€ | i 4 [2 BOTANICAL GAZETTE ee eee ee ee ee ee oe ee THE BOTANICAL GAZETTE EDITORS: JOHN MERLE COULTER anp CHARLES REID BARNES VOLUME XLI JANUARY—JUNE, 1906 WITH SIXTEEN PLATES AND SIXTY-TWO FIGURES CHICAGO, ILLINOIS PUBLISHED BY THE UNIVERSITY OF CHICAGO 1906 Mo.Bot Garagen 1906 PRINTED AT | The University of Chicago Press CHICAGO. EE OE ee VTE ee ees Oe TE EN FR a a ee en TABLE OF CONTENTS. : PA The nodes of grasses (with plates I and II) - Mintin Asbury Chrysler The bogs and bog flora of the Huron river valley (with sixteen figures) - Edgar Nelson Transeau Nuclear division in Zygnema (with Ebiten Ii ai IV) Mabel L. Merriman Effect of certain solids upoch the growth of seedlings in water cultures (with four —— E = ‘J. F. Breazeale Chemotropism of fungi - - - Harry R. Fulton The embryology and development of Riccia fee and Riccia crystallina (with plates V-IX) - - Charles E. Lewis A morphological study of Sargassum fili pendula. Con: tributions from the Hull Botanical Laboratory. L II (with plates Xand XI) - - - Etoile B. Simons Chromosome reduction in the microsporocytes of Lilium tigrinum (with plates XII and XIII) - John H. Schaffner Cytological studies on the Entomophthoreae. I. The morphology and development of iiss a plates XIV and XV) - Edgar W. Olive Cytological studies on the Pica phtbouee Il. Nuclear and cell division of seis ore (witht gay XVI Kee Edgar W. Olive Biological relations of ae nabe Il. Aor tion of water by leaves - V. M. Spalding New species of Californian plants (with two ro figures) Alice Eastwood New and noteworthy western plants. III. A, D. E. Elmer Some littoral spermatophytes of the Naples region - J. Y. Bergen New and noteworthy North American me of Tri- folium (with twelve figures) —- - Homer Doliver House Some studies regarding the biology of site and twigs in winter (with eight figures) - - Karl M. Wiegand The life history of Polysiphonia ioc Contribu- tions from the Hull Botanical matoeg LXXXIII - - Shigeo Yamanouchi The structure. and Sevaoplnsli of the bark in the Sassafras (with nine figures) - - - Howard Frederick Weiss BRIEFER ARTICLES— Notes on ak American Grasses. V. Some Trinius Panicum types CS - A. S. Hitchcock ‘ GE I IIo 373 425 vi CONTENTS [VOLUME XLI PAGE oe in Pallavicinia = - - - - J.B. Farmer 67 - - Andrew C. Moore 69, Notes on she relation between — of roots and of tops in wheat. Contributions from Hull Botanical Laboratory. LXXXI (with five figures) - - - - Edward Burton Livingston 139 New normal icicles for use in = sa aad ogy. III (with two figures) - W. F. Ganong 209 Notes on North American 2 et VI. $55: nopsis of Tripsacum_— - A. S. Hitchcock 294 The basidium of Amanita bsporiger vith seven-_ teen figures) - - Charles E. Lewis 348 The distribution and 1 habits of some common oaks = e ss 3 - - Picks Hill 445 CURRENT LITERATURE - - 71, 144, 214, 299, 353, 448 For titles of books et: see bisdete ar author’s name and Reviews. Papers noticed in ‘‘Notes for Students” are indexed under author’s name and subjects. News 65. eS a OE Oo ig ond ae a DATES OF PUBLICATION. No. 1, January 26; No. 2, March 3; No. 3, March 31; No. 4, April 28; No. 5, May 31; No. 6, June 30. ERRATA. 10, line 19, for (10) read (11). 14,.line 2 from bottom, for nternodes read internodes. 15, line 4 from bottom, for Contribution read Contributions. 75, line 16 from bottom, for East read West. 76, line 4 from bottom, for Perti read Petri. . 167, line 19, for stand read stain. 168, line 13 from bottom, for comma read in. 77, line 1, after was insert in. . 178, line 12 from bottom, for its read their. - 179, line 11 from bottom, after paraphyses insert Ste anameneee - 180, line 3, for The read Two. . 340, line 12, for heller read Heller. - 353, line 3 from bottom, for Petrostemonaceae read Podostemonaceae. - 379, line 13, for Alluim read Allium - lea MaDe a a aa a ea a Bete Paeee pear een Che Botanical Gazette HA Montbly Fournal Embracing all Departments of Botanical Science Edited by Jonn M. CovLtEer and CHARLES R. BARNES, with the assistance of other members of the botanical staff of the University of Chicago Vol. XLI, No. J Issued January 26, 1906 i MR EN Ti TE i a a el . CONTENTS THE NODES OF GRASSES (wirH PLATES I AND It). Mintin Asbury Chrysler - I THE BOGS AND BOG FLORA OF bei BRON, REVERS VALLEY Cigars SIXTEEN FIGURES). Edgar Nelson Transeau : NUCLEAR DIVISION IN ZYGNEMA (with PLATES I AND Iv). Mabel L. Merriman i | VC EFFECT OF CERTAIN SOLIDS UPON THE GROWTH OF SEEDLINGS IN WATER CULTURES (witH rour FiGuRES). J. F. Breazeale - - - - 54 BRIEFER ARTICLES NOTES ON Nor ae ae VY. Some Trinius PAnicum Types. A. S. Hitchcoc - - . - - - - - - - - 64 AC eat IN PALLAVICINIA. J. B. Farmer - - - - - - - 67 REPLY. An C. Moe -- - - - “ -—" - - - - CURRENT LITERATURE. BOOK REVIEWS - - + - - ee ee A EF UR HE Be THE ALGAL VEGETATION OF THE FAEROESE COASTS. ‘ PLANT DISEASES. REGENERATION. PLANT HISTOLOGY. BIBLIOGRAPHICAL INDEX OF NORTH AMERICAN FUNGI. MINOR NOTICES - - - es Snel men, Sew ae Maa ESE 76 NOTES FOR STUDENTS ~- - - - - - - - . - : - - 76 : NEWS - - . . - - - - - - - - - - - - - 80 6 Fal 42 tne tha Wats 1 oe 2 ae Cees + sho FTe ity of Chicago, Chicago, ll. Contributors are requested to write scientific and proper names with particular care, to use the metri System of weights and measures, and in citations to follow the form shown in the pages of het pokincae AZETTE. eparates, if desired, must be ordered i b= fear} of publication. -five separates of origina cles (without cove rs) will be furnished gr dditional copies and ccs (if desired) will be supplied The table below shows th ona cost of separates consisting o oe _ or t The cost may vary from ata and will depend upo ount of work cae press work, paper, bin ding, e etc. Separates euinien half-tones ~_ ost somewhat more than the rates given, the increase depending upon the number of cu nd the saci “of work sedated upon them. 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This policy ie debe HY to the man who wants to protect his fam the same time realize io himself a maberentel: pats ay return on the miums paid by him This is done by the apportionment of dividends every five years. e various options at the end ‘of the five-yeat — oct are exceedingly attractive and ve experience — of the Company shows that business m n and others e end of each five-year period, as the dividend is apportioned, the person ela has the choice © Cash, Reduction of Premium for five years, or a Paid- 4 up Addition to Polic | The Premiums are Fixed and Never Increase- | Policies issued on the Whole Life, Limited Payment ee Endowment plans. Send coupon for free informal”) ae about Five-Year Dividend Policy. NaMG oo isis sce eons te ee, Th P dential! PN i a iene ae SS ee e ru : Insurance Company of America POE Bye i es Occupaties 3.0.5. ep of New Jerstt Sheree eee) cu as a Stock Gompany by the State State — sae of Whole Life, Limited Payment DEN, Prest or Endowment Five-year Dividend Policy i is desired. Home Office, Newark, N. J., JOHN F. F. DRY VOLUME XLI NUMBER 1 BOTANICA (GAZETTE JANUARY, 1906 THE NODES OF GRASSES.! MINTIN ASBURY CHRYSLER. (WITH PLATES I AND II) ALTHOUGH the stems and leaves of grasses have received a good share of attention from anatomists, and the bundles of the internodes are perhaps sufficiently well known, the nodes have been largely neglected. The reason for this may be the supposed difficulty of unraveling the tangle of bundles found in»the node, or the obstacles which the sclerified tissues offer to the preparation of satisfactory sections. Yet the nodes are probably the most interesting regions of the grass stem, for they lack the comparative uniformity of the internodes. Since the application of the celloidin method to hard tissues the difficulties of cutting the necessary serial sections have been removed, so that we are now in a position to know intimately the structure of these critical regions of the stem. The object of the present account is to trace the course of the bundles of the grass stem, and to discuss the significance of certain structures which make their appearance at the nodes, in particular the amphivasal bundles and cambium. The investigation has been confined to forty-five genera, but since these represent the eleven largest tribes and there is a considerable degree of uniformity in structure, the account is believed to represent the family fairly from the standpoint chosen. The salient features may best be brought out by the description of types selected to illustrate certain points. The first to be con- sidered represents a medium condition as regards both taxonomic position and ecological relations. « Contributions from the Phanerogamic Laboratorics of Harvard University. No. 3. I 2 BOTANICAL GAZETTE [JANUARY Avena barbata.