims than constructive. Very recently, however, r. A. C. Seward has begun his great work on Foss¢Z Plants, but it has bags yet included the vascular plants; Professor R. Potonié has published his Lehrbuch der Pflanzenpaleontologie, a very compact presentation of the sub- ject; and M. R. Zeiller’s Eléments de Paléobotanique has just appeared. All of these books stand for the newer paleobotany, which is to reject uncertaln evidence and rest upon a foundation of morphology. chil What the morphologist wishes, however, is the omission of details w are not pertinent to him, and the compact presentation of what paleobotany has definitely contributed to morphology. This has been done, and appar ently well done, by Professor D. H. Scott, in a book which has just gi - His purpose has been “‘to present to the botanical reader those res fous paleontological inquiry which appear to be of fundamental pavenrere™ the botanist’s point of view.” Since the important results have related gr entirely to pteridophytes and gymnosperms, the Studies are ect gee them. It is certainly true that Professor Scott has presented in eee ably clear way just what botanists wish to know, but his data must be on paleobotanists. The book must be read for details, but some of the Striking suggestions for the consideration of morphologists are as e remarkable elaboration of Equisetales and Lycope ological Paleozoic is fully set forth, and should form a part of every morp”. presentation of these groups, the living representatives being sees quate for such a purpose. The most suggestive conclusion in t tion, however, is that these two groups, which seem to be S0 far a J > *Scott, DUNKINFIELD HENRY: Studies in fossil botany. eae 6d. with 151 illustrations. London: Adam and Charles Black. 1900. 7% [rovemBe® 352 1900] CURRENT LITERATURE 353 aebut divergent lines from an ancient stock represented by the Sphenophyl- les, Professor Scott regards the Sphenophyllales as worthy to stand asa gym- | Msperm group of primary rank. He would include in it not only Sphenophyl- lm and its acknowledged associates, but Cheirostrobus as well, a genus which strikingly combines the characters of Equisetales and Lycopodiales. _ Of course the Filicales are fully considered, a group of enormous Uutiquity, having held its own since the Silurian, and whose habit was so fredominant during the Paleozoic that many plants have been included with them whose real affinities are elsewhere. In general, Professor Bower's | Souping of Filicales into the Simplices, Gradatae, and Mixtae is shown to lave an historical basis. One of the most interesting sections of the book is that devoted to a con- Sileration of the Cycadofilices, a provisional Paleozoic group made to contain - fms which combine the characters of Filicales and Cycadales, and most of ; them first described as ferns. The representative genera are Lygino- - tendron, Heterangium, and Medullosa, whose habits and anatomical features _ % strikingly intermediate, but whose spore-bearing members are unfor- : ‘mately unknown, It is evident, however, that in them we have forms Ene confirm the conclusion that cycads have been derived from Aswas to be expected, the dominant gymnosperm group of the Paleozoic, : ee. is set apart as of primary rank, and, associated with the Cyca- 3 “gg seems to furnish the background for later gymnosperm develop- 354 BOTANICAL GAZETTE [NOVEMBER In discussing the general problem of the phylogeny of gymnosperms, the author fairly states the contending claims for its polyphyletic and mono- phyletic origin, but plainly inclines to the latter position, He sees no reason for imagining any connection whatsoever with the Lycopodiales, and still less with the Equisetales. The connection of Cycadales and Bennettitales with Filicales being considered as established, both through morphology and the paleobotanical evidence of such a group as Cycadofilices, “the derivation of the other gymnosperms centers about the extinct group Cordaitales,” whose affinities with the Cycadales are clear.—J. M. C Tropical Nature. COSTANTIN, the well-known French botanical philosopher, has issued a second volume in the series known as Bibliothéque Scientifique Internation- ale. His studies in various lines of botanical research, and especially his classic ecological papers on aquatic plants, have well fitted Costantin, both for his work on Adaptation-Evolution? and for the work which is the subject of this review.3 In the introductory chapters the author gives a realistic picture : of the virgin forest of the tropics, and vividly recounts the first sensations that one has in these new scenes. After a few remarks on tropical climates, Costantin plunges at once into the very deepest ecological problems of the tropics. He first discusses the origin of forests, and takes up some of the types of trees that are peculiar to tropical forests. An interesting chapter on leaves follows, and some of the most recent studies, as those on hydatl cs are introduced. After a few pages on flowers and fruits, the author give ad brief survey of the geological history of forests, holding to the older view that the present tropical forests are fragmentary, but yet lineal descendants of those which have passed away. . The lianas come in for an excellent treat- ment, four chapters being devoted to a discussion of the various classes cy their origin, Chapters follow on epiphytes, especially those found per sh tree tops; parasites of various classes; symbiosis, one chapter dealing wit) saprophytes, the other with the réle of ants and other animals in influenci"g = plant life. Saprophytism at one time would not have been given anon of symbiosis, but recent studies on mycorrhiza makes such ace haptet necessary. The strictly botanical part of the volume closes with ¥ pie ic on mangroves and one on island floras, the latter from the geogtP point. ‘nal a cally 0 Though the volume is entitled Tropical Nature, it is confined p << the plant world of the tropics, The last chapter is a curious excep p., with 177 ? Les végétaux et les milieux cosmiques (Adaptation-Evolution). 292 PR o figures in the text. Paris. 1808. ie >38 CosTANTIN, J.: La nature tropicale, 8vo. pp. 315- figs. 700. tee Alcan, 1899. 6 fr, pe 1900] CURRENT LITERATURE 355 _ deals with religious legends and traditions of the human race. This seems out _ ofkeeping with the rest of the work, which, though planned for a semi-popu- S lar audience, contains much matter of equal value to ecologists with the Works of Haberlandt, Schimper, and Warming. The figures are crude an # unsatisfactory in the extreme, when compared with Schimper. Though little _ thatis new is to be found in Costantin’s new volume, one finds there one of _ the best summaries we have of many topics, such as lianas and parasites. In the author's philosophy, also, is a great deal of valuable suggestion to all who _ ae interested in ecology.— HENRY C. COWLES. Organography. TO THE admirably selected series of German botanical works translated ’ p. 68, has crept in; now and then one finds ce, as “one would naturally expect that a lateral “toot removed from the shoot system and planted vertically and which rooted _ behave similarly,” p. 51; and we much regret that the incorrect form should be given further currency by the translator. But “S such as may be charitably overlooked in view of the great advantage of having this classic in English dress. We trust that the publi- d part (with its promised index) will be made possible at oy day by the appearance of the remainder of the second part from the author's hand. The imprint of the Clarendon Press is guarantee that the facture of the book is all that can be desired. —C. R. B. u NOTES FOR STUDENTS. i -H, Devaux has recently made an extensive study of lenticels, vad ea hg Concerns the physiological conditions of the growth and differentia- : Pi Of their tissues, the results of which occupy two thirds of a volume of the ales des Sciences Naturelles.6 4 ae nt B Organography of plants, especially of the A Ae: General gd Authorized English edition by Isaac BAYLEY ape eee Press, 1900 StaPhy. Roy. 8vo. pp. xvi+ 270. figs. 730. Oxford: 4 ': bs Gaz, 25:290. 1898. oy rails Siques ede H.: Recherches sur les lenticelles; étude sur les conditions a eat me Nat, Bot, yossement et de la différenciation de la cellule et des tissus. . VII. 121-240. p/.6. 1900. ed aaa y, 356 BOTANICAL GAZETTE [ NovEMBER The memoir is too voluminous to permit an adequate summary in the space at command, and the original should be available in every laboratory where research is prosecuted. Devaux gives a résumé of his work in the final chapter, from which some extracts are here made, Lenticels are found in all the great groups of vascular plants and on all their organs which have secondary growth. The primary number on the stem is in ratio with the vigor of growth (indicated by the length of shoots or internodes), with the total number of internodes, and with the numerical fank of each internode. The curve of their production agrees closely with the curve of elongation of the internode, though the maxima do not exactly coincide. In each internode the lenticels are almost always more numerous on the distal moiety. The dimensions attained by lenticels depends directly on their number per unit of surface. Two types are distinguished: (1) the closing layers are few in number, consisting of cells intimately united, with no (or small) intercellular spaces, being oftentimes comparable to true cork; (2) the closing layers are numer- ous, composed of rounded cells with many large intercellular spaces, and are like the packing cells (Fwd/zellen ; cellules comblantes), but suberized. The packing cells are alike in both. As to origin, lenticels are primary or secondary. Primary formed early and at points determined by an organ (stoma, rootlet, less often a bud). Secondary lenticels are formed later and at points not deter- mined by an organ. Whenever stomata are present there is a tendency t produce lenticels below them, usually in the cortex, sometimes in the per cycle. If stomata are very numerous, partial or complete abortion of net lenticels may occur. Some stems, wanting stomata, produce lenticels late, in the neighborhood of a bud after the fall of the leaf. lenticels are destruction to death or suberization of the cells. ation produces complete rupture of the closing layers, spring, followed by hypertrophy, death, or suberization (accompan™ a by sclerosis), and by centripetal displacement of the cambium or 18 | Boe r level. formation into permanent tissue, to be regenerated later at a deepe singe times a complete . osit¥ ae is occurs, not in winter only but throughout the year. Thanks to th ee they certainly serve in large measure in the general gascom the organs. But it would be false to say that the lenticels exist she pls exchanges. For (1) they are often absent or insufficient; (2) EF of the has porose regions different from lenticels; (3) open! eee af lenticels is not due to the needs of aeration. Rather they os och transpiration and automatic regulators of the internal moisture, ae - 1900) | CURRENT LITERATURE 357 plant uses efficiently for the proper gaseous exchanges also. (M. Devaux _ promises a further memoir on the general aeration of the plant by lenti- Finally, lenticels are defined as small limited regions of the superficial _ parenchyma in continual proliferation and continual development, capable of hypertrophy or of cicatrization, according to the conditions of external or _ internal humidity.— C. R. B ; ARECENT PAPER? on the embryology of Taxus daccata fills in some of ofthe gaps in previous accounts. The writer secured an abundance of wild material, but employed rather primitive methods in making his preparations, fixing in absolute alcohol, imbedding in celloidin, and staining in haema- toxylin. The following is a brief résumé of his work. The origin of the aril shows it to be a second integument. In tracing the development of the embryo sac the author was not able to get the earliest Stages. The two, four, and eight-celled stages of the embryo sac were not observed, but many sacs were observed in later stages. Free nuclear 1 _ tivision continues until there are about 256 free nuclei in the sac (the eighth division), when cell walls begin to appear. These cells rarely contain more than one nucleus, and have a regular six-sided appearance in optical section. ln the later stages of endosperm formation the cells are often multinucleate. The formation of archegonia begins at the end of May or the first of June, ) Dut archegonia in very different stages of development are soon found in the a Same prothallium, even embryos and young archegonia often appearing ether. The usual number of archegonia is from five to eight, but nine, "en, and even eleven were observed. Pollination occurs from the beginning ‘othe middle of March, and there is no so-called pollen chamber. By the of May three nuclei can be seen in the pollen tube, the tube nucleus, the huclei of the stalk cell and of the generative cell. Shortly before “vation occurs the generative cell divides into two very unequal cells. “ttilization takes place about the first of June. There may be several tubes, and several “archegonia” may be fertilized. The sex nuclei, ich are of about the same size, come into contact and then sink to the of the archegonium, where fusion takes place. Nuclear division then : ang until there are from ten to sixteen free nuclei at me base of the Pte ( ter which the free cells become arranged into ters, ; Fosette’’), the middle tier, consisting of suspensor cells usually ber but Sometimes more, and the lower tier, which is the embryo proper. long to a single ase three were F six in rest here are normally two cotyledons, but in one ¢ 3 -—CHARLES J, CHAMBERLAIN. PR ete, L.: Taxy Beitrige zur Kenntniss der Endospermbildung und zur Embryologie +4 bacrata L, Flora 86 : 241-288. pis. 15-79. 