Part 1
ANNALS OF BOTANY VOL. VIIT Orford PRINTED AT THE CLARENDON PRESS BY HORACH HART, PRINTER TO THE UNIVERSITY ANNALS OF BOTANY EDITED BY ISAAC BAYLEY BALFOUR, M.A., M.D., F.R.S. QUEEN'S BOTANIST IN SCOTLAND, PROFESSOR OF BOTANY IN THE UNIVERSITY AND KEEPER OF THE ROYAL BOTANIC GARDEN, EDINBURGH SYDNEY HOWARD VINES, M.A., D.Sc., F.R.S. FELLOW OF MAGDALEN COLLEGE, AND SHERARDIAN PROFESSOR OF BOTANY IN THE UNIVERSITY OF OXFORD D. H. SCOTT, M.A., Pu.D., F.RS. HONORARY KEEPER OF THE JODRELL LABORATORY, ROYAL GARDENS, KEW AND WILLIAM GILSON FARLOW, M.D. PROFESSOR OF CRYPTOGAMIC BOTANY IN HARVARD UNIVERSITY, CAMBRIDGE, MASS., U.S.A. ASSISTED BY OTHER BOTANISTS VOLUME VIII With XXIV Plates, in part coloured, and 5 Woodcuts Bondon HENRY FROWDE, AMEN CORNER, E.C. OXFORD: CLARENDON PRESS DEPOSITORY, 116 HIGH STREET 1894 <e “CLIBRS Ry iveR sake STANFORS ] HTS 84D ERRATA. Page 238, last line, for ‘and formed ’ read ‘ though they formed’ 248, line 12 from bottom, for ‘ tetrach’ read ‘ tetrarch’ 284, 285, 295 298, 299, 302, 306, 338, 349, 9” 341, ” 29 12 5; » omeat ‘it?’ I4 4) » for ‘sexual’ read ‘ asexual’ ary, » for ‘nuclei’ read ‘ chromosomes’ 5 from top, for ‘ not’ read ‘no’ 2 » » for ‘with’ read ‘without’ 13 from bottom, for ‘alternation’ read ‘ alteration ' 13. » » fJor‘small’ read ‘single’ ? ae », @ele comma after ‘ margine’ 9 from top, for ‘ Favellae’ read ‘ Favellis’ 5 from bottom, for ‘/rons’ read ‘ fronde’ 2 » »» insert comma after‘ minortbus’:; dele comma after ‘ tenus’ last line, for ‘ Sphaerosporae’ read ‘ Sphaerosporis’ line 2 from top, for ‘divisae’ and ‘immersae’ read ‘dtvisis’ and © smmersis’ line 7 from top, for ‘vestitis’ read ‘ vestita’ 342, ,, 24 from bottom, for ‘MYRIOGLOSSA’ read ‘ MYRIOPHYLLA’ CONTENTS. No. XXIX, March, 1894. CAMPBELL, D. H.—Observations on the Development of Marattia Douglasii, Baker. (With PlatesI and II) . . Dixon, H. H.—Fertilization of Pinus silvestris, (With Plates III, IV, and V) . . FARMER, J. B.—Studies in Hepaticae : On Pallavicinia decipiens, Mitten. (With Plates VI and VII) ; Peirce, G. J.—A Contribution to the Physiology of ‘the Genus Cuscuta. (With Woodcut 1, and Plate VIII) NOTES. MASSEE, G.—A New Cordyceps . . WaTson, W.—Absorption of Water by Dead Roots . No. XXX, June, 1894. BAKER, J. G.—New Ferns of 1892-3. . GiBson, R. J. HARVEY.—Contributions towards a Knowledge of the Anatomy of the Genus Selaginella, Spr.: Part I—The Stem. (With Plates IX, X, XI, and XIJ) SEWARD, A. C.—On Rachiopteris Williamsoni, sp. nov., a New Fern from the Coal-Measures. (With Plate XIII) y FARMER, J. B. and REEVES, JESSE.—On the Occurrence of Centro- spheres in Pellia epiphylla, Nees. (With Plate XIV) . NOTES. GREEN, J. R.—On the Germination of the Pollen-grain, and the Nutrition of the Pollen-tube . . GROOM, PERCY.—Botanical Notes, No. 6: On the Extra- floral Nectaries of Aleurites . . No. XXXI, September, 1894. Boov te, L. A. and WORSDELL, W. C.—On the Comparative Anatomy of the Casuarineae, with special reference to the Gnetaceae and Cupuliferac. (With Plates XV and XVI} PAGE 53 11g 119 131 133 207 219 225 228 vi Contents. Kny, L.—On Correlation in the Growth of Roots and Shoots . XSTRASBURGER, E.—The Periodic Reduction of the number of the Chromosomes in the Life-History of Living Organisms . PFEFFER, W.—Geotropic Sensitiveness of the Root-tip \ WAGER, H.—On the Presence of Centrospheres in Fungi. (With Plate XVII). . . . HoutMEs, E. M.—New Marine Algae. (With Plate XVIII) Bower, F. O.—A Theory of the Strobilus in Archegoniate Plants NOTES. WILLIAMS, J. LLoyp.—The Sieve-tubes of Calycanthus occidentalis. (With Woodcut 2) GREEN, J. R.