—At a short distance above a node the stem pre- sents a simple structure compared with that of many grasses, showing just two circles of collateral bundles surrounding a central cavity. The inner circle of bundles lies at the periphery of the central cylinder, which in this genus is not clearly marked off from the cortex. The bundles of the outer circle lie between groups of cells, which above the leaf-sheath bear chlorophyll; they are considerably smaller than the bundles of the inner ring, and from comparison with correspond- | ing bundles in other genera must be regarded as cortical bundles. In the upper part of a node these assume an amphivasal structure and immediately anastomose with one another and with bundles of the inner circle. But just at this level the structure of the stem is further complicated by the entrance of bundles of the leaf-trace, the course of which must now be described. The leaf-sheath in this species extends a little more than 360° around the stem, and contains, as do most of the genera examined, bundles of two distinct sizes: larger ones, alternating with others which are less than Nalf as great in diameter. These two kinds of leaf-trace bundles have a different course in the stem. The larger bundles undergo a pro- found modification as they enter the stem, as may be seen by com- paring figs. r and 2, in which the magnification is the same. As such a bundle enters the stem it rapidly increases in size, owing to increase in the number of xylem elements. Most of the added elements are tracheids with more or less suberized scalariform or reticulate thickenings, but some parenchyma cells are also present. These elements gradually extend around the sides of the phlcem until this is surrounded by xylem, except a small area on its outer side, which is generally occupied by sclerotic fibres belonging to the group which is so well marked im the sheath (fig. 1). In certain species, e. g., Lolium perenne, these fibres disappear, and the phlcem is completely surrounded by xylem. These bundles evidently must be placed in the amphivasal class. As fig. 2 shows, the phloem — enlarges very little, but the xylem increases so much that the bundle : may be five or SIX times as broad in the middle of the node as it is ~ in the leaf-sheath. These bundles are by far the most conspicuous — objects in a cross-section of the stem at this level. They have 2 spindle shape which is not due to their oblique course, for they slant — 1906] CHRYSLER—NODES OF GRASSES i. very slightly from the vertical and do not dip deeply into the central cylinder. The xylem consists so largely of tracheids running irregu- larly and mixed with parenchyma, that the mass has a considerable resemblance to transfusion tissue. Apparently, this region in a bundle forms an important water-storing organ. A further peculiarity of the bundle at this level is the presence of a distinct bundle-sheath or endodermis,? consisting of cells whose walls are reticulately thick - ened and suberized. As these larger leaf-trace bundles descend through the node, branches from the anastomoses mentioned above extend outward between the leaf-trace bundles, and probably anastomose with these, though the fusion is not so plain as in A. sativa. Below this level the bundles gradually resume the ordinary collateral shape, lose their endodermis, and run down in a single circle through the internode as already described. The smaller leaf-trace bundles also undergo some expansion as they penetrate to the boundary of the central cylinder, but throughout their course they may be dis- tinguished from the larger bundles, not only by their size but by their early turning outward into the cortex and running down to the next node as the cortical bundles described above. Though it is not plain in A. barbata that the larger leaf-trace bundles are joined, soon after their entrance into the stem, by other bundles of the node, in A. sativa and in many other grasses it may clearly be made out that on each flank of the leaf-trace bundle another strand applies itself, swinging through an angle, so that its phloern first joins on, then its xylem. In some species, e. g., Arundo Donax, two or more bundles join on each flank of the leaf-trace bundle. Certain features of the cortical strands are more clearly seen in Panicularia americana. The cortical nature of these strands is unquestioned, for they run quite outside-the central cylinder, in a wide area of lacunar parenchyma (jig. 7). As they reach the upper part of a node they anastomose with one another so as to form a transverse ring or girdle (jig. 8), which, at a slightly lower level, sends branches to the bundles of the central cylinder, forming nearly ‘or quite amphivasal bundles, though some of them very soon resume 2 The term endodermis is here used in the general sense employed by VAN TieGHEM, rather than in the histological sense proposed by other writers. 4 BOTANICAL GAZETTE [JANUARY the collateral structure which is characteristic of all the bundles of an internode. Further down in the node, the bundles of the leaf- trace enter the central cylinder and the smaller of these anastomose_ with branches of the nodal complex, and then turn outward to run down through the internode as cortical bundles. The larger leaf- trace bundles behave as in A. sativa. In Leersia oryzoides it may plainly be seen from a series of sections, that as each of the smaller leaf-trace bundles enters a node, it is joined by two small bundles from the nodal complex, and this rein- forced bundle proceeds downward through the sclerified cortex. Whether the smaller leaf-trace bundles run down through the cortex, or in the outer region of the central cylinder of an internode, cannot in all cases be determined with certainty, for the boundary of the central cylinder is often poorly marked, and the cortex may be a very narrow zone. Or the boundary of the central cylinder may be marked by a narrow sclerenchymatous ring, and the bundles may lie along this, projecting either towards the inside or the outside. But the position of these bundles inside or outside the central cylinder appears to be a matter of indifference, and in either case they pursue a different course from that of the larger bundles. In a general way it may be stated that the smaller bundles of the leaf-trace, after fusing with bundles from the nodal complex, run downward through the next succeeding internode in the cortex or at its inner border, and at the next node below join with the bundles of the central cylinder. Species to which this statement applies are: Zizania aquatica, Leersia oryzoides, Avena barbata, A. sativa, Panicularia americana, P. nervata, Agropyron caninum, Elymus americanus, Triticum sativum. The course of the leaf-trace bundles in the grasses, as here de scribed, differs in several respects from the course of such bundles in other families, even in so closely related a family as the sedges, recently described by PLowMAN (11). VaN TIEGHEM’s second class of cortical bundles (6, p. 751) corresponds the most closely, and is thus described: “Le faisceau médian de la feuille, qui en prend trois, entre directement dans le cylindre central, tandis que les deux latéraux descendent dans l’écorce pour n’entrer dans le cylindre 1906] CHRYSLER—NODES OF GRASSES 5 qu’au noeud suivant.” See further his remarks on the monocoty- ledons. Phalaris arundinacea, like Avena, has the bundles of its internode crowded into an annular area surrounding the fistular pith. As these reach the node they anastomose extensively, and at the same time assume the amphivasal condition, which is shown with especial clearness in the variety variegata, figs. 3 and 4, the latter more highly magnified in order to show the tendency for bundles to form nests of three or more, enclosed by an armor of sclerified fibres. These amphivasal bundles, though abundant in the nodes, are absent from the internodes. Arundo Donax may be mentioned as typical of species having several circles of bundles surrounding a central cavity. As the leaf- trace bundles enter the stem they swell out, though not to so great an extent as in Avena. The xylem completely encloses the phloem, and the usual suberized sheath of cells with reticulately thickened walls becomes visible. Farther down in the node the leaf-trace bundle is joined on each flank by one or more cauline bundles. The bundles of the latter class are provided with a sheath of heavily thickened cells, and some of them appear to pass through the ncde without anastomosing with other bundles, though this condition is rare in the members of the family which have fewer bundles. In accordance with the greater thickness of the solid part of the stem, the leaf-trace bundles penetrate more deeply into the central cylinder than in such genera as Avena, making their general course conform more nearly to the palm type of von Mont. The number of circles of bundles in an internode appears to be dependent on the size of stem characteristic of the species, and to have little value in estab- lishing relationships. Grasses with a solid stem conform even more nearly to VON Mount’s type, for the largest leaf-trace bundles penetrate nearly to the center of the stem, before curving outward and downward toward the periphery of the central cylinder. STRASBURGER (4) has given an excellent account of the course of the bundles in Zea Mays. He distinguishes leaf-trace bundles of five different ranks, and finds that the largest of these penetrate most deeply into the stem, while the smallest merely reach the periphery of the central cylinder. 