1899. 358 BOTANICAL GAZETTE ~ [NOVEMBER KAHLENBERG AND AUSTIN have been continuing the earlier researches on the toxic action of various substances on seedlings. They conclude! that the toxicity of acid sodium salts is greater than it ought to be if it were due solely to H ions and that the theory of electrolytic dissociation is unsatis- factory in explaining the poisonous action of these acid salts and of acids as well. The true explanation, they suggest, is very likely to a: found in the ability which the substances all have in common, to neutralize basic sub- stances. This explanation is independent of the theory of electrolytic dis- sociation. Kahlenberg also found himself unable to explain the sour taste of acids on the basis of the H ions? and holds the physiological effect tobe due to their chemical activity in virtue of the fact that the H is replaceable bya — metal of a basic radical. The more readily the H is replaced, the more — reactive the acids are and the more intense is their taste.—C. R. B. Dr. Gino PoLvaccr published a year ago the results of some of his researches on photosynthesis ® which have not been adequately noticed in this journal. His most important results are as follows: : Green organs of plants which grow in sunlight give the aldehyde reaction : with Schiff’s reagent. Under the same conditions fungi do not so react; nor do leaves, kept for some hours in darkness or in an atmosphere free of CO3. Formic aldehyde reactions are also obtained from expressed sap by proper treatment. He holds, therefore, that formic aldehyde is produced by gre? organs under the normal conditions of photosynthesis, and promises — the results of his researches on the process of its formation in a sec memoir—C, R. B. THE CORRECTION of a large number of typographical and other errors, both of omission and commission, in Engler and Prantl’s Pflanzery especially in the general index, will be found in Ad/gemeine Bot. — 1900: 110 e¢ seg. Otto Kuntze and Tom von Post are the ferrets. — e J inclined to magnify and multiply the errors, which they figure at 93 they have done a service for bibliographic work which will mc RB users of the Pflanzenfamilien some hours and much bad temper. —C. *Jour. Phys. Chem. 4:553-569. 1900. Jour. Phys. Chem. 4::533-537" me *° Atti Instituto Botanico de Pavia N. S. 73(1-21). 1899. NEWS. r Mr. Cyrus A. KiNG, formerly of Summerville, Mass., has been appointed _ instructor in botany in Indiana University. R. Witson Smiru, Ph.D. (University of Chicago), instructor in Mc- Master University, Toronto, during the past year, has just been promoted to the full professorship of botany in that institution. Dr. Oscar Loew, late of the Department of Agriculture at Washing- fon, has received and accepted a call to become again the professor of 4gricultural chemistry in the University of Toékyé. He sailed October 8. : - B. M. Duaear, assistant cryptogamic botanist of the Agricultural _ Experiment Station and instructor in botany in Cornell University, has returned from a year’s study in Germany and has been promoted to an assis- tant Professorship. PROFESSOR Dr. A. B. FRANK, director of the biological section of the Imperial Sanitary Bureau, formerly director of the institute for plant physi- in the Royal Agricultural School at Berlin, and author of a well-known text-book of botany and many physiological papers, died on September 27, at the age of 62. _ ABBE A, B. Lanexors died at St. Martinville, La. on August 1. His 7 long and indefatigable study of the flora of Louisiana was prosecuted in such yt as could be obtained in the course of his parish duties. He has sup- lied material most liberally for the study of specialists in all groups, and ot _ ane has been used for many new species which he discovered. His cryp- ‘ogamic Collections were probably left to the Catholic University at — : "gton, to which several years ago he gave his phanerogamic plants. bio UNDERSIGNED requests all foreign botanical writers to send to him a dd be ‘ach of their publications, as far as possible, and especially reprints _ Rals ong articles published in proceedings, transactions, and pert : ical ; learned societies, This request is made in’order that foreign = “oT may be reviewed promptly for Just’s Botanischer Jahresbe- aay bringing the same to the early notice and general use of all coe tad ROFEssor Dr. K. SCHUMANN, Botanischer Museum, Berlin, W. : “eg FOLLOWING report of the sixth annual meeting of the Botanical America, which was held in New York City, June 26 to 28, 1900, 359 360 BOTANICAL GAZETTE has just been received from the secretary. For the reading of papers the society met in joint session with Section G of the American Association for the Advancement of Science, June 28, in room 502, Schermerhorn Ha Columbia University. The meeting of the sections was called to order by the vice president, William Trelease, who announced the arrangements for the joint session, and called B. L. Robinson, president of the society, to chair. The retiring president, L..M. Underwood, then read his add “The Last Quarter: A- Reminiscence, and an Outlook.” The fu Il i text of address has been printed in Sczence. : Following is the program of papers ee : The significance of transpiration, C. X. Barnes Relationship and vari.bility of the Adirondack spruce, Charles Peck; Nuclear studies on Pellia, B. AZ. Davis ; a On the structure of the stem of Polytrichadelphus dendroides, Mrs. E. C. Brit fon, Observations on the group Yucceze, Wed/zam Trelease; Spermatogenesis in the gymnosperms, /. 47. Coulter; The pollen tube, and division of the — rae in pines the council), A/iss 7. C. Ferguson, On the homologies and probable origin of the embryo-sac, Son; Observations on Lessonia, Conway MacMillan ; Thigmotropism of roots, /. C. Newcombe ; Starch in guard cells, B. D. Halsted ; Coenogametes, B. A7. Davis ; The development of the archegonium, and fertilization in the (by invitation of the council), W. A. Murrill; The causes operative in the formation of silage, H. L R coc. A closed circuit respiration apparatus, wT: Russell and S. The officers for the ensuing year are: president, B.D. H president, R. A. HARPER; treasurer, C. A. HOLLICK; ‘Secres ATKINSON ; councillors, C. E. BESSEY, F. V. CovILLE. An important step was taken by the society in appointing consider the best means of realizing the purposes’ -of the advancement of botanical, knowledge.” Among other thin, : will consider the uses to which the accumulating funds of the put. The committee will report at the next annual meeting poe —GEORGE F. ATKINSON, Secretary. \Tonic and Nerve Food oe 50 odonr HORSFORD’S in a new size Acid Phosphate. 25c. When exhausted, depressed of the Liquid weary from worry, insom- The event of the la Or overwork of mind or year in dentifrices. body, take half a teaspoon of Beware of counterfeits dorsford’s Acid Phosphate in and substitutes of this, theworld’s best known i j alfa glass of water. getting the genuine i the stores. It necessa: send 25c. direct to the Propritos P. 0. Bo 247, New It nourishes, strengthens and im- | parts new life and vigor by supplying j te needed nerve food. 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WEBER WAREROOMS: AYAPAPAPAPAPAYAPAPAYAG KYALY EYE AP AH EH EDS hee PAP APAPAPAPAR EAE AYAP APKPEPADKHKH eye DORCHESTER, MASS. 108 Fifth Avenue, New York on Every Package Established 1780. 268 Wabash Avenue, Chicago 'v, eS papa ET eee eee 181 Tremont Street, Boston. You see the Cook @ has raiced t To take the Mayor to the show And Spotless Town is in surprise 7 see them win the cake-walk prize- e pa it cleans the “Stet RA eevee rer: XXX DECEMBER, 1900 ‘No 6- THE BOTANICAL GAZETTE a EDITORS JOHN M. COULTER anp CHARLES R. BARNES, WITH OTHER MEMBERS OF THE BOTANICAL STAFF OF THE UNIVERSITY OF CHICAGO ASSOCIATE EDITORS UR — University FRITZ N ae? ae ys - one R DECANDOLLE miversuy 2 . “Geneve ‘ VOLNEY M. —— it oe &TONT University of Michigan University ROLAND eer P ENGLER a Harvard University University of Bertin WILLIAM ‘TRELEASE eae GUIGNARD Botani ~ L . . aa de Pharmacie, Paris H. MARSA Dasa far ARPER EUGEN. WARM fe ersity 2 epenhagen Uni: Dra: of Wisconsin Imperia : cx Z Venice, Tokyé vere wirtROCk peu of Sciences 2 CHICAGO, ILLINOIS Ublishex bp the Giniversity of Chicas? be Sniversite of Cdicage Press co PYRIGHT 1900 BY THE UNIVERSITY OF cHicaGO = obtained 2 the only — | The Highest Toilet $ Botanical Gazette montbty Journal Embracing all Departments of Botanical Science ‘per year, $4.00 Single Numbers, 40 Cents subscription price must be paid in advance. . No numbers are sent after the expiration of the time paid for. No reduction is made to dealers or agents. FOREIGN AGENTS: ain — Wa... WESLEY & Son, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin on, 18 Shillings. SW. 46, Schonebergerstr. 17a, 18 Marks Issued December 20, 1900 CONTENTS ROMATIC SPINDLE IN THE SPORE MOTHER CELLS OF OSMUNDA ALIS. ConTRIBUTIONS FROM THE HULL Soe Lanna sane are xxl). R. Wilson Smith . 361 IN CHROMOGENIC BACTERIA. I. Notes ON THE PIGMENT OF anima G. Thi HROMOGENES (WITH SIXTEEN FIGURES). Z. J/. Chamot and hiry 78 ON THE DIVISION OF ase CELL AND lites oes IN Live deeet XXIII), J. M. Van Hoo 394 R ARTICLES. W SPECIES OF eee stosTEOMA (WITH THREE TEXT rier M. W. — ‘ pete INTERNATIONAL BOTANICAL CONGRESS 403 ON THE FLorA oF ar pte AND Sounns AT Beavrort, N. Cc. Duncan s. 40) ON THE ———_— OF chagpaseertise EBENOIDES ASA Seas. William R. bees 410 : : J - 416 ELEMENTS OF PaLronorany. PLANT DISEASES. 7 ; So ee 419 : oe ‘FOR STUDENTS ; : - : ns . : - a tes of lead-. y if sabi must be ordered in advance of publication. Not less than " separate cat of thes actual c ted, of which 25 (without covers) will be fur: a ef ne; ” (with or ) Cove 3 if desired) to be tae for by the author. epara of pent pense ae will also be su plied cost. 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Home Office: John — P . Write for information. a UME XXX NUMBER 6 CELLS OF OSMUNDA REGALIS. IBUTIONS FROM THE HULL BOTANICAL LABORATORY. R. WILSON SMITH. (WITH PLATE XXII} | HE cytology of the vascular cryptogams has received scant Muon at the hands of botanists, as a glance at the literature *eris, and Pteris. Osterhout’s (11) article on Equisetum lorable as the first complete account of the formation of a = uncontrolled by directive spheres or centrosomes. . He ~ tat the appearance of the spindle was heralded by a felt- * of threads about the nucleus, and that the threads later €d themselves into a multipolar spindle, and afterwards ag characteristically bipolar. These three publications, ter with that by Shaw (12) on fertilization in Onoclea, and aa Belajeff (2, 3, 4) and Shaw (13) which treat of the Tinut : of the cilia of spermatozoids, make up almost the total of © esearch into the cytology of the pteridophytes since troduction of improved methods of technique. 