—The Influence of Light on Diastase HuMPHREY, J. E.—Nucleoli and Centrosomes . No. XXXII, December, 1894. Davis, B. M.—Euglenopsis: a New Alga-like Organism. (With Plate XIX) . MOTTIER, D. M. —Contributions to the Life-History of Notothylas. (With Plates XX and XXI). . NEWCOMBE, F. C.—The Cause and Conditions of Lysigenous Cavity- formation SPALDING, V. M.—The Traumatropic Curvature of Roots. (With Plate XXII). . WRIGHT, C, H.—On the Double Flower of Epidendram ‘vitellinum, Lindl. (With Plate XXIIT) JOHNSON, T.—Two Irish Brown Algae : Pogotrichum and Litosiphon (With Plate XXIV) . . . NOTES, Bower, F. O.—On Apospory and Production of Gemmae in Tri- chomanes Kaulfussii, Hk. and Gr. (With Woodcuts 3, 4, and 5) . . . . Dixon, H. H. and Joy, J —On the Ascent of Sap PAGE 265 281 317 321 335 343 367 370 373 377. 391 403 423 453 457 495 468 INDEX. A. ORIGINAL PAPERS AND NOTES. PAGE BAKER, J. G.—New Fems of 1892-3 . : 121 Boop LE, L. A. and WoRSDELL, W. C.—On the Comparative Anatomy of the Casuarineae, with special reference to the Gnetaceae and Cupuliferae. (With Plates XV and XVI) . . . - 231 BowEr, F. O. A Theory of the Strobilus in Archegoniate Plants . 343 On Apospory and Production of Gemmae in Trichomanes Kaul- fussii, Hk. and Gr. (With Woodcuts 3, 4, and 5) : - 465 CAMPBELL, D. H.—Observations on the Development of Marattia Douglasii, Baker. (With Plates I and II) . . . I Davis, B. M.—Englenopsis: a New Alga-like Organism, (With Plate XIX) . . 377 Dixon, H. H. —Fertilization of Pinus silvestris. cwith Plates Ill, IV, and V) . . a1 Dixon, H. H. and Jory, J. —On the Ascent of Sap . . 468 FARMER, J. B.—Studies in Hepaticae: On Pallavicinia decipiens, Mitten. (With Plates VI and VII). . 35 FARMER, J. B. and REEVEs, JEsSE.—On the Occurrence of Centro- spheres in Pellia epiphylla, Nees. (With Plate XIV) . aIg GIBSON, R. J. HARVEY.—Contributions towards a Knowledge of the Anatomy of the Genus Selaginella, Spr.: Part I—The Stem. (With Plates IX, X, XI, and XI) . . . . . 133 GREEN, J. R. On the Germination of the Pollen- “grain, and the Nutrition of the Pollen-tube . . . - 325 The Influence of Light on Diastase . 370 Groom, PeRcy.—Botanical Notes, No. 6: On the Extra-floral Nectaries of Aleurites . . . - 238 Ho.MEs, E. M.—New Marine Algae. (With Plate XVIII) . - 335 HuMPHREY, J. E.—Nucleoli and Centrosomes . . 373 JOHNSON, T.—Two Irish Brown Algae: Pogotrichum and Litosiphon. (With Plate XXIV). . . . . . . - 457 JoLy, J.—See D1xon. Kny, L.—On Correlation in the Growth of Roots and Shoots’. . 265 MASSEE, G.—A New Cordyceps . . 11g MoTTIER, D. M.—Contributions to the Life History of Notothylas. (With Plates XX and XXI) . . . . . 391 Index. NEWCOMBE, F. C.—The Cause and Conditions of Lysigenous Cavity- formation PEIRCE, G. J.—A Contribution to the ‘Physiology of ‘the Genus Cuscuta. (With Woodcut 1, and Plate VIII) PFEFFER, W.—Geotropic Sensitiveness of the Root-tip REEVES, JESSE.—See FARMER. SEWARD, A. C.—On Rachiopteris Williamsoni, sp. nov., a New Fern from the Coal-Measures. (With Plate XIII) SPALDING, V. M.—The Traumatropic Curvature of Roots. (With Plate XXII). STRASBURGER, E.—The Periodic Reduction of the number of the Chromosomes in the Life-History of Living Organisms. . WAGER, H.—On the Presence of Centrospheres in Bung (with Plate XVII). . . . WATSON, W.—Absorption of Water by Dead Roots : WILLIAMS, J. LLoypD.—The Sieve-tubes of Calycanthus occidentalis. (With Woodcut 2) . . . . . . WORSDELL, W. C.—See BooDLe. WRIGHT, C. H.