6 BOTANICAL GAZETTE [JANUARY 4 The increase in complexity of the leaf-trace system over the condi- tion found in Avena, seems to be associated with the greater size of the leaf-sheath in Zea; just as a large stem generally has several circles of bundles, so a heavy leaf has a better developed bundle 4 system. Most of the bundles in an aerial node of Zea are collateral, leaving out of consideration the swollen leaf-trace bundles. STRAS- BURGER (4, p. 348) finds amphivasal bundles at the point of origin of axillary buds and adventitious roots; I have confirmed his obser- vation in the former case. Much larger and more numerous amphi- vasal strands are however to be found in the nodes of the axillary branches bearing the ear of corn. One of these bundles is represented 5 1 in fig. 5. It is only in the leafy part of the branch that these occur, j for in the “cob” the bundles are collateral, with an exceedingly well- . developed phloem, doubtless associated with the transfer of elabo- rated food. The amphivasal bundles of these branches are as usual bundle fusions, and their occurrence in the reproductive axis of a plant showing few elsewhere, seems to be a point of some significance, especially when we consider that Zea is probably a highly organized member of the family. Zizania aquatica merits special attention on account of certain features which may be regarded as primitive, e. g., the six stamens. In an aerial internode a narrow cortex surrounds the hollow central cylinder, and the two are separated by a ring of sclerified cells. Partly imbedded in this ring are a number of small bundles, some of which project into the cortex and on aceount of their origin must be regarded as cortical bundles. All the bundles are collateral, and those inside the sclerotic ring lie at different depths within the central cylinder. As would be expected from the aquatic habitat of the plant, the xylem is reduced; in fact, in some instances, it is represented only by a cavity, and in all cases it has its vessels very slightly lignified. The phloem does not share in this reduction. As the node is approached the bundles at the periphery of the central cylinder anastomose, at the same time becoming amphivasal, and a number of transverse strands join up some of the inner bundles of the stele with one another. At this level the leaf-trace bundles enter the stem; they are of at least three ranks, and of these the largest penetrate into the central cylinder, enlarging on the way; ee eee ee Se te ee ee ee Ee ee ae i a ee Cee ee 1906] CHRYSLER—NODES OF GRASSES 7 owing to increase in the xylem elements, which come to enclose the phloem more or less completely, as already described for Avena. A little further down they are joined on the flanks by several bundles from the internode above, each leaf-trace bundle with its tangle of contributing bundles forming a complex bunch of vascular tissue. A suberized endodermis surrounds the leaf-trace bundles in the node. The second rank of bundles of the leaf-trace also enter the central cylinder, where they are joined by other bundles, but soon return to the periphery of the central cylinder, where they give rise to some of the bundles which run through the next internode below on the border line between the sclerotic ring and the cortex. The smallest bundles of the leaf trace go no deeper than the sclerotic ring, and here fuse with other bundles found in this zone. Thus, the course of the bundles of different rank agrees with what STRASBURGER found in Zea. In the lowest part of the node, the very numerous bundles of the sclerotic ring anastomose and proceed downward, greatly reduced in numbers, while the bundles inside the central _cylinder also become much fewer, owing to completion of the fusion of the large leaf-trace bundles with cauline strands. The basal region of the stem has a cortex which differs from the aerial parts in being much broader and more spongy, on account of the large intercellular spaces. At any of the basal nodes the central cylinder is bounded by an endodermis, consisting of a single layer of rounded cells with suberized walls (fig. 6). Inside this is a narrow zone of vascular tissue, whose elements run circularly; then a wide zone, consisting of small bundles running vertically, and so closely packed together that it is generally impossible to dis- tinguish their limits. Bounding the two vascular rings on the inside is a band of sclerified cells which are in contact with the pith. As the photograph shows, leaf-trace bundles make their way into the central cylinder through wide gaps in this four-layered ring, and it may be clearly seen, even in unstained sections, that along the edges of such a gap the external suberized endodermis is continuous with the inner sclerified layer. All of the large leaf-trace bundles pass through such gaps, but the roots leave the central cylinder without causing a gap, as has been observed in plants of various groups. In the pith of the central cylinder are scattered many 8 BOTANICAL GAZETTE [JANUARY bundles, nearly all of which are amphivasal, and where the nodes are crowded, as they are at the very base of the stem, the amphivasal condition is retained by bundles from one node to another, though in the more elongated internodes, found a little higher in the stem, only collateral bundles occur. The contrast between the upper and the basal nodes is indeed striking, for the former show no amphi- vasal bundles running longitudinally in the pith, except the enlarged leaf-traces. This feature of the aerial node may be partly accounted for by the thinness of the diaphragm in which run the anastomosing bundles, which are generally amphivasal. AMPHIVASAL BUNDLES _ Although these have been repeatedly reported as occurring in the subterranean stems of monocotyledons (see STRASBURGER, 4, p. 348, footnote; DEBARY, 2; SCHULZE, 5)3, the only references to their occurrence in grasses that have been found are by STRASBURGER (4) and Jerrrey (9). Duvat-Jouve (1) figures the rhizomes of many grasses, but shows no amphivasal bundles. Yet an cxamina- tion of the nodes of some of the same species shows that the bundles in question occur here. Two sorts must be distinguished in this family: (1) the swollen portion of a leaf-trace bundle, the xylem consisting largely of a mass of tracheids running irregularly; (2) the type usually figured, the xylem forming simply a ring of vessels. The features of the first class have been described under Avena. Such a bundle is always enclosed by an endodermis which generally has pitted or reticulated walls, and shows, in addition to the ordinary metaxylem elements, a large number of reticulated tracheids, which almost or quite enclose the phloem. All the species examined show these bundles, from hydrophytes such as Zizania to xerophytes such as Ammophila, and there appears to be no relation between the size which a bundle attains at its widest part and the condition under which the species grows. If, as already suggested, these bundles serve to store up water, it might be expected that they would be poorly developed in aquatics, but the only peculiarity of the bundles found in such plants is the slight lignification of the xylem, a char- 3Since the above was written, Horm has reported the occurrence of amphivasal bundles in the rhizome and also the aerial stem of Croomia pauciflora (Amer. Jour. Sci. 20:50-54. 