361 362 BOTANICAL GAZETTE [DECEMBER Impelled by the meagerness of our knowledge, and by the want of accord among such papers as have been published on this subject, the writer two years ago began collecting and examining the sporangia of Lycopodium, Selaginella, and vari- ‘ous ferns, to discover some of the forms most favorable for cyto- logical investigation. None was found so good as Osmunda. All three of the species occurring in the northern United States were collected, but only O. vegalis was found to be fixed and pre- served in a condition suitable for the purpose, and it alone has received any considerable attention. For want of time a full examination of the reduction divisions was not possible, and therefore attention was centered only upon such features as it was hoped might be followed in detail in the time at disposal. Accordingly in this paper only the origin, structure, and fate of the achromatic spindle are to be considered, together with such comments upon the changes in the chromatin as seem necessary for clearness. A more extensive account of the chromatic ele- ments of the spindle is reserved for a future occasion, after there has been opportunity for further study. The methods of fixation, embedding, and staining require a brief mention. As fixing agents, chromo-acetic acid (1 per cent. chromic acid, 0.75 per cent. acetic acid), and Flemming’s woake solution were employed. Chloroform was used as the medium of transfer to paraffin. The sections were cut 5 or 10M in thick- ness. To differentiate the fibrillar structures most clearly, = ous combinations of the following stains were tried: nigrosi, Delafield’s haematoxylin, erythrosin, iron-alum-haematoxy!im, acid fuchsin, methyl-green, iodine-green, safranin, gentian-violet, orange G. Of these stains the two combinations which saith yielded the best results are iodine-green and acid-fuchsin, - was found especially advantageous in dealing with the — elements, and safranin and gentian-violet, which was mos effective in bringing out the details of the achromatic figur The sporangia of Osmunda make their first app the latter part of summer, and continue their growth 10 ae autumn months. By mid autumn those of O. cinnamomed & 1900] ACHROMATIC SPINDLE OF OSMUNDA 363 0. Claytomana have reached the mother cell stage, in which con- dition they lie dormant during the winter. The division into spores takes place in the spring, usually in northern Indiana between April 15 and. May 1. O. regalis does not reach so advanced a condition before winter as do the other species. Cell divisions in the sporogenous tissue occur in the spring, and itis not till after the middle of April that the mother cells are differentiated ; the division into spores is effected about three weeks later. ‘Amore exact statement will be instructive. April 6, material collected showed occasional karyokinetic figures in the sporangia. April 21, first collection in which no divisions were found in the ‘porogenous cells; the tapetal cells were in active multiplica- tion, May 5, many nuclei were in synapsis, the first seen. May 10,most of the nuclei were in synapsis, but some in first division. May 13, in most cases both divisions were completed, and often - trace of the spindle remained.’ This indicates that the rest- ng and early spirem stages last for about two weeks, and the ‘YMapsis stage for three or four days; and that the two divisions “eelfected in quick succession within a period of two or three days, The difficulty of obtaining the various stages of the second tivision also affords evidence that the second division follows . quickly after the first, and is completed in a much shorter time. © resting mother cell (jig. z) is angular in form, and Presents a rather large nucleus. The cytoplasm is evenly dis- : tibuted as a delicate network of stainable matter with irregular ‘Wellings at the points of intersection; it cannot be regarded as . mig the Structure of foam. The nucleus has a structure aly that of the cytoplasm, differing only in the seo a8 =a of its meshes, the larger size of its granular thicken "85 and its more intense reactions with stains. Two or three cleoli are usually present. In the cytoplasm are numerous a Staining granules of variable size, each usually lying a : ‘stained area, as if the granules had suffered contraction in | be added that the collection of May 13 was not from the same locality F earlier dates. 364 BOTANICAL GAZETTE [DECEMBER the process of fixation. Of the nature and function of these granules, which in staining properties resemble nucleoli, little was learned except that they are not starch. They are not peculiar to any phase of cell division, though rather more numerous in the resting mother cell, or the prophase. They cannot be extra-nuclear nucleoli, for there is no increase or dimi- nution in their size and number contemporaneous with the dis- solution or renewal of the nucleoli. Moreover, the nuclear membrane is yet unbroken, when, in the later prophase of divi- sion, the nucleoli disappear from view, and therefore, if the nucleolar substance passes out into the cytoplasm at this time, it must do so by osmosis, and any relation between it and the extra-nuclear granules must be conjectural. To the inquiry if they might not be the source, at least in part, of the material of the achromatic spindle, no satisfactory answer can be given; it may be so, but positive evidence is lacking. The mother cells enlarge for about two weeks after they have reached their full number, gradually separating from one another and becoming more and more rounded in outline. The nuclei also grow larger and pass first into a spirem condition 10 which the chromatic ribbon is not a single thread but much branched and anastomosed, and later into synapsis. The details of these changes need not be considered in this paper. Meanwhile, the cytoplasm has retained its reticulate arn But towards the end of synapsis a change is discernible, an this change is the first indication of the future spindle. It begins as an aggregation of material immediately about boas nucleus, causing this region to stain more deeply with gentian- violet. The structure of this accumulation of cytoplasm, a it is first recognizable, is so delicate as absolutely to baffle rs powers of the microscope; nothing definite can be said << regard to it, except that it appears to be very finely granu es especially in preparations fixed in Flemming’s solution. eet similar layer about the nucleus of the pollen mother pes Cobaea scandens, Lawson (8) proposed to give the name a karyoplasm, in virtue of its position. Since Strasburger § 1900] ACHROMATIC SPINDLE OF OSMUNDA Bia 5. kinoplasm has the advantage of more general acceptance, it will be employed here to designate not only this differentiated layer of cytoplasm but also the spindle forming material in all its modifications. As the kinoplasm increases in quantity, it begins to assume asomewhat definite outline and texture. It becomes distinctly granular, and the granules are often disposed in short rows which tun nearly concentric with the periphery of the nucleus, but are intertangled more or less confusedly (figs. 2, 4). In cells which have been fixed in chromo-acetic acid a more distinct fibrous appearance is presented. In some cases the fibers appear to form a loose mat; in others they are so related as to resemble a delicate meshwork with the meshes flattened towards the nucleus. Though the appearance is such as to suggest that the fibers and meshes are only a modification of the cytoplasmic reticulum, the writer was unable to trace the steps of such a transformation. The kinoplasmic material, both in its earliest form and in that assumed after cell division before its final re-transformation into reticulate cytoplasm, could not be distinguished as either retic- ular or fibrillar, but was very indefinitely granular. The reader will not fail to discover the similarity of these “onditions to those described by Osterhout (11), as the prepara- tory Steps of spindle formation in the spore mother cells of Equisetum, although in Equisetum no layer of granular matter Was observed. But the similarity stops here. The spindle in munda does not pass through a multipolar stage, nor is there at any time a zone of radiating fibers about the nucleus, such as were seen by Osterhout in Equisetum, and by Belajeff (1), Mot- - (9), Lawson (8), and others in the pollen mother cells of “atlous seed plants. Tripolar spindles, though occasionally met mith (fig. 7 0), were of so rare occurrence that their beginning or fate could not be traced, and it is certain they are not normal ‘ages in the development of the spindle. . The changes in the outline of the kinoplasmic mass ar aad and easy to follow. When it has become distinctly § i structure its fibers, all the time increasing in quantity at e very fibril- 366 BOTANICAL GAZETTE | DECEMBER the expense of the outer cytoplasm, begin to collect in greater abundance at opposite sides of the nucleus and to be pushed up into dome shaped prominences. This condition is very well shown in fig. 3. The threads of granules run nearly tangential to the nucleus, extending out some distance at the side, and it is obvious even at this time that the axis and the bipolarity of the spindle are already determined. It is rarely the case, however, that both poles are equally prominent from the begin- ning; commonly one develops considerably in advance of the other. A curious relation between the position of the chromatin in synapsis and the first formed pole of the spindle was observed so frequently that it cannot be regarded as acci- dental. In elongated cells in which the nucleus is situated excentrically, the first pole of the spindle is formed on that side of the nucleus where there is the greatest amount of cyto- plasm, while in synapsis the chromatin was almost invariably observed to be gathered within the nucleus to the side farthest from the center of the cell; that is, the chromatin in synapsis and the kinoplasm are gathered towards opposite ends of the cell. Such a relation between chromatin and kinoplasm may be true of all the mother cells, but it can be traced only in those cells which have the distinction of a long and a short axis, since the polar- ity of the spindle is not apparent until some time subsequent to synapsis. These peculiarities will be referred to again when the second division is considered. Every care was taken to avoid the illusion into which an observer might be misled by the effects of irregularity of infiltration of the fixing or other reagents, and the conclusion was reached that these relations have a real exist ence and significance in the living cell. The writer inadvertently had an excellent opportunity - examine a condition of the cell somewhat resembling SY ae, a Finding that the young sporangia, which are ensheathed ¥ 7 close covering of hairs, did not sink readily in the killing ee he first moistened some of them for a moment with om? They sank immediately, but afte: wards turned out to be ight unfit for study. It was perfectly easy to trace the path of 1900] ACHROMATIC SPINDLE OF OSMUNDA 367 invading alcohol. Both cytoplasm and nucleoplasm were pushed forward ; but no one could mistake such a condition for synapsis if he had once become acquainted with the latter. Contrary to what might be looked for, it was the resting cell which suffered most by this treatment; the spindle fibers, the chromosomes, and the spirem thread were not seriously affected. It will now be necessary to refer briefly to intranuclear changes. The chromatic material, emerging from synapsis, gradually unrolls and extends itself within the nuclear cavity into a much coiled spirem, apparently of one continuous thread, forno ends can be seen in uncut parts of the nucleus. The spitem shortens and thickens, and after a time is segmented into long irregular chromosomes, which continue the shortening and thickening process already begun in the spirem. It is easy to see that many of the chromosomes are split longitudinally into pairs, and the two parts of a pair are often twisted loosely about fach other. They take up a peripheral position in the nucleus, being apparently in close contact with the nuclear membrane, = halves usually remain attached to each other for a time, iving rise to Xs and Ys, or loops (as in fig. 5), according to the mode of attachment. But while the shortening is still going on, ‘ome of the pairs fall apart; otherwise how can the number of chromosomes shown in fig. 7 be accounted for, if Strasburger’s \15) estimate of twelve? be correct? This number has been ‘ounted in a few cases in uncut nuclei of the age shown in fig. 5. ad also in polar views of the late anaphase in which twelve ughter chromosomes have been seen symmetrically grouped tthe pole. But in nuclei of the condition shown in fig. 7 the Number of separate chromatin masses is quite variable, four- # aoe or sixteen being most common, and even as a _, osnty have been counted.3 If these numbers indicate a fa ing “Part of some of the chromosome pairs, there must be a ae | Sig reunion before their arrangement into such an ee “S that shown in fig. 8, in which of the twelve chromati | See Rote at the end of the paper. 