—-On the Double Flower of Epidendrum vitelhinum, Lindl. (With Plate XXIII) B. LIST OF ILLUSTRATIONS. a. PLATES. I, I. III, IV, V. VI, VII. VIII. 1X, X, XI, XIL XIII. XIV, XV, XVI. XVIL XVIII. XIX. XX, XXL. XXII. XXIII. XXIV. On Marattia Douglasii (CAMPBELL). Fertilization of Pinus silvestris (D1xoN). On Pallavicinia decipiens (FARMER). Physiology of Cuscuta (PEIRCE). Anatomy of Selaginella (G1BsoN). Rachiopteris Williamsoni (SEWARD). Centrospheres in Pellia epiphylla (FARMER and REEVES). Anatomy of Casuarineae (BOODLE and WORSDELL). Centrospheres in Fungi (WAGER). New Marine Algae (HOLMES). Euglenopsis (DAVIS). Notothylas (MOTTIER). Traumatropic Curvature of Roots (SPALDING). Double Flower of Epidendrum vitellinum (WRIGHT). Two Irish Brown Algae (JOHNSON). 6. WoopcuTs. 1. Physiology of Cuscuta (PEIRCE) . . . . 2. Sieve-tubes of Calycanthus (WILLIAMS) . . . 3-5. Apospory and Production of Gemmae in Trichomanes Kaul fussii (BOWER) . . ‘ : PAGE 403 53 317 207 423 281 321 119 367 453 66 368 . 466-7 Observations on the Development of Marattia Douglasii, Baker. BY DOUGLAS HOUGHTON CAMPBELL. Professor of Botany, Leland Stanford Junior University, California, U.S.A. —_~¢-0—= With Plates I and I. ==¢¢—o— HILE collecting in the Hawaiian Islands during the summer of 1892, I was fortunate enough to find near Hanalei, upon the island of Kauai, a large number of very young plants, of Marattia Douglasii, and with them a suf- ficient number of prothallia with embryos to make it possible to study the principal points in their development. Until Farmer's! recent paper on Axngiopteris, no account of the embryogeny of the Marattiaceae has appeared, except a brief mention made by Luerssen*? of the arrangement of the primary organs. His material was however too scanty to throw any light upon the early divisions of the embryo. The growth of the prothallium is very slow and the development of the embryo correspondingly late, so that in artificial cultures more than a year must ordinarily elapse before embryos are developed. This probably accounts for the fact ! Farmer, On the embryogeny of Angiopteris erecta, Annals of Botany, Vol. vi, No. XXIII, Oct. 1892. 3 Luerssen, Handbuch der Systematischen Botanik, Vol. i, p. §&2. (Annals of Botany, Vol. VIII. No. XXIX. March, 1894.) 5 f 2 Campbell—Observations on the Development that the embryogeny has not been worked out before, as the spores germinate without difficulty. The Marattiaceae at the present time include four genera, with about 25 species, confined exclusively to tropical regions. In a fossil condition they are much more numerous and diversified than at present, and according to Solms-Laubach ' comprised the great majority of the Carboniferous and pre- Carboniferous ferns. Their great antiquity, as well as their morphological peculiarities, point to their being primitive forms, and this makes any information concerning their life- history of especial interest. As I have endeavoured to show in several other papers, there is constantly accumulating evidence pointing toward the Eusporangiatae as the forms from which the leptosporangiate ferns have descended, and we shall see that this view is still further strengthened by a study of Marattia. Up to the present time, we have almost no information concerning the embryogeny of the homosporous Eusporan- giatae, except Farmer’s! recent paper on Angiopierts, and this being the case, it is hoped that the present paper may have some interest, as adding somewhat to our knowledge of this important group. Marattia Douglasit is the only representative of the order found in the Hawaiian Islands. It is a large fern with massive leaves, two to three metres in length. It is common in the damp forests at an elevation of 300-400 metres, and occurs upon all the larger islands. The prothallia were found growing thickly upon clay soil near the plants, and were easily distinguished from ordinary fern-prothallia by their fleshy consistence and darker green color. Most of them already had young plants of various sizes attached to them, but a number were found with young embryos. The material was fixed with one per cent. chromic acid, and then gradually transferred to alcohol for study after my return to California. ! Solms-Laubach, Fossil Botany, pp. 142-152. 