19¢5). Be ee 1906] CHRYSLER—NODES OF GRASSES 9 acter shared by all the bundles of such plants. Quite commonly, the bundles are surrounded by a layer of parenchyma rich in chloro- phyll. This suggests that the node is an active assimilating organ. Bundles of the second class—amphivasal bundles as usually under- stood—are found in the nodes of the great majority of the grasses examined, but are especially numerous in the following species: Coix lachryma, Paspalum stolonijerum, Panicum sanguinale, Sor- ghum halepense, Leersia oryzoides, Phalaris arundinacea, Calama- grostis canadensis, Avena barbata, Panicularia americana, Panicularia nervata, Festuca arundinacea, Triticum sativum. The fact that they are practically confined to the nodal regions, suggests that they are associated with bundle fusions, and this assumption has been amply borne out by observation. Further, since the bundles which fuse are usually traces of leaves which come off higher up, it appears that the occurrence of such bundles is to be referred to the leaves. The closed mode of venation, prevalent in the monocotyledons, involves that a large number of bundles shall run down parallel to one another through the petiole or sheathing base of the leaf. In the grasses the numerous bundles are accommodated in the leaf-sheath, which frequently encircles the stem for somewhat more than 360°. The large number of bundles cannot at once find room in the vascular ring, which we may believe constitutes the primitive stele in both dicotyledons and monocotyledons, according to the results of JEFFREY (8), and CHRYSLER (10). Hence the leaf-trace bundles, or some of them, pass into the inside of the central cylinder, and sooner or later join other bundles. It will be readily seen that a bundle lying in the pith has a better chance to orient itself with regard to some other bundle which it may join, than has a bundle which merely fits itself into a gap in a vascular ring (e. g., the leaf- trace of a fern such as Adiantum). Hence it is not surprising to find that before two bundles of a monocotyledon fuse, they swing around, so that phloem fuses with phloem, and the xylem accordingly surrounds the compound bundle. How far beyond the point of fusion of the bundles this amphivasal condition persists, is a feature which varies greatly. In most of the grasses the collateral structure is soon resumed, but the example of Zizania shows that, at the base of the stem where the nodes are crowded, the amphivasal condition 10 BOTANICAL GAZETTE [JANUARY may continue through several internodes. This probably accounts for the prevalence of these bundles in the rhizomes of monocotyle- dons, where they were first observed. A comparison of the aerial and subterranean nodes of the grasses under study has not yielded 4 results of great significance; most species show no noticeable differ- ence in the number of amphivasal strands in the two cases. But in Andropogon jfurcatus, A. scoparius, Chrysopogon avenaceus, Zizania aquatica, and Phleum pratense, the amphivasal strands are distinctly more numerous in the basal nodes. No examples of the opposite condition have been found. Querva found in Gloriosa (7) that the amphivasal bundles are connected with the origin of a branch. Among the grasses, Phalaris arundinacea, Paspalum — stoloniferum, Sporobolus Wrightii, Coix lachryma, and Zea Mays : show these bundles at the point of origin of branches, but in other species only collateral bundles could be discovered at these places. Too much importance should not be attached to the few cases named, in view of the fact that the sedges uniformly show amphivasal bundles associated with leaves and not with branches, as PLOWMAN has _ shown (10). This is one reason for considering the grasses a more specialized group than the sedges; in fact it may be premised that the amphivasal condition originally connected with leaf-traces has in the Gramineae spread to the branches. The occurrence of amphi- vasal bundles in the leafy reproductive axis of Zea, while they are rare in the main stem, deserves emphasis. While many of the grasses show amphivasal bundles in all the nodes, in this highly | developed genus the bundles in question have nearly disappeared from the ordinary nodes, but .have persisted in the conservative region named. STRASBURGER proposes (4, p. 348) a physiological explanation for the occurrence of these bundles, viz., that the amphi- vasal structure is favorable for the taking up of reserve materials stored in a rhizome, but this explanation is not in accord with the accepted view that elaborated food is carried not by the xylem, but by the phloem. Examination of serial sections leads to the opinion that the mechanical necessities of bundle fusion rather than con- siderations of absorption of food have been the determining factor in producing these bundles. If we accept the view advanced above, that these bundles are to 1906] CHRYSLER—NODES OF GRASSES Ti be associated with the large number of leaf-trace bundles of the monocotyledons, their phylogenetic significance is considerable. The ferns and dicotyledons have a comparatively small number of leaf-trace bundles; amphivasal bundles are absent in the former and rare in the latter group, but are widely distributed in the mono- cotyledons, which accordingly appear to represent a more recent and highly specialized group. In the most highly organized members of the Gramineae, such as Zea, is shown a tendency to reduce the number of amphivasal strands, but even in such cases they may linger on in the reproductive axis. JEFFREY (9) has called attention to the fact that in highly organized families, such as Tridaceae and Orchidaceae, these bundles disappear even from the reproductive axis. CAMBIUM. A feature of the vascular bundles of Avena barbata, not so far mentioned, is represented, in fig. 9, which shows a bundle from the stem at a distance of about 1™™ above one of the upper nodes. That the tissues are immature is shown by the presence of protoplasm and a nucleus in certain of the vessels. The shrunken protophloem is represented by the dark band at the outer edge of the bundle, and the metaphloem as usual has its elements arranged irregularly. Between these elements and the vessels are a number of rows of flattened cells, radially arranged, corresponding well to the cambium of dicotyledons. This feature is not confined to the young stem, as is seen in jig. 10, which represents a bundle from above one of the lower nodes of the same plant. A few tangential divisions are to be seen in the leaf-trace bundle shown in fig. 1, though cambial activity here is slight. Toward the node and farther up in the internode and sheath, this peculiarity is not shown by the bundles, but at the regions mentioned most of the bundles have a more or less clear indication of cambium. In the leaf-sheath of Andropogon argenteum (fig. 11), at a distance of 1-2™™ above its insertion on the stem the larger bundles show an unmistakable cambial layer. From the small amount of phloem or xylem showing radial arrangement of its elements, it appears that the cambium is functional for only a short time. Sections. through the leaf-sheath, cut 5™™ above the one shown in the figure, still exhibit a layer of radially arranged cells exterior to the xylem, 12 BOTANICAL GAZETTE [JANUAR but the cells have thicker walls and a rounded outline, indicating that in that region the period of activity of the cells is past. The stem of this plant does not show good examples of cambium, nor do A. scoparius or A. jurcatus show the feature, even in the leaf-sheath. A further example is shown in fig. 12, which represents a bundle from the stem of Erianthus Ravennae 1-2™™ above the level of inser- tion of the leaf-sheath. A similar appearance is presented by the bundles of the sheath. The leaf-trace bundles of Zizania frequently show a large amount of their phlcem radially arranged, in spite of the fact that dicotyledoncus aquatics generally show a marked reduction of their cambium. The examples so far cited include only the more striking instances of cambium found in the family. More or less plain evidences of cambium have also been observed in the following species: (1) In both stem and leaf-sheath; Coix lachryma, Panicum crus-galli, Avena sterilis, Lolium italicum, L. perenne. (2) In stem; Tripsacum dactyloides, Miscanthus sinensis, Pennisetum longistylum, Panicum sanguinale, Leersia oryzoides, Sporobolus Wrightii, Calamagrostis canadensis, Arundo Donax, Avena sativa, Briza maxima, Panicularia americana, Bromus inermis, Triticum sativum. The occurrence of a cambium