3Guj 8nard counted as many as twenty-two. 368 BOTANICAL GAZETTE [DECEMBER masses eleven are clearly double. Whether in the case of the larger numbers all the groups represent pairs, or whether some of them are single chromosomes, has not been determined. It is by no means certain, however, that the number of chromo- somes in the equatorial plate is so constant as theories of nuclear division which are accepted at the present time require us to assume. There is a corresponding development of the spindle fibers coincident with the maturation of the chromosomes. The spindle acquires a more distinctly bipolar form, and its fibers, which become coarser and longer, run continuously from pole to pole. During the prophase they have the appearance of knotted cords or strings of loose beads. These features can be seen in fig. 5, and also in fig. 6, which represents a section of a cell cut suffi- ciently deep to remove one chromosome and part of the equato- rial region of the spindle. But in the metaphase and anaphase the knotted appearance is no longer recognizable. The fibers then appear as stout uniform threads or rods ( figs. 9, 70). The same statement can be made of the second division ( figs. 17 78); that is to say, the fibers at the time when they are aed tioning in the separation of the chromosomes are of uniform diameter and texture. They are rows of granules at all other times, either when they are disappearing or in process of forma- tion. It is evident that the achromatic spindle is wholly of plasmic origin. If any nuclear material takes part in its formation it can do so only after passing through the nuclear membrane There can be no direct union of linin or other nuclear substance with the kinoplasm, for in nuclei, as far advanced in the prophase as those shown in jigs. 5 and 7, the chromosomes are the ee Stainable constituent remaining; no fibers of any kind can made out, nor any trace of the nucleolus. : A comparison of the shape of the spindle, before and after the dissolution of the nuclear membrane gests that this period is marked by a sudden change- the nuclear membrane is still present, the spindle has - as seen just , sug While poles 1900] ACHROMATIC SPINDLE OF OSMUNDA 369 rounded and ill defined, and its breadth relatively great in com- parison with its length. But in all cells from which the nuclear membrane has disappeared, the spindle is seen to be consider- ably narrower, and at the same time longer and more sharply pointed. To explain these phenomena it seems necessary to suppose that a pressure, exerted by the spindle fibers upon the nucleus, is sustained by the nuclear membrane, and that when the membrane finally gives way, its collapse is attended by a sudden diminution of the diameter of the spindle ( figs. 7, 8) together with a corresponding increase in length. At the same time, the pressure of the fibers crowds the chromosomes close together. If the fibers be conceived of as curved elastic rods, this action becomes intelligible; that they are rigid enough to exert pressure is probable from the fact that the second spindles frequently cause a widening of the ends of the mother cell, Pushing out the wall so as to give it the form in section of a figure 8. One effect of the sudden collapse of the nucleus is that the chromosomes, which were all necessarily on the inner side of the spindle fibers, are most of them forced to the outside. No distinction of central and mantle fibers was possible, though Very great care was exercised at this point of the investigation. It is clear from fig. 9 that the original fibers, which run merid- lonally, still remain so after the separation of the chromosomes. €peated searching under a magnification of 2250 diameters failed to reveal any fibers other than those which run from pol ‘0 pole, or any thickening of the fibers towards the poles, as if ‘ome portion attached to the chromosome were undergoing con- fraction. How then are the chromosomes propelled ? Inasmuch °S there are no fibers discoverable by which they can be pulled, 'S it not possible that they have a power of motion in them- Selves ? pole This hypothesis is not absurd. It is quite as reasonable to ume an automobility of the chromosemes as @ contractility of ; ‘pindle fibers. The initiatory separation of the — 2 i + e : n mo Pairs by longitudinal fission implies a power of moveme 370 BOTANICAL GAZETTE | DECEMBER which is entirely independent of external tension; and the same inference is possible from the gradual shortening of the spirem to form the chromosomes, and from the re-expansion of the chromatin in the daughter nuclei. Meanwhile, the outer region of the cytoplasm loses its reticulate structure (still evident in figs. 3, 4), and passing through a con- dition in which it appears to be a diffusely staining mass becomes converted into granules, as in figs. 8 and g. This change, which is comparable to that preceding the organization of the kino- plasm into fibrillae, advances gradually from the spindle out- wards towards the cell wall. Directly after the metaphase the granules of this outer zone arrange themselves into rows stretch- ing from the poles out towards the equatorial region of the cell. This condition is shown in fig. g, and again in the second division in figs. 77 and 78. These rows of granules, which soon become more threadlike, will be spoken of as secondary fibers, to distin- guish them from the primary fibers that form the central frame- work of the spindle. They are not mantle fibers, for they do not enter into connection with the chromosomes, and are only just coming into existence when the latter begin to withdraw from the equatorial plate towards the poles. The secondary fibers multiply rapidly, and by the end of the anaphase fill all the outer part of the cell; in fact all the cytoplasm seems to have been exhausted in producing them. Almost immediately there begins a breaking down of the spindle fibers, indicated in the first place by their reassuming the appearance of dotted threads. In all cases the primary spindle is the first to disappear; then the secondary fibers break down and all the stainable cytoplasm is of granular texture ( figs: 71,13). Acell plate is first formed, which however comes to nothing ; it can be traced through figs. 12-18, 21. Though the second division follows quickly after the first, there is a sufficient interval between to allow the mere of a nuclear membrane and the partial reorganization of ee chromatin. The chromosomes after assuming a very eer cal pattern about the poles unite into a chromatic mass wht 1900] ACHROMATIC SPINDLE OF OSMUNDA 371 takes the shape of a shallow cup with the concavity towards the center of the cell. If there is any synapsis in the second division, it is repre- sented by the case of fig. 73. It will be interesting to compare this with the conditions prevailing at synapsis of the mother cell. In fig. 77, as in the synapsis formerly described, the chro- matin is bunched on the side of the cell most remote from the greater mass of kinoplasm. In this case it is easy to under- stand how these conditions have arisen from those preexisting. Whether the relation of the chromatin to the kinoplasm is deter- mined in the mother cell in the same manner, by the position of the last preceding spindle, it is impossible to say, om account of the long rest which the mother cell passes through. If it is so determined, it argues strongly for the organic continuity of the Kinoplasm, or at least of some specific substance which retains @ definite position in the cell and reacts on the cytoplasm so as 0 cause it, at the proper time, to be transformed into a fibrillar texture and organize the spindle. It should be remembered, however, that the first visible signs of approaching division in the mother cell are intranuclear, hence so far as we can judge ftom appearances, if the stimulus to division originates from ‘ome special region or structure, and not from the activity of the cell as a unit, we must assume that the primary impetus pro- feeds from the chromatin. Apparently the preparations within os, and those in the cytoplasm, “1 equal rapidity relatively to each other, with cases in which the second spindles are well formed, while the nuclear membrane is still unbroken, and others in which the ag membrane is gone and the chromosomes of ane second well organized, while the spindle is still very genes Vatiably, however, the development of the second spindle Siege by an accumulation of material along those eee : | © daughter nuclei which were in contact with the old spin e. : fibers first appear in this region also, and the spindle as *en in sections across this part of the cell is bipolar from the the nucleus for the sec- do not always progress for one may meet 372 BOTANICAL GAZETTE | DECEMBER beginning. The poles are not diametrically opposite each other, but both lie on one side of the nucleus, as shown in fig. 75". Invari- ably, too, the axes of the second spindle are at right angles to that of the first; that is, though they may be at any angle with reference to each other, they always lie in planes which are parallel to the first cell plate (figs. 75-18). In the preparations selected for drawing, the second spindles are either parallel (figs. 14-17, 21), or at right angles (fig. 78). It must not be understood, however, that these cases are typical in this respect ; they were chosen because in them both the spindles are in view at once. Undoubtedly the material used for the building up of the second spindle is obtained from the disintegration of the first, not as fibers, however, but as granules; and in this abundance of granular matter ready formed in the cell, we may perhaps see an explanation of the rapidity of the second division. he phenomena attending the metaphase of the second divisions agree completely with those of the first division. There is the same absence of mantle fibers (figs. 17; 18), the Same continuity of the fibers from pole to pole (fg. 1g), the same excess of fibers over the number of chromosomes (/igs. 18,19). In the anaphase there is the same elongation of the chromosomes into crooked or lobulated rods, the same beaded appearance of the fibers as soon as the chromosomes have passed to the poles (fig 2r), and the same development of secondary fibers which run from the poles towards the equator. = The beginning of the secondary fibers is shown in fig. ae they are more abundant and longer in figs. 20 and 2, In rt it is seen that those put out from the poles of one dee ee those from the other. The secondary fibers so meeting unite PY their ends into continuous threads, which connect the Sas of the primary spindles. In this way four secondary SP is : are formed; and thus the four daughter nuclei are now aca by six symmetrically placed spindles, of which two ae ers and four secondary. Meantime, there has been a slight gi ‘is tion of the primary spindles so that their planes, which were 1900] ACHROMATIC SPINDLE OF OSMUNDA 373 frst parallel to one another, come to lie at an angle of 60°. hhis results in placing the four daughter nuclei equidistant from me another in the tetrahedral arrangement which is character- stic of fern spores. Whether the rotation is brought about by the action of the secondary fibers, or by some other influence acting on the nuclei, could not be determined. In fig. 27 two primary spindles are shown, and some of the seconday fibers not jet united by their extremities. In jig. 22 are one primary spindle (that on the left side) and two secondary ; the axes of the other three spindles would make angles of 60° with these or with the plane of the paper. The disappearance of the spindle has been followed with stat care in order to discover, if possible, in what manner the kinoplasm is metamorphosed into ordinary cytoplasm. It was ‘ound that the middle fibers of all the spindles are the first to disintegrate, the material being used up in part to build the cell Mate. After the spindles are no longer distinguishable, a con- ‘derable amount of granular or amorphous matter remains on the inner side of each nucleus (jig. 24); finally this disappears and the cytoplasm assumes a reticulate structure throughout lis 25). All attempts to identify the granules of kinoplasm with microsomata, or the fibrous matter of the spindle with the Flop lasmic reticulations, were unsuccessful. The fibers of the ‘pindle are composed of modified cytoplasm, which in the ansformation loses its characteristic structure and becomes “structureless in appearance, then granular, then fibrous, and ‘ning to its normal condition reverses these steps. SUMMARY. ; The achromatic spindle originates wholly from cytoplasmic Materia] (kinoplasm) which accumulates about the nucleus in *YMapsis or spirem stage in the form of an indefinitely gran- mass of stainable matter. n- : The Kinoplasm becomes distinctly granular; then se Aig ‘ é ‘range themselves into short rows concentric wa Q ssed in Uclear membrane ; finally the rows of granules sr aR 374 BOTANICAL GAZETTE [DECEMBER greatest abundance on opposite sides of the nucleus, foreshadow- ing the development of a bipolar spindle. Usually one pole is formed considerably in advance of the other; and in cells cut parallel to their long axis, it can be seen that the first pole (the greatest accumulation of kinoplasm) is on the side of the nucleus remote from the chromatic mass of synapsis. _ The spindle is bipolar from the beginning. Némec’s (10) generalization, therefore, that sporogenous cells as compared with vegetative cells are characterized by their spindles passing through a multipolar phase, does not hold good of Osmunda. The fully formed spindle shows no distinction of central and mantle fibers, and no bodies which can be interpreted as centro- spheres ; all the fibers run from pole to pole. The dissolution of the nuclear membrane is attended by a sudden narrowing of the spindle anda corresponding increase in length. During the anaphase new (secondary) fibers, not to be con- founded with mantle fibers, are put forth about the poles and meet in the equatorial region of the cell. In the late anaphase the primary fibers, and soon after them the secondary fibers, begin to disintegrate, taking the appearance of beaded threads, and then of granules; at this time all of the stainable cytoplasm of the cell appears granular in texture. The spindles of the second division at first have their axes parallel to the first cell plate. They are constructed out of the granular products arising from the disintegration of the first spindle. The phenomena of the second spindles exactly repeat those of the first, except that four secondary spindles are formed by the union of the secondary fibers