3 Loc. cit. of Maratha Douclass, Baker. GERMINATION OF THE SPOREs. Fresh leaves with ripe sporangia, as well as prothallia and young plants, were collected and were carried back to California, reaching there after more than two weeks con- finement in a tin collecting-box, in good condition. Spores were sown and germinated promptly, but the subsequent growth was very slow. No embryos developed from the prothallia, but the latter, as well as the young plants, have grown perfectly well under bell-jars in the laboratory. As the germination of the spores, and the devclopment of the prothallium, has already been exhaustively studied by Jonkman!', my study of these was mainly confined to the apical growth of the older prothallium, and certain points in regard to the development of the sexual organs. Jonkman showed that the prothallium differs mainly from that of the leptosporangiate ferns by its more massive character. It closely resembles such a liverwort as Pellia, for example, having a thick midrib that merges gradually into the thinner wings, which, however, unlike most ferns, are more than one cell thick, except at the very edge. In artificial cultures the growth is very slow, and it may be a year or more before the sexual organs are mature. If fecundation does not occur, the prothallium seems capable of unlimited growth, and may reach a length of two centimetres or more. In the species studied by me, the older prothallia were always decidedly elongated and not orbicular like those of Angiopteris described by Farmer*. These old prothallia, like those of Osmunda, often develop adventitious shoots (Fig. 4 6), and in all ways give evidence of almost unlimited capacity for independent growth. In the older prothallia, the midrib is very massive and projects strongly from the lower side where the arche- gonia are produced in great numbers. The dark green 1 Jonkman, La génération sexu¢e des Marattiacées, Arch. Ncerlandaises T. XV Pp. 199. * Loc. cit. p. 265. 4 Campbell— Observations on the Development colour of the prothallium is much like that of Anthoceros, and also recalls strongly, as do the prothallia in other particulars, those of Osmunda cinnamomea. According to Jonkman, the prothallia of the Marattiaceae, like those of the other homosporous ferns that have been investigated, usually at least, grow by a two-sided apical cell which is afterward replaced by a group of marginal initials. In all the specimens at my disposal, the marginal group of cells was already formed and was often wider than is commonly the case in the Polypodiaceae. The form of these initial cells is much the same as in other ferns studied by me. When seen from above (Fig. 1), they are more or less oblong in outline, but in vertical section (Fig. 3), they are nearly hemispherical. As usual, the basal segments are cut off by a wall that extends the whole depth of the prothallium, and the segment is then divided by a horizontal wall into a dorsal and ventral cell of nearly equal size. From the latter, the projecting cushion of tissue on the ventral side is formed. This arises