in the region just above the nodes in grasses recalls the well-known power which members of this family possess of bending upward at these regions if the stem is by any means laid horizontally. In this connection it is of interest to note that PLowMAN (11) has found only traces of cambium in the sedges, and in line with this, the stems of sedges are unable to right themselves if bent over into a horizontal position. Miss ANDERSSON (3) has reported the occurrence of a more or less plain cambial zone in the young plants of representatives of many monocotyledonous families. She calls attention to the similarity between the bundles in the seedling of Lilium and those in Ranun- culus. The only grass referred to is Zea, in which the mature _ bundles often show a radial arrangement of the cells between xylem and phloem, as is illustrated by the well-known figure in SACHS’ Text-book. From the occurrence of cambium in the tuberous stems of Gloriosa, QuEvA (7) has already concluded that the mono- cotyledons are derived from dicotyledons. In the case of the grasses it would seem that the cambium possessed by the ancestors i 4 1906] CHRYSLER—NODES OF GRASSES 13 has been retained in the regions where it is of use. On the other hand, it may be argued that we have here the first appearance of a feature which in the dicotyledons becomes prevalent. But why should the cambium appear only at the nodes, where it is of use ? It may be regarded as almost axiomatic that the need for a struc- ture is not a sufficient cause for its appearance. So it seems more reasonable to read the evidence in the way first suggested, viz., that we have here a relic of a structure which was present in the ancestors of the grasses, but has disappeared from most parts of the plant and from most families of monocotyledons, and_ is retained above the nodes of grasses in connection with their power of bending at these regions: Thus the evidence favors the derivation of the grasses from ancestors having a cambium. The stele of the grass stem has evidently departed widely from the primitive protostele or siphonostele. It has been repeatedly shown that reproductive axes are able to retain ancestral characters. An examination of this region in seventeen species of grasses belong- ing to seven of the tribes has failed to disclose any instances where the stele presents the primitive type described by PLOwMAN (11). This result seems to confirm the opinion derived from other con- siderations, that the Gramineae represent a more specialized group than the Cyperaceae. These considerations may now be stated categorically: (1) The grass family has adapted itself to every habitat, from salt marsh to pampas, and shows every gradation in habit from the bamboo downward. The sedges are prevailingly hydrophytes, and few of them attain a considerable size. (2) The hollow stem characteristic of most grasses has probably been derived from a solid stem such as is present in the sedges and most monocotyledonous families. (3) Amphivasal bundles are not found in so large a proportion of species nor are. they as numerous in an individual in the grasses as in the sedges. (4) In practically all of the grasses the leaf-trace bundles are of at least two ranks, while the sedges show no such distinction. (5) The floral axis of the grasses dees not present the simple type of stele shown by some sedges. 14 BOTANICAL GAZETTE [JANUARY On the other hand, the open sheath of the grass leaf may be con- sidered to be more primitive than the closed sheath of the sedges. Further, the cambium found in the grasses is here considered to be a primitive feature. But on the whole the Gramineae are to be _ regarded as a more highly specialized family than the Cyperaceae, though the families are evidently very closely related. Certain anatomical features of the grasses, such as the distribu- tion of the amphivasal bundles, seem to have an important bearing on the phylogenetic position of the family among monocotyledons, but since the anatomy of the group is as yet very imperfectly known, a discussion of this point would be premature. SUMMARY AND CONCLUSIONS. 1. The grasses depart considerably from the scheme propcsed by von Mout for the course of the bundles, chiefly owing to the stem being hollow in most cases. The leaf-trace bundles are of at least © 4 j two ranks; of these the largest penetrate most deeply as they enter the central cylinder, generally receiving one or more bundles on each flank as they pass downward to the lower part of the node; the smaller leaf-trace bundles do not penetrate deeply into the central cylinder, but after anastomosing with other bundles pass downward either in the cortex or at the inner border of this. At the next node lower, these cortical bundles anast mcse with one another, and then with the bundles of the central cylinder which have come from fusions ‘with leaf-trace bundles at the next node above. 2. The leaf-trace bundles, especially the larger ones, undergo a marked change as they enter the stem. This consists in the appear- ance of a distinct endodermis, and in an increase in the xylem, leading to the formation of a greatly swollen amphivasal bundle. Below the node these bundles resume the collateral type. 3- Amphivasal bundles of the ordinary type, though absent in ihe ce internodes, are very commonly found in the nodes, and arise by fusion of collateral bundles which are generally leaf-trace — bundles. In some species they are more numerous in the nodes at the base of the plant, and where such nodes are crowded, the bundles may retain the amphivasal condition through successive internodes. The presence of amphivasal bundles in reproductive branches of plants in which these bundles are scarce in ordinary — aa 1906] CHRYSLER—NODES OF GRASSES 15 nodes, points to their being an ancestral feature, which, in highly organized members, has disappeared from most parts of the plant, but is retained in the conservative flowering axis. It appears that the amphivasal bundles so characteristic of monocotyledons, in all probability made their appearance in connection with the entry of numerous leaf-trace bundles into the nodes, but that secondarily, in certain instances, they are found to be related to branching. 4. A well-marked, though generally short-lived, cambium occurs in the bundles just above the node or near the base of the leaf-sheath in certain grasses. This fact is considered to lend support to the view that monocotyledons have been derived from some group possessing a cambium, probably the dicotyledons. 5. The anatomical features of the grasses point to their being a more highly specialized family than the sedges. This investigation has been carried on in the Phanerogamic Laboratories of Harvard University. I am indebted to Professor G. L. GoopateE for material, and to Professor E. C. JEFFREY for material and for advice during the progress of the work. HARVARD UNIVERSITY. LITERATURE CITED. Norte.—Practically nothing bearing immediately on the subject of this research has been found in the older literature, so it is not cited here. References to it may be found in DEBAry (2), and the text of Kny’s Wandtafein. 1. Duvat-Jouve, M. J., Etude anatomique de quelques Graminées. Mém. Acad. Sci. Montpellier 7: 309-406. 1 2. DeBary, A., Comparative anatomy of the phanerogams and ferns (trans.). 1884. ; 3. ANDERSSON, S., Ueber die Entwickelung der primaren Gefassbundelstrange der Monocotylen. Review in Bot. Cent. 38:586, 618. 1889 4. STRASBURGER, E., pce den Bau und die Verrichtungen der Leitungs- : nen. Jena. 1 5. Scuutze, R., sieht zur vergleichenden Anatomie der Liliaceen, Haemo- doraceen, Hypwicidoak: und Velloziaceen. Bot. ‘Jahrb. 17: 295-394. 1893 6. VAN TiecHEM, Ph., Traité de Botanique. Paris. 1891. 7. Queva, C., Contribution & l’anatomie des monocotyledonées. I. Tra- vaux et Mém. Univ Lille VII. 22:1-162, pls. 1-11. 1 8. Jerrrey, E. C., The morphology of the central cylinder in the angiosperms rans. Can. Inst. 6:1-40. pls. 7-11. 19920. 16 BOTANICAL GAZETTE [JANUARY 9- re E. C., A new key to the phylogeny of the monocotyledons. Science N. 5. 37: eee. 1903. 10. CHRYSLER, M. A., The development of the central cylinder of Araceae and Liliaceae. Bor. GAZETTE 38: 161-184. pls. 12-15. 1904. 11. PLowMan, A., The comparative anatomy and phylogeny of the Cyperaceae. Annals of Bot. (ined.) vol. 20. EXPLANATION OF PLATES I AND II, PLATE I. Fic. 1. Avena barbaia; leaf-trace bundle in the leaf-sheath 1™™ above its insertion on the stem. 130. 