put forth during the anaphase. The primary spindles become rotated about each other so 4 to bring the four daughter nuclei into the tetrahedral arrangement. Cell plates are formed across the six spindles (two primary and four secondary), and in connection with them the separating walls of the spores are laid down. 1900] ACHROMATIC SPINDLE OF OSMUNDA 375 Such a relation between the fibrillae of the kinoplasm and the cytoplasmic reticulum as Blackman (§) reports in Pinus, and Lawson (8) in Codaea scandens, could not be verified. Between well developed spindle and cytoplasm are the three stages, (1) dotted fibers, (2) granules, (3) amorphous kinoplasm (structure too delicate for the microscope to reveal). The same phases in reverse order were traced in the first formation of the spindle. This investigation was conducted in the Hull Botanical laboratory of the University of Chicago during the spring and summer of 1899. The writer, while assuming full responsibility for the views expressed, takes pleasure in acknowledging his indebtedness to the members of the Botanical Staff for their courtesy and encouragement, and especially to Dr. Bradley M. Davis, under whose more immediate direction the work was undertaken, +" Since the foregoing account was written (in February 1900), a com- pehensive work on karyokinetic problems (15a) has been issued by Stras- burger, in which, among other topics, he discusses the divisions of the spore other cells of Osmunda, and the general formation of achromatic spindles “plants. He distinguishes two types of spindles, those possessing ened “mes and those without such controlling centers. The latter, which are ‘racteristic of higher plants, are again subdivided into multipolar polyarch ‘pindles, such as those of the spore mother cells of Equisetum, and multi- tar diarch spindles, such as are common in various spermatophyte root- "8S; and he see § the same origin and reaction to stains, iz and mantle fibers, but supporting and attracting fibers pi oo). He pronounces against a power of movement in the af “toth oo Selves, and attributes their withdrawal from the equatorial 2 . : = _ of the attraction fibers. This conclusion is quite eee ne “is use t expressed in the preceding pages. To the view that the ae 'P to help complete the achromatic spindle, my observations, t — Not “ : too “ontradictory, are not altogether favorable; the spindle nee 376 BOTANICAL GAZETTE [DECEMBER nearly complete before the disappearance of the nucleolus. In his renewed observations on Osmunda, Strasburger has noted the large and varying num- ber (20-22) of chromatic groups in the spore mother cell. Each such group he regards as a chromosome pair. He finds the number of chromosomes in the prothallial cells is not so constantly twelve as he formerly (15) stated ; it may reach sixteen or more. It is therefore certain that the current view as to the constancy of chromosome numbers cannot be maintained, at least as regards Osmunda. MCMASTER UNIVERSITY, Toronto, Canada. LITERATURE CITED. 1. BELAJEFF, W.: Zur Kenntniss der Karyokinese bei den Pflanzen. Flora 79: 430-442. 1894. : Ueber den Nebenkern in Spermatogenen Zellen und der Sperma- togenese bei den Farnkrautern. Ber. d. Deutsch. Bot. Gesells. 15 337-339. 189 3. ———: Ueber Spermatogenese bei den Schactithalnee Ber. d. Deutsch. Bot. Gesells. 15 : 339-342, 1897. : Ueber die Cilienbildner in den Spermatogenen Zellen. Ber. d. Deutsch. Bot. Gesells. 16: 140-144. 8. 5. BLACKMAN, V. H.: On the cytological features of fertilization and related phenomena in Pinus sylvestris. Phil. Trans. Roy. Soc. B. 190: 395- 426. 1898. 6. CALKINS, G. N.: Chromatin reduction and tetrad formation in pteri- dophytes. Bull. Torr, Bot. Club 24: 101-116, 1897. 4. GUIGNARD, L.: Le développement du pollen et la réduction chromatique dans de Naias major. Arch. d’Anat. Microscopique 2: 455-599 1899. 8. Lawson, A. A.: Development of the karyokinetic spindle in the pollen mother cells of Cobaca scandens. Proc. Cal. Acad. Sci. Bot. 1:169. 1898. : Origin of the cones of the multipolar spindle in Gladiolus. Bor. GAZ. 30: 145-152. Ig00. 9. MorTTiger, D. M.:: be zur Kenntniss der Kernthei Pollenmutterzellen einiger Dikotylen und Monokotylen. Ja hrb. wiss. Bot. 30: 169-204. 1897. toa. NEMEC, B.: Ueber die Ausbildung der achromatischen Kerib figur im vegetativen und Fortpflanzungs Gewebe der hdheren Pflan zen. Bot, Centralbl. 74: 1-4. 1898. ear ering der Kern und Zelltheilung. 77: 241-251. 4. 8a, lung in ae Bot. Centralbl. 10b. BOTANICAL CAZETTE, XXX. + Vetoeid. Lith Anst.7 BAPun ke LeIpeS SMITH on AGHROMATIG SPINDLE. : 1900] ACHROMATIC SPINDLE OF OSMUNDA 377 1. OSTERHOUT, W. J. V.: Ueber Entstehung der karyokinetischen Spindle bei Equisetum. Jahrb. f. wiss. Bot. 30: 159-168. 1897. 12, SHAW, W. R.: The fertilization of Onoclea. Ann. Bot. 12:161-285. 1898. 13. -: Ueber die Blepharoplasten bei Onoclea und Marsilia. Ber. d. Deutsch. Bot. Gesells. 16:177-184. 1898. 14. STEVENS, W. C.: Ueber Chromosomentheilung bei der Sporenbildung der Farne. Ber. d. Deutsch. Bot. Gesells. 16: 261-265. 1898. 15, STRASBURGER, E.: The periodic reduction of the number of chromo- somes in the life history of living organisms. Ann. Bot. 8: 281-316. 1894. See also Biol. Centralbl. 14: ——. 1894. ———: Ueber Reduktionstheilung, Spindelbildung, Centrosomen, und Cilienbildung im Pflanzenreich. Jena. 1900. EXPLANATION OF PLATE XXII. The figures were outlined by the aid of an Abbé camera under the mag- hification given by a Zeiss apochromatic oil immersion lens 2™™ aper. 1.30, in combination with a Zeiss compensation ocular no. 8. In all cases the preparations were studied with the higher oculars, 12 and 18. Fic. 1, Mother cell with resting nucleus. Fic. 2. Mother cell with nucleus in spirem stage, and the kinoplasm dis- Unctly granular and fibrillar. IG. 3. The same, somewhat more advanced, showing the bipolarity of the kinoplasmic mass. 1G. 4, A section which cuts off only a portion of the nucleus. Fic. 5. Chromosome stage. FIG. 6. Portion of a cell, showing part of the equatorial r spindle, egion of the Fig. 7. Later chromosome stage. Fig. 8. Chromosomes in equatorial plate after the disappearance of the nuclear membrane. Fig. 9. Anaphase, showing continuity of the primary fibers from pole to pole, and secondary fibers extending out from the poles. FIG. 10. A tripolar spindle. Fig. imary spindles on the left side; 1G. 22. Late anaphase: one of the pr : : d out of the secondary fibers. the two “two other spindles shown have been forme 1GS. 23-25. Successive telophases. STUDIES ON CHROMOGENIC BACTERIA. I. NOTES ON THE PIGMENT OF BACILLUS POLYCHROMOGENES. E. M.CHAMOT and G. THIRY. (WITH SIXTEEN FIGURES) Tue Bacillus polychromogenes was first isolated from a well water of Nancy by Macé? in 1894, and since its first discovery he has met with this same organism on five different occasions in well and conduit waters of that city. Two years after its discovery it was again isolated from the same well, and was found to possess the same characters as in the previous case. The organism was then described by one of us under the name of Bacille poly- chrome.? These six colonies, found at different times, have varied neither’ in the original colonies nor on subsequent cultivation ; varieties are, therefore, still unknown. Neither has it been possible to obtain variation by culture methods, for although one of the original colonies has been grown in the laboratory since 1894, part of the time in America, no change has been observed. It has also been impossible to obtain a non-chromogenic variety = spite of all attempts. It seems more than probable that this beautiful species will be met with by other investigators, and therefore, although the present article has to deal with the pigment, a few words regarding the characteristic features of the bacillus may not be out of place. The B. polychromogenes was so named because of its power of giving a multiplicity of colors on ordinary culture media. On such media the organism produces at times blue, at peculiar de *Mack, E.: Traité pratique de Bactériologie. Ed. 3. 849-852. 1897: Atlas Bactériologie, f/ 29. ; iét *TuIRY, G.: Sur une bacterie produisant plusieures couleurs. C. R. de la Soc! de Biologie, 7 Nov. 1896. ee Contribution a l'étude du polychromisme bactérien. Archives d. physiol. ¥-9 284-289. 1897. 9. 1897 oe [DECEMBER 1900] CHROMOGENIC BACTERIA 379 others red, green, violet, purple, or yellow. These colors, as will be shown, can generally be controlled, thus permitting one to obtain at least part of the colors with certainty. This interesting fact las already been pointed out by the authors.3 In addition to the pigment described below, colored insoluble microscopic crystals are also generally found on solid media. These crystalline aggregates are made up of irregular radiating clusters of fine needles of a deep blue color, and are probably not due toa crystal- lization of the pigment, but to crystals of some other substance stained by it. The composition of these crystals will be dis- cussed in a future communication. ; The organism liquefies gelatin, solidified blood serum, albu- min (white of egg), fibrin, etc. In peptone solutions no indol is | produced, neither does the bacillus produce gas in fermentation tubes filled with glucose or lactose peptone solutions at room femperature nor at incubator temperature. The bacilli themselves are generally colorless, rarely they appear red or blue; in some of these latter cases the organism i uniformly stained, at others times only minute intra-bacillary slanules are colored. The bacillus is polymorphic; not only does its form vary according to the nutritive media employ ed, but often there is great variation in form observed in different Portions of the same medium. It has not been possible thus far to obtain with any degree of certainty a constant form on any culture medium, not even when employing one of definite com- Position (7. ¢., a medium in which the source of nitrogen 1s eet Peete etc. but a chemical compound of known composition Such as asparagin or other bodies). In general the organism “sumes the form of a short rod rounded at the ends, at other limes it is spherical; again, long curved giant forms with swollen “tds are seen. The bacilli are sometimes isolated, oogenenarn stouped (diplo-bacilli, chains). Coccus forms are ee of of a = staphylo- and diplo- forms, or as tetrads and chal “ight cells 3 ator, E. et Tuiry, G.: Bacille polychrome. * Communication 4 la Réunion Biologique de Nancy, Cultures et Spectre du Pig- Feb. 1898. 380 BOTANICAL GAZETTE [ DECEMBER Usually the organism is motile, but the motion is always slow. Motility is to be seen in colored as well as in colorless individuals. It stains well by the ordinary methods and retains the stains by the method of Gram and that of Claudius. Greater morphological details would take us beyond the scope of the present note, namely, an account of some of the results of the study of the pigment produced by this bacillus on potatoes and on gelatin. Growth on potatoes and the pigment formed.—On a medium as variable in composition as potatoes of different varieties and different ages of growth, it might be expected that there would be considerable variation in the colors of the pigment and nature of the growth of a chromogenic organism. The B. polychromo- genes shows the effect of such changes in a most marked manner. Potatoes upon which this organism grows are colored variously yellow, greenish, red, violet, blue; the last color predominating but not constant. It was soon found, however, that a beautiful deep blue could be obtained, almost without fail, if the potatoes were first soaked in a dilute solution of sodium hydroxid (0.25 per cent. to 0.50 per cent.) containing a little calcium phosphate for twenty-four hours or less, depending upon the thickness of the pieces. Since this medium has served as the basis for the isolation of the pigment, and is being constantly employed by us in the study of the pigments of other chromogenes, it may not be out of place to describe our methods of preparation. As large tubers as possible are chosen, such as are known to become mealy and porous on boiling. They are well washed in cold water, using a brush to aid in cleansing, and are dropped into boiling water with the skins on, and boiled till just cooked through. The water is then poured off, the potatoes allowed to cool somewhat, pealed, cut into slices 1 to 2™ thick, and dropped into a dilute solution of sodium hydroxid, where they remain about eighteen hours. The supernatant liquid is then poured off, the slices drained and transferred to glass boxes tole aa o diameter and 49 to 50™ deep, with loosely fitting covers (in other words deep Petri dishes), a little water is added, and the 1900] | 900 CHROMOGENIC BACTERIA 38 I _ medium sterili . ee an eat days in succession in streaming steam, ipon such a piece of ure of B. polychromogenes is inoculated "produced, which SE sae a deep blue very soluble pigment the medium, colori iffuses slowly through the whole mass of into fifteen da iyi the latter an intense indigo blue, In from lass, and has aerate matter has penetrated the entire ee ot Dito 2 it