abruptly by the multiplication of cells in the ventral segments some distance behind the apex. The superficial cells of the prothallium, especially upon the upper surface, have a strongly developed cuticle which makes it necessary to use a good deal of care in imbedding the pro- thallia for sectioning. Numerous root-hairs grow from the lower surface, especially from the midrib, and fasten the prothallium firmly to the ground. The sexual organs are found in large numbers upon the monoecious prothallia. The antheridia are formed first, and mainly upon the lower surface of the midrib, but also occur upon the upper surface of the prothallium. The archegonia appear later, and so far as my observations go, are confined entirely to the lower surface of the midrib. Upon the old prothallia they may be formed in great numbers, and as they turn dark brown when they fail to be fertilized, are readily visible to the naked eye as dark brown spots thickly studding the broad and thick midrib. Both antheridia and archegonia differ much from those of the leptosporangiate ferns, and the of Marattia Douglasit, Baker. 5 archegonia, at least, seem to give a possible clue to the origin of the sexual organs of the Pteridophytes. THE ANTHERIDIUM. The general structure of the antheridium and its divisions have been so completely described and figured by Jonkman that they will be passed over very briefly here. The antheridium arises from a single superficial cell which first divides into an inner cell, the mother-cell of the sperm-cells, and an outer cover-cell. This latter divides by several curved vertical walls which intersect, and the last wall cuts off a small triangular cell (Fig. 7, 0), which is thrown off when the antheridium opens and allows the sperm-cells to escape. The inner cell, by repeated divisions, gives rise to a large number of polyhedral cells, the sperm-cells. Before these are completed, however, cells are cut off from the adjacent cells of the prothallium, completely enclosing the mass of sperm-cells (Figs. 7-10). In microtome-sections of material stained with alum-cochineal, the transformation of the nucleus of the sperm-cell into the body of the spermatozoid is easily followed. The nucleus of the sperm-cell, at the time when the division is complete, has the appearance of an ordinary resting nucleus, but no nucleolus is visible. The first sign of the formation of the spermatozoid that could be seen, was an indentation upon one side followed by a rapid flattening and growth of the whole nucleus. The cytoplasmic prominence, which according to Strasburger ' is the first indication of the formation of the spermatozoid, could not be detected in material stained with alum-cochineal, and although in some- what later stages (see Fig. 11, a) a slight elongation of the forward end of the spermatozoid was sometimes noticed, it was vague and its limits uncertain. The main part of the body stains strongly with alum-cochineal, and for some time shows unmistakably the nuclear structure, and is sharply differentiated against the colourless cytoplasm. In the later } Strasburger, Histologische Beitrage, Heft IV, p. 116. 6 Campbell_—Observations on the Development stages, the body becomes quite homogeneous. In the nearly full-grown spermatozoid, the body is much narrower and tapers to a fine point at the forward end (Fig. 12). From a careful study of all stages, it seems certain that only a very small part of the forward end is of cytoplasmic origin. Unfortunately, I was unable to get any free spermatozoids, so that it was impossible to test them with Strasburger’s' and Belajeffs?