2. Same; swollen and amphivasal condition of a leaf-trace bundle in the middle region of node. X 130. Fic. 3. Phalaris arundinacea; section through upper part of a node; near the outside are the leaf-trace bundles, alternating large and small; just internal to these are the nests of amphivasal bundles. X30 1G. 4. Same; one of the nests of amphivasal bundles more magnified; on three ie are leat bic bundles. X 150. Fic. 5. Zea Mays; one of the large amphivasal bundles from a node of x reproductive branch. 115. Fic. 6. Zizania aquatica; part of stele and cortex in basal region of the stem. Two gaps with their leaf-trace bundles are visible. 35. PLATE II. Fic. 7. Panicularia americana; section 1™™ above node, showing two cortical sae X45. ame; section from upper part of the node, just above insertion of Kskakoa. The cortical bundles are connected by a ring-shaped anastomosis. x45. Fic. 9. Avena barbata; bundle from a young stem 1™™ above insertion of the leaf-sheath, showing cambium. X 150. IG. 10. Same; bundle from the same position in a mature stem. 150. Fic. 11. Andropogon argenteum; bundle from the leaf-sheath 1-2™™ above its insertion on the stem, showing cambium. X 150. Fic. 12. Erianthus Ravennae; bundle from the stem a short distance above anode. X150. et Spe 5 eae Sen BOTANICAL GAZETTE, XLI “ O80 Aan eave Bg A SSagket BS EL A €99%3%E.689'S be ? ind {.¥ $664 HA hd “9 A A Osaat wcteg eS 8@>': OH L) srstai tons Sc © # é SLRS OA Ta N CARED he 23,085 « 172, Osha yereee sue a,, ’ CR OT ad pak ae OO e4 pri tet 0) rat FG S ~Ps Fi te fe © eae as poe, oo. to. CAs BMse eo Gasce a. oa ~~ ae 8, eae: p Aad Se tf 2. a: a © gs % is ‘ ‘ al : in ek * » é . Ae a e.: A ee D4 « e.. @*# M. A.C. photo. PLATE J \) hl AN) Ua ay. oe Rat SNR > . Gat ret ¥ Ye >. ‘ a wh | WA (¥ i 6 . ai tree ee Bales © yg =¢ Baynes 7 Fa Epes CHRYSLER on NODES OF GRASSES BOTANICAL GAZETTE, XLI PLATE if <2 av)” ' Pe A tees, 8a, ' wy y i ~, afte ACTY Oe ey f ey! Py MING TCR Poe € e* ws Dew): oe eer, & SES 504 a M. A.C. photo. CHRYSLER on NODES OF GRASSES THE BOGS AND BOG FLORA OF THE HURON RIVER VALLEY. EDGAR NELSON TRANSEAU. (WITH SIXTEEN FIGURES) [Concluded from p. 448.] IV. The ecological characteristics of the bog flora and their causes. The plants occurring in the bog habitat are almost all perennials. In the case of the herbaceous vegetation, the winter is passed by means of subterranean rootstocks. The shrubs are in part evergreen and in part deciduous. The tamaracks and the two birches are deciduous, and the black spruce and pine are evergreen. Most of the herbaceous and shrubby forms multiply abundantly by vegetative shoots of one form or another. The length of the underground stems of the shrubs is proverbial, but is best appreciated by one who has attempted to dig up one of them entire. In con- nection with the competition between species for space in the habitat, this is of the greatest importance. A luxuriant growth of cassandra furnishes the most :favorable situation for the development of sphag- num in this vicinity. Its profuse branching affords a framework for the upbuilding of the sphagnous layer, its shade properties do not interfere with the photosynthetic work of the moss, and it protects ‘it from the drying effects of wind and direct insolation. Where such associations occur, the difficulties presented for the germination for most seeds, and the efficiency with which competition is combated, are evidenced by the fact that among the tree species only the tama- rack, spruce, and pine are successful invaders. All of these plants send out adventitious roots from the stems and branches, and so keep pace with the upward development of the moss. The absence of poplars, willows, red maples, and elms in such undisturbed situations must be in part attributed to the completeness with which such terri- tory is controlled by the cassandra-sphagnum association. ECOLOGICAL ANATOMY. Aside from the purely aquatic forms which have received much Botanical Gazette, vol. 41-] [17 18 BOTANICAL GAZETTE [JANUARY ecological attention, it is of interest to look at the anatomical char- acteristics of certain of these plants. Eriophorum virginicum may be taken as a type of this group, and also of the sedge zone vegetation in general. The culm is very slender and erect, leaves flat, and very narrow, perennial by root — stocks. Stem: epidermis very thick-walled and cuticularized. As development proceeds, certain radial rows of the primary cortex cells have their walls thickened, and served to connect the tissues of the central cylinder with those of the three-or four outer layers of hypodermal cells which also become thick-walled. Between these radial groups of cells lysigenic air cavities are formed. Root: epi- dermal cells in part thin-walled and in part secondarily thickened, no definite arrangement of the thick-walled cells apparent; internal structures closely resemble those of the stem; no mycorhiza present. Leaj: outer epidermal cell walls strongly thickened and cuticularized, radial and inner walls less so; lysigenic air spaces traverse the leaf longitudinally; a very thick layer of stereome adjoins the leptome, decreasing to one or two cell layers on the hadrome side of the bundle; chloroplasts massed among the outer layers of the cortex, but occur throughout. Sarracenia purpurea.—Well known for its insect-capturing pitchers. Stem: epidermis and first hypodermal layer thick-walled; lysigenic air cavities throughout pith and cortex; resin deposits confined to the epidermis and one or two hypodermal cell layers, but where wounded heayy deposits of resin take place in the exposed and underlying cells. Root: cell walls firm, resinous bodies present — throughout, but especially prominent in the two outer cortical layers, in which the cell walls are also strongly thickened. Leaj: epidermis thick-walled and, slightly cuticularized; stomata on both sides of the lamina, with guard cells strongly cuticularized and slightly protuberant; resinous deposits throughout; inner face of lamina with strong downward pointing bristles. Oxycoccus macrocarpus.—Stem: pith thick-walled, with resinous bodies; a thick layer of broad-celled bast forms a complete cylinder within the epidermis. Leaf: margins revolute, upper epidermis without stomata, heavily cuticularized, radial walls thick, wavy; hypodermal collenchyma of two or three cell layers on leptome side 1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 19 of midvein, one or two cell layers on the side of the hadrome, develop- ment of the stereome cells also smaller on hadrome side; palisade of two cell layers; lower epidermis covered with wax, especially at * the stomata, guard cells slightly elevated. Mycorhiza present in the larger roots, wanting in the hairlike branches, no root hairs. - Andromeda polijolia—Leaf: margins revolute, upper epidermal cells thick-walled, radial walls undulate, no stomata; lower epidermis supplied with unicellular short stiff hairs, and covered with wax, stomata slightly protuberant, strongly cuticularized beneath mid- vein; palisade of three layers of long narrow cells; stereome strongly developed above and below vascular bundle; on the ventral side this adjoining three layers of large thin-walled air cells and a one- | layered hypoderma. Root: resinous deposits throughout, no mycor- hizal fungi found. Chamaedaphne calyculata.—Leaj: margin slightly revolute, epi- dermis thick-walled, heavily cuticularized, cuticle rough, no stomata on upper surface; ventral epidermis covered by shield-shaped multi- cellular hairs, and a deposit of wax; cuticle unusually thickened beneath the midvein, guard cells sunken, subsidiary cells protuberantr palisade tissue of four or five layers. Root: inner and radial walls thickened, cortical tissues thick-walled; resin deposits in vascular bundle and cortex; no mycorhizal fungi found. Chiogenes hispidula—Lea}: margin revolute, epidermal walls very thick, cuticle present, papillate, palisade not strongly developed; mesophyll cells in part thick-walled and in part thin-walled; resinous bodies in the epidermis; stomata slightly protuberant. Stem: resin Present in cortex; mycorhizal fungi in the epidermis of the smalle; Toots and throughout the cortex of the larger. Vaccinium corymbosum.—Leaj: cuticle present, epidermal walls hot thickened, palisade of one layer, mesophyll tissues with resinous bodies, cuticle of ventral surface papillate; abundant unicellular irs on lower epidermis few on upper; leptome side of mid-vein adjoined by three layers of stereome and two or three layers of hypo- dermal collenchyma,.on the hadrome side reduced to two of stereome and two of collenchyma, cuticular papilli usually developed beneath the midvein and at edge of leaf. Root: cortical tissue with resin, mycorhiza present. No resin deposits found in stem. 20 BOTANICAL GAZETTE [JANUARY Salix sericea.