uniformly and so intensely that the potato, the more oa almost black. The more porous the the pigment is panei 3 and uniformly is it colored, since the color remains a ieee Sbsenee of air. For some time it purple, and th the — intensity, then it becomes violet longer able to aie OWlns either to the organism being no the various as « pigment to replace that being reduced by the production 2 substances present in the potato, or owing color begins to ee reducing agents by the culture itself, the Ridly, the color Decolorization proceeds more and more s being “s aa the surface of the medium exposed to the ‘Boumes a dirty oe disappear. The culture medium finally Mo thin slices and eee Generally, cutting up the potato tction of a bl SEponne them to the air leads to the pro- ‘00 old, ue again by oxidation, providing the culture is not quite soluble in The bi hi, ue pigment is very soluble in water, her, chloroform, - Mlle alc Yenzine, = a in strong alcohol, in et ‘Ttnately extractio ef therefore, ‘5 the best solvent; but unfor- reducing Dcian n with water removes such an amount of. “ohol, the str ces that it was found necessary to employ dilute Mesent in th ength of the latter varying with the moisture | e culture to be extracted. In or s cut i socpassesages the pigment, the piece of culture medium der that n slices and exposed to the air for a short time in formed, “a much of the blue pigment as Po : OWed to . alcohol is then poured ovet 7 Fed off, and or some hours, the blue alco alli aarenetebater of the solvent is ong as any coloring matter can be re holic solution 1s made. This moved. The 382 BOTANICAL GAZETTE [| DECEMBER alcoholic solution of the pigment is then filtered through a care- fully cleaned bacterial filter (Chamberland, Kitasato, d’Arsonval, or others). The first portions of the filtrate are rejected, owing to the fact that a chemical action takes place at first, doubtless owing to air present in the pores of the tubes; the result is the production of a beautiful purple-red liquid. » After some 50%, more or less, have passed, the filtrate passes unchanged in color. It is then evaporated to a syrupy consistence at 50° to 60°C,; a higher temperature leads to reduction and decomposition of the pigment at this stage, owing to the presence of large amounts of sugars extracted from the potatoes. The thick deep-blue liquor is precipitated with strong alcohol (about 98 per cent.), the supernatant liquid poured off, and the precipitated pigment dis- solved in a very small amount of distilled water and again repre- cipitated with alcohol. This process is repeated as long as the alcohol seems to extract anything from the pigment. At this point the pigment is precipitated in such a finely divided condi- tion that it refuses to settle completely, and cannot be retained on filter paper ; a very small bacterial filter is therefore employed for the separations. The material is then carefully removed from the filter tube and dried. This dry amorphous powder has a grayish-blue color, and is completely soluble in water to 4 beautiful pure-blue color. Although we have reason to believe ‘that it is still impure, the amount of impurity is doubtless ° small that the pigment thus separated can be used as the basis of comparison, and for the reactions given in this paper. It can not be made to crystallize, and is insoluble in all ordinary sol- vents, such as ether, petroleum ether, benzene, chloroform, amy! alcohol, etc. oe If the blue aqueous solution is treated with a trace of acid, the blue is changed to a violet, a trifle more acid leads to = production of a beautiful purple (royal purple). An excess 0 acid gives rise to a red with more or less of a purple tint. = sensitive is the compound to acids that carbon dioxid cause a change of color. It was at first thought that when se? acids were employed, a color change resulted which was differet £900 ] CHROMOGENIC BACTERIA 383 from that produced by inorganic acids. Later experiments seem to indicate that this is more probably due to a difference of intensity of action. Ammonium hydroxid causes a change similar to that pro- duced by acids, but the red color in this case is of a different tint from that obtained with the latter. The addition of acids to the purple-red ammoniacal solution restores the blue color, but if the acids are added in great excess the red tint of acid solutions results. It is worthy of note that a decided excess is necessary. Fixed alkalies (potassium, sodium, barium hydroxids), in small amount, first produce a violet tint; if a little more of the teagent is added a pure blue results, but the color is somewhat paler than that of the original solution. When added in excess, the fixed alkalies give rise to a grass green solution. When the pigment has not been carefully purified, or when filtrates directly from a culture are employed, the change to green is much more tapid, and the amount of alkali required for its production is less. If the pigment is quite free from foreign bodies the green is rather persistent, but when impure rapidly fades away, leaving a yellowish liquid. The process of decolorization begins at the bottom and gradually extends upwards until the surface is teached; here, being in contact with the oxygen of the air, the color persists. If the yellowish alkaline liquid be shaken with air it immediately turns green, then blue; and if the agitation be continued there results a blue solution of almost the same inten- Sity as the original. Allowed to stand undisturbed a reverse change is observed, namely, rapid decolorization passing through a green. The blue can be restored even after several days by shaking with air, The addition of alcohol to the yellow- ish solution produces a dirty yellow precipitate which ae blue the instant it comes in contact with oxygen. This phe- Momenon explains why it is that porous potatoes pce mee Pigment, and why cutting the colored medium into thin and €xposing it to air before extracting, gives 4 larger quantity © the coloring matter; for it seems to be obvious that we have to do here with a case of oxidation. 384 BOTANICAL GAZETTE [DECEMBER Fixed alkalies added to the red solution resulting from the action of acids on the blue first restore the blue color, then, if in excess, produce a green, which in turn disappears as has just been described; and in like manner shaking with air or addition of hydrogen peroxid restores the blue. From neither acid nor from alkaline solution will solvents such as petroleum ether, ether, chloroform, benzene, amyl alcohol, etc., extract any color- ing matter. = As to the important question whether the organism produces the blue pigment or a compound which turns blue in air, the writers do not yet feel justified in advancing an opinion. When the clear blue solution (obtained by dissolving in water the coloring matter isolated from potatoes by the method described above) is placed before the spectroscope, a fairly well defined absorption band is seen in the neighborhood of the Dline. The maximum intensity of this absorption band has approxi- mately a wave-length of %=594, in the case of the purest pig- ment thus far obtained ; its width and intensity varies, naturally, with the concentration and thickness of layer of the solution examined. There is also a slight darkening of the spectrum in the red, and a similar cutting off in the blue extending to the far violet. In the red this absorption seems to begin somewhere from A=680 to A= 690, but is so gradual that no satisfactory measurements can be made. In the green, blue, and violet the increasing absorption is so gradual that no reliable decision can be made as to just where the absorption begins. Fig. I gives the absorption spectrum of solutions 10™ thick, containing i liter of the purest pigment obtained; fig. 2, same solutions 1n layers 25™™ thick; fig. 3, same pigment in solutions of “ae wed liter, examined in layers 10™™ thick. In 25™™ layers the absorp: tion bands of these last solutions are too intense to permit of their being represented on the same scale as those figured. Blue solutions obtained by mere filtration of water extracts of colored potatoes usually show more marked absorption in the red an violet ends of the spectrum than do solutions of the pigment separated as previously described. Fig. 76 shows the spectrum Sent eae NG TRO TEES Ee ee ee eS i ane epi ee Oe NX + personal 144, 216, 360 st A. 418 work of 71, 141. Bes Bicknell E, work of 7, 210 oodle, L. A. Borraginacez, Giirke on 135 Botanical Society of seabion 216; annual meeting of fee ea pomeasng work of 71, 210 Bray, W of 428 of travel 125 2 work of 135 , N. L., personal 430;. work of 71, vi Brom nygirne 135 Brown f Unive aaity, bequest to 431 eyes Segoe ow ae sich capil- lar Coville: boa ‘ombuliforme a. nen 20; naviculare 20, 123; pecuaeniage: trum 2 Berne ae on 71 Burgess, E. S., work of 431 Busse, Walter, personal 142 a Cactacee, cna er of ted Cactex, Bran Caesalpinia corr, patie iis California a, Congdon on pits of 135 Callirrhoé alceo tee 182 rtical 205 Camptos =e “rhizophylus x Asplenium — Cam s, E gies Wok of 405 Creuset dic Capillitium, formation of BLT Cardam hg i 130 ! 63 Carleton, M. A., personal 216, 288 Caryophylaces of Mexico 0 58 Carpinus Caro) Castillo, D. del, work of, 405 heailiets, Piper 0 on 210; Suksdorf on 425 Catena, Chodat on 135 BOTANICAL GAZETTE [ DECEMBER Cedar, von Schrenk on diseases of 429 Cell aivision in raigo+ arians 217 Cell se e, development and function of 73; Celtis oni, Hill on 424 Cephalotaxus edie ee: Arnoldi on 426 Cerastium sor Nate Carieasns on papier of 288; Swingle on breeding of 2 Cereus enneacanthus, roots of 350; gigan- teus, roots of 34 Ceterach one x Phyllitis scolo- pendrium a apie c, J., 66, 138, 140, 208, 211, Characee, Goetz on egg i ay gas has: of pollen tube. Lidtoles on hiidnodiin Botrys 406 Chevalier, A., work of 405 China, flora of centtal: Diels opsidiaan f Ab. Cohnii 101; inches 107; oo, I Chisrecade Cohnii 100; sarcophyci 103 Chodat, R., el of 135, 405 Ch rchill, J. R., personal 142 Chrysopsis, Pollard on 8s Chytri idi ale > Gobi o n 426 Cissus anh 40 adop gee ae dene: 164 +» per Col Collettrichum gosypil i Collins, J. F., perso’ 143 Composite, double Totiberee in 252; Hieronymus on 135 Congdon, J. W., aes 135 Congress, International "Ee tanical 40 Coniferae, Worsdell on ovules ‘of 13 Conti, Pad eork of 424 : Contributor s: Atkinson, G. Bar 78 B. 57; Cou Coulee, IM ee at. eine 138, 130, 140, 141, 209, 210, 212, 213, 215, 270, 280, 282, 285, 287, 354, 418, 419, 420, 421, 422, 425, 427, 429; Cowell, John F. 348; Cowles, Henry C, 355, 417, 28, 429; Cranefield, » 131; Dayis, Bradley M. 139, 427; arcege i M. W. 403; Fairchild, D ; Harper, R. . 217; Hefferan, lane wae Holzinger, John M. 125; pase Duncan S. I, 410; Kraemer, Henr G ace 206; Moore, George eed 117; Nelson n, n, Elias 122; Perrot < 403; ion, ‘Gitean 3575 Rimbach, August ; Robertson, Charles Sch » K. 359; Smith, R. Wilson ars Thériot, I, 12; Thiry, G. 378: Timber- lake, H. G. 73, 154; Van Hook, J. M. Tg ‘Waugh, F, A. 206; Wiegand, Karl °o o Aven 64, see 25 EE ceailaria majalis 25 ok Coreopsis aristosa 130 lan vdalis cava 9; nobilis 186; ochroleuca seca’ s “La nature tropicale” 354 Cunlter, J. G. 70 Coulter, J. a 67, 71, 133, 136, 138, 139, 140, 141, 209, 210, 212, 213, 215, 279, 280, 282, 285, 287, ie ey 419, 420, 421, 422, 425, ih Coulter, S. E, A Coulter. Stanley, ‘6 aislages of plants of ra 21 res 2 pe ca 360 Cowell, Tein Co wles, Henry od 355,417, 428, 429; per- Ba ctela I rataegus pefeai is 323; Alle iensis 3 Michauxi 342, 343; Pinetorum 343; ; otendiiniie 345; ru- sororia straminea 345; Cremer, M., work of 211 Crown n-ga all, Toumey on 213 Cru ruciferae, insects and 130 ~ INDEX TO VOLUME XXX la 344; Sargenti 339; senta 341; 336; venusta rena of Mexico sng uciflorae, Martel o Cpa afin flex 195; multi- pon t Paisaiooal tot 195; plese 195 iyineaas of W yoming, Nelson on 434 ycads, Lang on 2 Cystopus candidus, W ager ein I, 25, 73, 145, ote sty, 252, I, 39 Gime F., work of 287, D pre Torre, C. G.,and Harms, H. “Ge- 279 a Si iphonogamarum ” 67, Ganesan wor . a 405 Dasya eleg Davis, Bradley w 139, hd personal 142 Davis, C. A., work of 4 Davis, K. C., work of 82 Day, Dav id F isher, aerpent of 347 Deane, Henry, work of 2 te geri Carvin besiaas 182; Davis 282 : occidentale, Ramaley on 282 cul ophagus Ene ou on 2 ae bulbifera Laisemonies oy wor ve Desm Sin rteri a : Scvtie 16; cratiliond DeVries, fees k of 212, 405 pee tatesertek Dlpapicem 13; __ pelluci- dum een: — 14; Howei15; laxi- retis 14; vari iieanaeeiiia, hinvicts 123; subcom- 0 122 Dicranum, _—— cased alustre 153 ries phora Ravenel Scofield on 282 Dicyota dichoto: Diels, L., wor we ot m 15: oe AS: 405 ; Diseases, anthracnose :. pat 48; 0 forest trees, conjuse n 288 , M. W. 40 103, aoe “Resin ducts and —- cells of Abies and Picea” 418 grey montana 189; Ye Dryopteris “etn x marginalis 414 alia B. M., personal 288, 359 436 BOTANICAL GAZETTE Durand, work of 424 Dutailly, work of 405 “Earth and Air” 288 Eaton, Daniel C., a 142 Pe tao siliqulos $s 409 Echino = Wislizeni, roots of 349 Beology, tery on 428; Jones on 424; Ule 28 tus n Characez, Goetz on 214 Eleutera, Stuntz on 71 Embryolog gy of Taxus baccata, Jager on 357 agree Nathanson on aioli Manes 138; of Sg lig pellucida 1; of Canna Indica 39; of Convallaria majalis 25 ; of Potamogeton as lg 31 Endosperm o of Peperomia pellucida Engler, A., “ Das paanventeiok i ; Me Die natiirlichen ee nzenfa alien” 279, 358, 420 nographien _afri- oe. Hiatoentmien 419; work of 135, 2 Enzymes of sesiet, Cremer on 211 Eri nee acris a preri nS 3; Droebachensis 198: gla s mens seeleieeut cae cna eus 199; i rtus 198; _Philadelphicusy double fertiliza- tion in 254; Piper on 210; strigosus Soable cei in 254; trifidus 63; Yellowston Eryngium, Britten sc Baker on 135 Erysiphez, Freeman o innesota 282 Erythronium, iniden- 176, 180; mesa- choreum 180 Eryt throtrichia safe glaberrimum on: ebpiouenss Wy- racillimu oink re 194; mingense I oe pega 73 Evans’ “ Alaskan Hepaticae ” 419; work cits: Deane and Maiden on 210 Euphe bin norpponitolts 407 F Fagus, Diels on 424 rasa D. G. 130 Fax n, Edw in, personal 142 Fem hybrid Fern i a +» personal 142; work of 13 Fertilization, double in Caltha palustris, Thomas on 425; in Compositae, double 252; of Batrachospermum, Osterhout on 139 [ DECEMBER Sais serge tc Museum Mi ; Her n, work of 2 . Pucidel, sepotdan gymnan ris 123; gymnandrus 16; rabbenibatie ‘bunt 16 jahault, work of 404, 405 aes Gillot on local 405; of southeast? Minnesota, Wheeler 0 282; of Lake Nyassa, Engler on “ss Flowers pier inse 130 seen of the United States, Gannett on Forestry, ipa re of 430 Fo Noll Formic aldeh ries pte on a Iton on Fox, Wm. J., personal ne ork of 71 n, E- Fuitillasia Semaoieciath; cell plate of 83; Meleagris 175; tenella, double fertili- lization in 252 Fritsch, Karl, work of 425 Fuligo septica, nuclear division in 217, 2 Fungi, Hennings on African 135; Patter- son on 7I G Galloway, B. T., personal 143, 431! Galls, Kiister on anatomy of 140; origin of tannin in 2 annett, Hen oe f 215 Garden, Missouri Botadii cal 143; Journal of the New York Botanica Gaurella oot ; guttulata 351 Gauropsis 351; canescens 351; guttulata 351 Gelidium crinale 409 Gentiana, cruciata 185; Pollard on 70; uberula 186 penn a. Noll on 134 Gerber, w of 405 ak ations nd light, bgerpeerd on 215 Gesneriacez, Fritsch 0 Geum, Dutail on ee Gidon, Ginkgo, Sew and Gowan on 139 Gladiolus, ralkipules asia? in 145 __ Gleos sagen fructigen 2; malicor- ticis 48, 57; serra §2. Graphaitwan thermale 121; Wrightii 122 Gobi, work of 426, 427 ee a ne 1900 | ‘Goebel’s “ ae = of plants” Goetz, Geo P ‘ au x0 y> ersonal 430 Griffiths, David, personal Grimmia, alpest pas a ore 19; mollis 123; mo montana Idahensis 18 ; pseudo sonetage 18; subsulcata 18, G saaetia squarrosa 182 Grinnellia Americana ie of lower pects eo on 422 ( a G rk of a 5 , Arnoldi on 426 : jymnostomum, curvirostre commutatum ; curvirostre scabrum 12; microsto- Wimmerianum mum 13; murale 13; 13 H Halacsy’s “ Conspectus Florae Graecae”’ Halsted, < PG e Joe - ae Harper, 17; person 6, 360 Hartig’ s is “Lehrbuch der Seated OES heiten”’ 417 Heckeria senbryo = of 9 Hefferan, Mary 2 tb aelaag bi re 215 ceo : annuus 257; Nawaschin on tis rimus 186 Heller's * . “Cato af North American Plants 43 Hemerocallis “ula, cell pe of 83 Henderson, L. ork of 135 Hennings "P., pa os 135 pee Evans on Alaskan phani on 70, 135 ae Castillo, Flahault, and Moulle- 425; Ste- farin on 4 Merbariam, oF H.N. Patterson 144; Rocky in 18 9 Herbe. perennial 171 Heuchera ovaltiolia 118; saeco 117 Se Chodat on 135 Hollick, C. A., personal 216, 360 Holzinger, John M. 125; work of 425 INDEX TO VOLUME XXX , 430 : 41S: 18; oe nee 437 Hottes, C. F., pager 143 Hua, work 0 » 405 Huber, on A onlans flora 405; work of 136 Hunn and Ba cae “ Amateur’s practical garden book” Hy, work of 405 Heaciithae. candicans 176; orientalis, cell late of , Hyatt, James, tal Hybrids, Asplen m < Camp- s eizophys ney "Léveill and tds Shibata n 42 Hymenophyllace, Boodle on 427 Hymenophytum, 5 ni on 70; ee tum 12 crostomum 13; muraie 13 Hypnea musciformis 409 Hypnum, a m 124; Cardoti 124; cupressiforme resupinatum 23% fluitans brachydietyon Holzingeri 124; Halle 22; molle Schi perianum 23; ochra ceum uncinatum 124; pallescens agi polygamu 4; resupinatum 23; Schim- perianum 23 stellatum t 124; uncinatum 21; uncinatum subjulaceum Holzingeri 5 Hypoxis hirsuta 175 I Idaho plants, Henderson 0 lex, Cassine 406; Lane ie opaca 407 Infusoria, Yasuda I e of 83 nsects, flowers aig 130 Iris versicolor, cl kage “Tentamen Florae Ito an atsum Lutchuensis ” i J Jager, L., work of 357 elite’ s* oom of the flora of Long johannes ; * Das Aether - Verfahren ” jokneee, Duncan 5. I, 419; work of 208 _E., work of 424 Jun uncus Juni ards pate 406, 407 K eeler’s “ a conti trees” I Fy ates Ke el work of of I "G. G., cite ae Keanedy, Pp. B., personal 142 438 BOTANICAL GAZETTE King, Cyrus A., personal 359 ie Britton on oe of 405 Know » work of 141 Kau s Handbath der Bliithenbiologie”’ 143 peeleng work of 286 i Kuhnisteta, candida 182; t cones 182 Kuntze, Otto, w ne of 358 Kiister, work eye L Labiatae, Giirke on 135 Laboratory table, Roitowlew on 21 Lacinaria, ee 182; squarrosa 183 “pe Wij. G.2 £wWerk, ok of 212 1S.fh. death of 359 arix, Americana, cell plate of 83; Euro- paea, cell plate of 83 Larkspur, Wheeler on 28 Lawson, Anstruther A. 145, personal 72 Leaves, Clos on floral 4 ioe Lemmermannia, Chodat 135 Linnaeus’s Systema Rakeisé: Britten on 2 Lenticels, Devaux on Lepidium, apetalum 190; pubicarpum Leskea pas Lesquerevia pap nia, fuscescens asi. littoralis 318; 21 oe and germination, ier og on 215; and eg Palladine on 21 Lilium, longiflorum, cell a ou ion in 252; Philadelphicum, double fe rtiliza- ion i perbum 181; Live ihe bie of the cell and nu- son eet Lloyd, of 71 ee ade “ rhe of North America”’ 143 se Livingston, er E. 285, 289, 424; per- | DECEMBER Loesener, th, ier of 135 Loew, E., Person 143 bases cod 359; work of 214 poe W. #H., ig of 429 Longyear, B. O., k of 4 Lotsy’s “ "Alkaloids i in Cinchons ” 279 Lounsberry’s “A guide to the trees” 132 Lupinus, humicola Tetonensis 120; Pol- Lycopodium, Lloyd and Underwood on Lythracez, Koehne on 424 M Maceration fluid, abate: on 215 Macfarlane, J. M., personal 143 Machaeranthera, superba 197; canescens. alpin aize, Webber on xemia in 284 akino’s Teoney Florae ibs gaa 418 alinvaud, Ernest, work of 70 amillaria, . Brandegee on 210 aples of North America 143 ede aes division of the cell and nuc n 394 Marsilea, Nathanbos on parthenogenesis. Sasessssssss ) 2 ea at ae co < Martel work of 405 : Massee, George, ‘wrk of 213 Matthiola, Conti on x “3 -, 415; work of 71 McFarland, I. Hotac e 206 Mcllvaine’s “One Thousand American Fungi” I Medeola Virginiana 181 Megalopus, Engler on 209 rh eh 43 Mertensia, amoena 195; Fendieri 196;. foliosa 196; lanceolata 196; oblon ngi- folia 196 Ppa bbl 176 Meyer, of 212 Mtiactivon Floerkeanum Henrici 12 Micrococcus, agilis ; a new chromo genic 261; carneus ene carnicolor 2693. OO eT ae aS ene ee a eae RE pe ees Ie a fe nS SE oS (RE eae 4 ; 3 d 1900] cumulatus 268; fragilis 268; hae todes 267; ps cus 267; rosaceus ei flav roseus 268; a us 272; roseo- persicus 268; rubellus 267; rubens 267; rubigino 267; genres 267; sub- carn 268 Sicrometry, international unit of 404 Bip est: Insulae A cLigeeane ” 280 Mimulus, Solace fo 425 Minnesota, Ery siphon. ee man on 282; Wheeler on flora of southeastern 282 Miyake, K., work of 141 pace determination of i onoclea, Step ani on bee es embryo sac a. 25 An Gs pal ge sare, Guan Thom s 100 orocco, Camus on ora of 405 orris, E. L., work o orus rubra 407 losses, new North American of North America 12; von a Hi on teeth of 21 Moullefarin, work of 405 Muhlenber at ip , anew Serna 430 Murbach ‘ ates Boudier on 405; Dang n 405; instruction foncai og pide on 405 a eh gd ee SSssass S55 agereag lpr ve ola in on nuclear divi- of 282; nuclear and cell division ZI7s Sturgis on 71 Myrice Gale 407 Myrmec eee Buscalioni and Huber on 136 N Narcissus bg Lidforss on 210 ‘Nash, George V., work of 71 athan anew , wor rk of 138, 213, 282 Naturalist ; , a new journal 43! Neckera , Stunts n7I wey er work of 71, 424 209 Neovossia, Iowens 273; Moliniae 273 New _ rk Poteulcad Garden, Journal of Nitophyltum, Nott on 2 1, Dr. F., work of 134, 211, 214 Nomenciatire Cook on types in ic ; International Botanica al Con g§ n 404; 1 ees flere o South America fe INDEX TO VOLUME: XXX 439 North Carolina, Johnson on flora of 405 Nothocaleis cuspidata 182 Nott, Charles P., work of 287 oe cle eus, and cell plate 73, 154; Chodat 405; division in Fuligo varians 217; in wei ms 405; Nawaschin on divi- sion of 283 CEn erie ee 351; canescens 351; humifus ieee Kellerman on non-indigenous flora Olive 2 Woy eres 142 Ono, . work of Onoclea ae lis 408 Opuntia, fulgida, Serietiols of a 351; fulgi da, roots of 349, 350; non —— of 351; Whipplei, rode a Orehis, latifolia 735 —. 182 Ree ea arcticum 19; Idahen oF ellii esi 2 papillosum 20 Gon a cinnamomea 362; Claytoniana pee gran: s 408; spore mother cells of sci tir Qo E.; pgs se Osterhou re ae of 139 Ovules es Cease Wardell on 137 P Padina pavonia 409 how E., — 430 Palladine, work of Ponavicieias. Stephan n 70 almer and Kee soe "#Stracture of the salen irdle” Paris’s “Index Bryans ae a) ) <5 mn B22". 3) Qu ns —-_ 2 nN lea, Sapp 138 yong has he ae of 79; herba- m of a Pe a Fis oi of 431 Pelza hassnuspeced 414 Penstemon, Suksdorf on 425 Peperomia pellucida, endosperm and em- Peace physiological observations on 171 Pericycle, Fi scher on 286 Periodical, secaemational for names 404 eronspora parasitica, Wager on 210 Perrot, E. 4 Persea Carolinensis 407 Personal; Ambronn, H. 72, 288; Appel, Otto 143; Arthur, J. C. 72; Atkinson, 440 BOTANICAL GAZETTE . 216, 360; Audubon, J. J. 430; Bailey, W. W., 143; Bessey, C. E. 144, nee ato, Busse, Walter 142; Britton, ; Carleton, M. A. 216, 288; Church, ie Ry 142: Clements, Beck: Hitchcock, As: S; 6, 360; H ottes, C. F a ut Moor AC. fe » 142; Olive, E. W. 142; Rimbach, A. t 144; regi OH. F543; i 2, 216; Schaffner, . ‘ rwood, L, M. 430 bb, J. E. 72; Williams, E. F. 142; ‘Woods, AS Fs a32- 2ukal, H, 216 Petrocelis filth 100 Peucedanum foeniculaceum 182 varepy Duis nd on 424 Phaius Blumei, abig Soien on 213 Phascum cuspidatu um Phaseolus ee _ cell plate of 83 Phlebodiu ureu x vulgare ale esantachntin Phlox, anaieola a 63; Su Mstiort oe noe: rede in botany and in : hordell Diced Miyake on 141; Polacci n 0 Phragmites communis 273 Phyllitis scolopendtium X Ceterach offi- ine enpeiiieule 63; interme- Physmatomyces, Rehm on 424 Physalis longifolia 182 Pinus Ta eda 407 Piper, eure sac of 9 hae lacie | DECEMBER Piper, C. V., work of 2 Pisum s sativum, = slate of 83 Plantago m Plaxisodioghors easiiees Nawaschin on 282 ; Plasmodium, Nawaschin on formation of = tanthera montana 182 Plate, sd es opis and function of the cell 73, Plowright, work of 405 Polacci, Gino, ‘york of 358 Polla rd, Ory SP of, 70, 425 Pollen other: alle a Gladiolus 148 Pollination and seed dispersal, Ule on 28 Polyides cheese Polypodium incanum 407; vulgare ele- gantissimum x Phlebodium aureum Polyporus carneus, sti ioe on 420; juni- perinus, Schrenk o Po byitiekaid aculeatum mee angulare 411 Polytric hum Jensen, Holzinger on 425 Potentilla, Piper on 210 Ss eton pieces $ 31 Prantl’: s “ Lehrbuch der Botanik” 66 Prasiola leprosa 104 ere: Schaible on effect of air 209 n, Carleton E. 3 oe a, Wiegand o Praean Bokorny on e uories vot 286; a light, Palladine on 214; vacuoles 7 gymnosperms, Arnoldi on 308 Protobasidiomycetes, —_ on 213 Protoccoidez, Chodat on 135 Protoplasm, Kny on ‘eteicalinhe living 28 Pseudoleskea atrovirens 22; Best on 71, eon ; — cone | Holzingeri 124; patens- mere ico: Poralel eseinchtn 182 Psorotheciopsis Rehm Pterogonium gracile Californicum 21- Pylaisia polyantha drepanioides 21 Pythium tenue, Gobi on 427 apt aquatica 407; ene on 274, 276} gra 407; virens 407 at Raciborski, M. et, personal 430 Radais, work of Rafines es Watern Minerva” 216 ———— 1900 | Ramaley, Francis, work of 282 ee mie on 282 Re k of 424 che: Nea, ise of 135 a of Mi chigan Academy of Science Review s: Bailey’s “Cyclopedia of Ameri- Horti n 132; Costantin’s “Ta nature tropi- Coulter’s * ‘Catal ogue ‘e the sient of Indiana” 421; r .and : 279, 420; nographien afrikanischer Pflanzenfamilien ” 4 vans’ “ Alas- Hepa a SAL Os ebel’s “Or ig’s Pi omideena a Pflanzen- Jelliffe’ 8 Phe ai of the flora of Long Island ” 420; Johannsen’s “ Das Aether- Verfahren” 280; Ke ee ler’s “ Our native trees” 133; Koning’s “Der Tabak” 278; Lotsy’s “Alkaloids in i sion 279; Lounsberry’s “A gul uide to oO dberg’s “Flora of Montana” 61; Scott's * Studies in fossil sige ase der Botanik” 67; Urban’s “ Flora of the West areas ea ie rburg’s it M u ia”? 207; Wie «Ty Die Rohstoffe des Paivmeurticht ” 66, 2795 d’s “Cycadeoidea of th Black ae oe ee “ “The cell” 65; Woo tigm ” 981; Zeiller’s “Sti ose . Hianents paobotaniane : INDEX TO VOLUME XXX ; 441 Rhizidiomyces ichneumon, Gobi on Rhode Island Summer School for nature ah udy 143 hus Toxicodendron 406 Ries leptanthum 119; saximontanum Richter, Oswald, work o Rimbach, Augu patie personal 144 Roberts, H. F., personal 1 rad gah Charles 1 136 Robin ., personal 142, 216 Rolland, work 0 ee Roots of Cactacez Rosa, gross eserrata a pisocarpa 120; Woodsii I Rostowzew, S, oe . rey Rowlee, W. W., sr atl iecuas ioe Natdenahe on 213 Rydberg, P. A. “Flora of Montana” 61; work of 71, 425 > al Adansonia a 408 n s, Loew on 214 Salvia, Fernald on 136; pratensis 186 Salts, Ono on effect on growth 422 u Sargassum vulgare 409 seared aera Johnson on 208; em- Saxitaga, brinediinils 119; re 8h 118; Piper on 210; Wiegand on Schaffne " Nips oe of 431 Schaibie, ¥ r., work 0 Schenley P Park, botanical ation 430 Schinz, Hans, work 0 Schrenk, eee yon, asuabe! 