—Leaf: upper epidermal cells small, strongly cuticularized; mesophyll compact, palisade of two layers of long narrow cells; stomata on under surface, guard cells sunken beneath the slightly protuberant companion cells; hypoderma of five- or six-cell-layers on hadrome side, and eight layers on leptome side of midvein. Root: resinous bodies present in medullary rays and cortex, the latter consisting of thick-walled cells; no mycorhiza. Ledum_ groenlandicum.—Leaj: upper epidermis rugose, with scattered unicellular hairs, margins strongly revolute, cuticle present, cell walls thickened, the radial walls being broadly undulate; lower epidermis covered with a thick cuticle and a felt of long multicellular and short unicellular hairs, glandular hairs usually present near the small veins, stomata protuberant; palisade of three or four layers of broadly oblong cells; beneath vascular tissue of midvein and between the mestome bundles occur large air.cells which may form lysigenic air cavities in the older leaves. Root: mycorhizal. Larix laricina—Leaf: bifacial, deciduous; epidermis thick- walled, slightly cuticularized, guard cells sunken beneath the com- panion cells; palisade tissue developed toward the dorsal surface, two layers thick showing a radial tendency, stereome reduced to a few cells beneath the leptome; two resin ducts near edges of leaf. Root: composed of mycorhiza, resinous deposits throughout, cortical tissues early destroyed by fungus. When grown in culture solutions and well aerated, normal roots with root hairs are produced. Picea Mariana.—Plants in bogs are stunted. Leaj: epidermal cells thick-walled, cuticle present, guard cells sunk beneath the companion cells; mesophyll cells compact, of a more or less radial palisade type. Root: mycorhizal, resin deposits throughout, cortical tissues destroyed by ieee Normal roots are developed under culture conditions. Pinus Strobus.—Plants very much stunted-in the bogs, leaves shorter and thicker. Leaj, epidermal walls so greatly thickened that scarcely a lumen remains, beneath this a hypodermal layer of thick-walled cells; mesophyll cells compact and of the usual lobate type. Root: mycorhizal, cortical tissues traversed by the fungus hyphae; resinous deposits throughout. Stem: annual rings narrow Sr 1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 21 and distorted, resin bodies throughout cortex and meristematic tissues of the wood. To summarize these characteristics, it is evident (1) that epidermal and hypodermal tissues are thick-walled; (2) that for the ccnserva- tion of water these are reinforced outwardly by a heavy cuticle, by coverings of wax and air containing hairs; (3) that resinous bodies are found in the roots and leaves of many of the plants; (4) that there is a general reduction in the size of the leaves, and that these are frequently revolute-margined; (5) that palisade tissue is quite uni- formly developed; (6) that mycorhizal fungi are present in the roots of most of the plants; (7) that, when compared with the xero- phytes of dry sand plains (25, 6), they show a similarity in respect to the reduction in size of the foliage, in the development of external protective coverings of the sub-aerial parts, and in the presence of palisade tissues, but are very different in the matter of root develop- ment and character of root structures. To account for the peculiarities of the bog vegetation various theories have been brought forward. KraiMman (28), in accounting for the xerophilous character of the plants of arctic swamps, which include several species common to American bogs, lays stress upon two factors: (1) the low temperature of the .moist substratum, and (2) the presence of drying winds. The former influences the plants by decreasing the power of absorption, the latter increases the rate of transpiration. The plants of such habitats must therefore be protected against the loss of water by the subaerial parts. SCHIMPER (44, p. 11) in classifying the natural habitats in which xerophytes occur mentions among others “peat, bogs, because of the humous acids in the soil.” On page 18 he says: _ The xerophilous character of the vegetation of peat moors has hitherto been considered an incomprehensible anomaly, and yet the rich supply of humous acids in the soil furnishes a condition for its occurrence as cornprehensible as it is necessary. The presence of Scotch pine-and heather on both dry sand and on Wet peat is thus not more remarkable. than is that of Ledum palustre, Vaccinium uliginosum, and other peat-plants on the cold dry soil in the polar zones. Further (p. 124) the statement occurs that ‘‘on the very acid humus of moors the vegetation assumes a decidedly. xerophilous character, because . the. humous,.acids impede the. absorption of water by the 22 BOTANICAL GAZETTE [JANUARY roots.” However, in describing the arctic vegetation (44, pp- 11, 715), he follows the suggestion of KrHLMAN, a conclusion to which he had come independently. GANONG (16) also accepts KIHLMAN’S explanation for the xerophilous nature of the raised-bog flora of New Brunswick. | In the study of the structural adaptations of these plants and the causes of their occurrence in bog areas, several questions arise. Are these two factors, cold substratum and acidity, efficient causes of xerophily ? Do they act, in the case of the bogs of this region, with sufficient strength to cause xerophilous modifications in the plants there found, or to permit the growth of only such forms as are xero- philous ? The last question may be answered from field observations. They indicate that most low-ground plants grow quite as well on the bog substratum as on the ordinary swamp soils, and that the swamp species of this vicinity may all be found at one place or another grow- ing on bog soils. It would seem that here the bog substratum is no more efficient as a selective agent than are the swamp soils. The only cases which have come under my observation in south- ern Michigan which will throw light upon the question of the effect- tiveness of the temperatures and acidity in the production of xero- philous adaptations is in the case of Picea Mariana? and Pinus Strobus. These two plants both show reduced size of stem and leaf, in the Oxford bog, when compared with plants growing on the margin. But to what extent this may be due to sterility of the bog substratum rather than to temperature and acidity is indeter- minable at this time. . EXPERIMENTS. To answer the question of the efficiency of a cold substratum and soil acidity to produce xerophily, experiments have been in progress for approximately two years. The difficulties in the way of experimentation along these lines are numerous. The means for controlling soil temperatures in bodies of soil sufficiently large for experimentation with the larger bog plants are practically beyond the possibility of a university laboratory. When it is further realized ons The so-called P. brevifolia Pk. This form is certainly no more deserving of a distinctive name than is the bog form of the white pine. 1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 22 that the experiments should extend over a series of years in the case of the shrubby forms, the problem becomes still more com- plicated. Cold bog water Cold nutrient solution Warm nutrient solution Warm bog water Pic. 12.—Average plants from the several cultures of Indian corn, From photo- graphs. ‘ In order to test the relative effects of humous acids (of the con- centration found in the bogs of this vicinity) and low substratum temperatures, experiments were made in the form of water cultures 24 BOTANICAL GAZETTE [JANUARY and with a peat substratum. All of the bog water used was brought to the plant house from the First Sister Lake. The acidity of the water varied from .o005 to .co23 normal acid, as measured by 7 / 100 KOH solution. WATER CULTURES.