143, 288; work o Schuh, R. E work of 135 Schumann K. 359; work of 135 C. $., work of 282 Seolopendriom vulgare X Ceterach offici- Scott's is “ Sudies in fossil botany ” 352 Sedum 18 : peg anpassel and emcee a Ule on. Heinricher on w _Rhabdonia tenera _. Rhadinocladia, oak on 135 Rheotropism, Juel on 136 ., work of 142 —_ A. ee — sf 139 442 BOTANICAL GAZETTE r,.(. L., work-of 1 work of 428 Silene, Tetonensis 117; Watsoni 117 Silphium a fertilization in 256 a 69 Sisyrnchium, Bicknell on 71, 210; Cali- ornic Sisymbrium, canescens a microtites 59; umbro 60; eye tonii 59 ork of 71 Diels on 424; rotundifolia 407; srnnenens 407 Smith, C. H., bequest 431 Smith, R. Wilson, 301 personal 359 Solanum, Fernald "Solidago, dilatate 3 ots multiradiata sco- i m 197; Pollard on 70; semper virens 4 bt canescens 64 Sorus, development of in Lessonia litto- ralis 332 Sphaeropsis malorum 48 Sphagna, W renee On 210, 213 Spindle, ee 361; multipolar in Gladiolus 45 : on in myxomycetes 217; sel Hag alls e * deslesnmen regalis 364; Na Stahl, E., on gorhac ts za 6 Stangeria as gigs Lang on 212 Stapf, work of 4 Staphylea penlians Starch, snr of evergreens in winter, et2 ly: Stenanthella, a a on 425 * Stephani, Franz, duh ~ 7°, 135 » pers Stigeoclonium, Sieg Poh form in 289 €o ? Stipa avenacea, mechanics of awn 113 Stipe in Lessonia Stone’s “ Plants of Lake Quinsigamond ” Strasburger, Noll, Schenk, and Schimper’s “Lehrbuch der Botapik ” 67 Stuntz, S. C., work of 71 _ Suksdor . N., work of 1 135, 424, 425 i ct D. B. » personal 2 Symphy¢ ogyna, "Stephani te a T Table, Rostowzew on laboratory 215 ‘Tannin in galls, origin of 274 [DECEMBER Taxonomy 70, 135, 209, 4 Tendril, Noll on lalate of coiling 214 Terminalia chebula, —_ on 276 Texan fungi, Long on 429 Thalictrane Davis on a Thaxter, Roland, personal a Thelypodium auricuiatum Thermostat, zapek on prlegs Ty is 4 - 37 homas, Ethel N. , work of 213, 428 sara Molinia 27 a erlake, H.G. 73,1 A ee Rydber rg o Scar Torrey Day, celebration of Toumey, J. ennsege 725 work of 213 af = a nsendi oxicity, of acids an rip ts, True on 140; sodium salts, Kahlenberg and Aus- n 358 t 35 Travel, notes o prone T = , Wm., personal 216 Trentepo ohlia virgatul 409 Trichia fallax 217 Tri chostomum, dicranoides 17; macro- 17 401 Triplochitonacez, Schumann on I Tripterocladium leucocladulum campto- carpum Trisetum, rah = by Triticum v Troximon, Sts on 424 True, R. H., nal 142; work of 140 Tulipa, Pe ag iectiltiation 5 in 252 Types in botanical nomenclature, Cook 427 U Ule, work of 428 Ulva lactuca 409 ! Underwood, L. M., personal 430; work of Urban’s “ Flora of the West Indies ” 421 Uredinew, Plowright and Magnus on 405 Utricularia, Meister on 70 V Valerianella, Suksdorf on 424 an Hook, J. M. 394 Variation, De Vries on 405; Gallardo on * statistical methods in 405; Krasan on 405 Veratrum, Rydberg on 425 - Verbena, ‘hastata I (a8 Miche 130; stricta 130; urticifolia 130 , | 1900 | Verbenacee, Giirke on 13 Vergnica peregrina, Heiaricher on 215 Viburnum, Suksdorf o 405 Viola, ainplcfaia 193; Pollard on 70; Thorii 193 Violet, Po llard on 425 Vitis rotundifolia 407 W Wager, Harold, work of rh Warburg’s “ Monsunia ” Ward’s *‘ Cycadeoidea of the Black hills” 41 Warnstorf, C., work of 21 Washington plants, Shkedort | A 135, 424, - Washingtonia divaricata 6 Water, eee! ological larger tre of the Chicago draina age 143; plants, Kny on Po, cellular plasma in 28 Waugh, F. A. Webb, J. E., ie rsonal 72 Webber, H. . vor of 138, 2h Sbhebitoa cee ie mutata cullat rinata 20 Lal ao ace si Weisia, viridula es vaca gymnosto- moides 13; Wimmeri West Virginia plants, Morris on 425 INDEX TO VOLUME XXX 443. Wheeler, wt tg + bet of 282 Wiegand, K M. 25; work of 210, 425 Wiesner’s *“ Die Ro hstoffe des acme pr eadl 79 n, E. ’ de, work of 71, 405 Wildem Williams, "f mile F., personal 142 pi a pateihen n 425; Rowlee on 71; Sea n 425 Wilson's “ = “The cell” 65 Asc F., personal 431; ‘ Stigmo- e” 281 Wordsdell, W2G;, et of 137 Wright, W. F., work of 424 Wyoming plants, Nelson on 71; new species of 117 Xenia, deVries on 212; Webber on 284 ¥ Yale pps 3 School 430 Yasuda, Atsushi, work of 285 Yeast, ova on enzymes of 211; deter- mination of 404 Zeiller, thes Eléments de Paléobota-: nique ”’ 416 rade marin Zukal, Hugo, acah of 216 Zygadenus, Rydberg on 425 A Tonic and Nerve Food HORSFORD’S Acid Phosphate. When exhausted, depressed or weary from worry, insom- nia or overwork of mind or body, take half a teaspoon of Horsford’s Acid Phosphate in half a glass of water. It nourishes, strengthens and im- parts new life and vigor by supplying the needed nerve food. Sold by Druggists in original packages only. Joe odont in a new size 25C. of the Liquid The event of the year in dentifrices. Beware of counterfeits dentifrice. Insist upon getting the genuine at the stores. It necessar send 25c. direct to _ Pia P. 0. Bo 247, New York City. ; WML & sie ’ NEW YORK LONDON NNENS ENN TALCUM Delightful After Bathing A Luxury After Shaving A POSITIVE RELIEF FOR CHAPPED <— CHAFIN d all afflictions - a Peek Removes all odor of perspirati Get MENNEN’S. (tbe theme a little higher in price, , per rhap orth Ss substitutes, efuse all = liable to do har . Sold e every where, or mailed for 25 cents. — — ARD MENNEN CO., Newark, N. J. Awarded “GRAND PRIX” Paris Exposition 1900 (16 West 23d St. 1.166 Broadway. 504 Fulton 8t, 169 Tre: — St. wae ¢ ey tse 748tate St. pur New York: Brooklyn: Boston: Philadelphia; Chicaco: Rely on Platt’s Chlorides as your household disinfectant. An odorless, color- gases, thus prevent- ing sickness. Sold in quart bottles only by druggists and high- B, Platt, Platt Street, New York Ri ere es MADE FROM THE BEAN’ E.. PURE! HEALTHFUL! STRENGTHEN Sold at om oneness and b CERS EVER THERE - | Waukesha Hygeia Mineral Springs Water * 2 FROM IT 18 MADE Boro-Lithia Water, Gin- ger Ale, and Wild Cherry Phosphate. De Me a et Se 47 to 53 E. Kinzie St., CHICAGO, Il. Telephone in 605 and 608 | Waukesha Water Co. We will deliver anywhere in the S. we can reach by express [we will pay express charges] a case containing enou TOILET PAPER To last any average family a full year; finest satin tissue, A. P. W. BRAND. ever saw better at anything like the price per rooo sheets, we will refund the lene: Our Ss mearerey is ood —we a are the largest makers in the world, and originated the pirkiented roll, A PW, Paper Company Montgomery St., Albany, N.Y. WW one Sheets and Booklet mailed FREE | | : 8 ’ ’ q . ‘ 4 4 } IT KEEPS THE STOMACH SWEET Dr. Alexander Haig, London, in “ Food and Diet,” says: © Records Srom all sides show that the less animal Jlesh a people take, the better do they comz out in trials of force VZA . e "aa “2 production, and CSPeciury in endurance.” The og 4 i Same distinguished authority also says: © The proof of the poisonous nature of meat lies in the beneficial results of refraining Srom it,” Lt seems only common sense to eat less meat and more Quaker Oats. This delicious food contains all of the food-elements of meat and none of its unwholesome qualities, It is at once the most perfect and most economical food. Easy to Buy and Easy to Cook. Sold by all dealers in sealed pack- O sesecnsese BERRA APR AIORR AOR IOIRPER Babbitt’ S QOOOOOOOO QOOOOOO0O O O O) e O 6 $ 00 # eo : alk al %, 9 QQQQOQOQOCOQQOOOOOQOOOOOO OORQQO ES) QOOOO QOQOOOOOO OOO DOSOOOOOOO is sc 0QO0OK) “3 sisusesccesteeceasetccaoet ste oe OX ye 2000) CO 20 Babbitt’s Best Soap sessscccsosccgn 3, ” 9 : 238 Babbitt’s |17'76| Soap Powder 222 & 449 O ° Babbitt’s Pure Potash or Lye 6 2 : Babbitt’s Best Baking Powder 3 2299000092800202280220090099220222QQQ002 SERS Hr esbesSecosscbosooseoe ee YOORQGSGHOHGGS OO elelele 539988888000 Quality DOOOOOOOOO SBOESEEEEEESS [Jn i SOSSR ISSA Soc3 Unttormity &-35c25 DOOCOOOOOO SOOOCOOOL) 355: OOOHOOSSS Economy SOOSORK soon eo OOO DODOOOSOONNK O88 5 Strength SOSSSRIIOS of 208 3 siecenceee 8885 SSSSSSIIOONT LES SOOOO OOOO Se 5555 Moe 8. Beat Ree JOOOOOOOOOOOO age dass ae oy ERE is style—approved—original—pupular. Quality and nae wngon ng the finest; with cola, shirt and cuffs all under brand, made to fit each. other, which makes colla 2 cus cost 25 c It does not pay to Shi , $1.50, or He 00, dpendice on the & Sind 5 _ want hay your furnlener: UNITED SHIRT AND COLLAR CO., Makers, TROY, N.Y The Improved BOSTON GARTER The Standard for Gentlemen ALWAYS EASY The Name ‘“ BOSTON GARTER” is stamped DE on every loop. The . CUSHION BUTTON «_CLASP Lies flat to the leg—never Slips, Tears nor Unfastens. SOLD EVERYWHERE. Sample pair, Silk 5c, Cotton 25 Mailed on receipt of prices OST CO., Makers gg Mass., "U.8.A, : ME-EVERY PAIR WARRANTED -@ cre Absolute Security and comfort which the Dr. Deimel underwear affords in the most trying climate is remarked upon by all its wear- ers. Those whose powers of resistance have been weak- ened by the pernicious use of enfeebling flannels derive new strength from this ideal linen undergarment. > All true Dr. Deimel am- ples of the cloth free. ¥ We also make the finest Dress Shields in existence. Can be washed, are odorless. A guarantee with every pair. ¥ THE Deimel Linen-Mesh System Co. 491 BROADWAY, NEW YORK 2202 St. Catherine St., 11! sages ctor St., Montreal i. sco, Cal 10-12 Bread St., 728 15th St., N. W., London, E. C- Washington, D. C. WON A DIPLOMA OF THE AND POSSIBLE PRIX fe NAL ENTY-FIVE MEMBERS, AND IN WITH TWENTY OTHER TYPEWRITERS. The —— pone US kona Co., USE, N.Y., U. A. The No. 2 “* New Manifolding ”’ HAMMOND TYPE- WRITER IMPROVED 4 4 METHOD INCREASED POWER AUTOMATIC . BLOW SUPERIOR RESULT /t also has a number of Valuable Mechanical Improvements. 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Controls All Pain, Bleeding and Inflammation. There is only ONE fF “POND’S S EXTRACT and haga tage knows its purity, strength and great goats! value. yoo wage e weak, watery Witch Hazel ov Fepre ented to be ‘ POND’ S EX ‘the TRACT. They ain ‘‘ pie alcohol,”’ wtiie ies the skin, and, taken internally, is a deadly roe Get POND’S EXTRACT, sold oNLY in SEALED BOTTLES in BUFF wrappers. POND’S lsc partbebirn abe Serbs ab gs sod bruises piles, however severe. It is a specific = generally co in in all skin diseases, a es quick relief to bur Genuine bears blue signature on label. # ve and approred 5, % amin Or tm oy gh os ———— EIS Departmen sy mee B —— am nadand approved ay the examined Oo approved tne Scentifx Ovepart ment ana i Sate OF BEL? Registered by U,$, Patent Office s ae 3 ~ ¢ : ae * 24 Or att “ ‘ f WN | x ITHIA WATER OF VIRGINIA, Springs Nos. 1 and 2, IN COMPOSITION APPROXIMATES THE Chis rp =< Blood Serum. ; ~*~ A BLOOD FOOD AND NUTRIENT. John V. Shoemaker, M. D., a D., Professor of Materia Medica and Therapeutics in the Medico-Chirurgical College of Phila- delphia, etc. New York Medical Journal, July 22, 1899 (extract). ‘*An additional advantage and extremely importart reason for the PECULIAR EFFICACY of the BUFFALO LITHIA WATER lies in the fact that its composition approximates that of the Serum of the Blood; therefore it is admirably fitted for absorption into the blood current and immediate incorporation with the watery portion of the nutrient fluid. it becomes at once identical with the BLOOD SERUM. These are quali- _ ties which far surpass those possessed by any extemporaneous solution | of a single chemical preparation, as when a lithia tablet, e. g., is dis- solved in water for immediate administration. When we speak of a dose, it is of a quantity altogether relative, and what the physician emphatically desires in a dose is therapeutic efficiency. This we ave in the TTHIA oot ‘Those who have made use of this water and carefully noted its ef- fects have often been surprised at the results obtained from amounts so small, according to chemical analysis, of Lithia and the accompanying Salts. The explanation of this extraordinary activity ts doubtless to be found in the conditions just adduced. . poet Both of these waters are powerful Nerve Tonics, and No. 1 1s also a potent Blood Tonic, and is especially indicated in all cases where there is Poverty or Deficiency of Blood. In the absence of these Symptons, No. 2 is more especially indicated. WATER is for sale by Grocers and Druggists generally. ‘ : : . »stions. sent to any address. Testimonials, which defy all imputation or questions, sent to any add _ PROPRIETOR, BUFFALO LITHIA SPRINGS, IRGINIA Springs are open for guests from — koe 58 eee Railway They are reached from all directions over the Danville Division of the ou : oS i? * iL ea S a ae _—_ Sa eee Por ie U at ere se A. 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KF RYAPAYAPAPAPAPAPAPAPAPAYAAY AY EY AY mths AF ey ey er CyrCeee ee ee ee , alicttack. shack alah ahech thas desk themh.hemsheh theta Pp 2 yy Oot Ni Ny ¢ ~* POO Fe ee ei Good for the Spotter of ! ,He spotted a Baot on ie a] fs at