—(1) The plants were grown in four-liter battery jars covered with a plaster of Paris plate, having five one-inch open- ings for the passage of the plants and one of smaller size for a ther- mometer. Four such jars were employed in each experiment, two containing a 0.2 per cent. Knop’s solution, and the others bog water. One of each was further maintained at a lower temperature. The cooling was accomplished by passing tap water through 15 feet of quarter-inch (4.5"xX7™™) glass tubing, arranged in a coil within the jar, somewhat below the surface of the liquid. The sides and bottoms of the jars were covered with black paper, and those which were to be cooled were further surrounded by white paper and sphagnum. Daily readings of the temperatures of the air, warm-water solutions and cold-water solutions during the warmest period of the day were recorded. In this way the maximum differences between substrata and air were obtained. As these temperatures were not constant they exaggerate, to a slight degree, the average differences in temperature. Thus, four conditions were obtained which are comparable: (1) warm nutrient solution (temperature approximat- ing that of the air of the plant-house), (2) warm bog solution, (3) cold nutrient solution, and (4) cold bog solution. » Fig. 12 shows the results of one of these experiments with corn. The photograph was taken eighteen days after the experiment was started. When the cultures were set up, the plumule had developed to a length of 2 inches (5°™). The air temperatures during the period of experimentation averaged 18.8° C., that of the warm cultures 18.7° C., and of the cold cultures 10.8° C. - It is to be noted that under these conditions the best growth of the leaves and roots occurred in the bog water. But a reduction of 8° in the substratum temperatures caused a diminution in the devel- opment of both leaves and roots; the plants in the nutrient solution and the bog water being equally affected. When all of the plants had developed five leaves, it was noted that in the case of the cold cultures the two lower leaves had withered. This experiment was 1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 25 repeated with corn, white lupine, and bean under similar conditions, with similar results. The greater development of roots in the case of the warm bog water may be due to the presence of a poison in very minute quantities; but this I have been unable to prove. (2) A third culture was then made in which five plants of corn were grown in each of the four water culture conditions, and in addition in four similar conditions, using a mixture of sphagnum and peat for the substratum. Wooden boxes 2 feet long, 1 foot wide and a half foot deep (60 X30 *15°™) were constructed, and two were lined with galvanized iron. The bottoms of the unlined ones . were perforated so as to allow of easy drainage. The lined boxes served for the undrained conditions. Further, in one of the drained and in one of the undrained boxes, 40 féet (12™) of glass tubing, bent into coils, the joints: ‘being connected» by rubber tubing,were arranged so that a constant flow of cold water, for lowering the temperature, could be maintained. The water level in the undrained bog substratum was kept just below the surface. The water was obtained from the bog at First Sister Lake, but occasionally all were watered with distilled water. The amount added to each box was practically the same. In order to keep the solutions in the water culture jars at the same acidity as in the undrained boxes, the water was siphoned off and transferred once a week. Care was taken in this — transfer to aerate the water in the boxes as little as possible, while that of the jars was aerated at irregular intervals by means of a bulb. There were thus produced eight conditions, in which it was possible to test the effect of the acidity of the bog water, of aeration (drainage) of the substratum, and of low temperatures. As a result, it was found that the growth of roots and leaves was best in the warm bog water, in the warm nutrient solution, and in the drained warm peat substratum. Reduction in size of both roots and leaves occurred in the cold bog and nutrient solutions, and in the drained cold and undrained warm and cold peat substrata. But the plants in the undrained cold peat showed the most marked reduction in size. The conclusion was reached (1) that humous acids (acidity varying from .coo5 to .0023 normal acid) have no effect upon corn in the matter of leaf and root development; (2) that low temperature and lack of aeration of the substratum both cause reduction in size; 26 BOTANICAL GAZETTE [JANUARY and (3) that when low temperature is combined with poor aeration the effect is very marked. This experiment was repeated with peas, and the same result was obtained, although the effects were not so marked (jig. 13). The roots in the undrained substrata were killed when they attained a depth of a half inch (12™™) below the surface. (3) In order to test the effects of drainage and of low temperature on bog species, another set of cultures in peat-sphagnum substrata was made. The apparatus used consisted of two flower-pots and . two glass dishes aproximately a foot in diameter and three inches Fic. 13.—Effect of the several conditions upon the development of pea seedlings. All are average specimens. From photographs. deep (30X7.5°™). ut BOTANICAL GAZETTE [JANUARY BIBLIOGRAPHY. . Battey, V., Tamarack swamps as boreal islands. Science, N. S. 3:250. 1896. Barsour, E. H., Report State Geologist of Nebraska 1: 193-198. 1903. Betz, J. O., Influence of temperature on the rate of nitrification. Ann. Rept. Wis. Agric. Exp. Sta. 18:224. Igor. BLanck, E., On the diffusion of water in humus soils. Landw. Vers. Sta. 58:145. 1903. Review, Exper. Sta. Rec. 14:848. 1903. . BiatcHiey, W. S. and AsHuey, G. H., The lakes of northern Indiana and their associated marl deposits. asth Ann. Rept. Dept. Geol., Indiana. gol. . Britton, W. E., Vegetation of the North Haven sand plains. Bull. Torr. Bot. Club 30:571. 1903. CoutTeEr, S. M., An Pee si of some typical swamp areas. Rept. Mo. Bot. Gard. 15:4 . Cowres, H.C; een peste of Chicago and B eeeh i Bot. AZ, 31:145. I9o!. Davis, C. A., A contribution to the natural history of marl. Jour. Geol. 8: 485. Igoo. , A second contribution, etc. Jour. Geol. 9: 491. 1901. . Davis W. M., Elementary meteorology. Ginn & Co. 1 DeCortots, E. G., Investigation of the composition of soils rich in vege- table matter. 29th Ann. Rept. Ont. Agric. Coll. 1 993. . Dryer, C. R., Lessons in physical geography. Amer. Book Co. rgor. Dumont, M. J., Les causes d’infécondité des sols tourbeux. Compt. Rend. Acad. Sci. 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Snyper, H., Report on composition of muskeag soils. Bull. 81, Minn. Agric. Exper. Sta. 1903. 49. Sraut, E., Ueber den Einfluss des sonnigen oder schattigen Standortes auf die Ausbildung der Laubblitter. Jenaische Zeitschrift fiir Naturw. 1883: 16. BOTANICAL GAZETTE [JANUARY . StocxpripcE, H. E., Rocks and soils. J. Wiley & Sons, New York. 1895. . Srupart, R. F., The climate of Canada. Scot. Geog. Mag. 14:73. 1898. ~ Tarr, R.5., The A es geography of New York state. Macmillan Co. New Mark, ~ ravior, F. B:, pea ee of Erie-Huron beaches with outlets and moraines in southern Michio. Bull. Geol. mer. 8:31. 1897. Topp, J. E., The moraines of southeast South Dakota and their attendant evades Bull. 158, U. S. Geol. Surv. 1899 . TRANSEAU, E. N., On the geographic distribution and ecological relations of the plant societies of northern North America. Bot. Gaz. 36: 401. 190 , The development of palisade tissue and resinous deposits in leaves. Sejm N. S. 19:866. 1904. WAGNER, G., Observations on Platygonus compressus LeConte. Journal Geol. 11:777. 1903. . WesER, C. A., Ueber die Moore, u. s. w. Jahresbericht der Manner vom Morgenstern 3:1-23. Review, Bot. Cent. 88:17 WELD, L. H., A peat bog and morainal lake. ak GAZ. 37:39. 1904 - WHEELER, H. J., Results of many experiments on “acid upland soils” are to be found in the 6th, roth, and 12th Ann. Rept. of the R. I. Agric. Exper. Sta. 1893, 1897, and 1899. Also see bulletins no. 46, 47, 49, 66, 69, 71. go and 95 of the same station. - Wottny, E., Die Zersetzung der organischen stoffe. Heidelberg. 1897. NUCLEAR DIVISION IN ZYGNEMA. MABEL L, MERRIMAN. (WITH PLATES III AND Iv) THE species of Zygnema chosen for this investigation possesses a nucleus unobscured by chromatophores, and hence one in which division stages can be easily followed. No zygospores were found in the material, so the species could not be identified with any degree of accuracy. The number of pyrenoids are normally two, one on each side of the nucleus. The material was gathered from the same locality, the margin of a brook, during the months of August and September of two successive years. The filaments were studied in a living condition to make sure of the presence of dividing nuclei, and were then killed in chromacetic acid and the weaker solution of Flemming for later study. The greater part of them were killed in the evening, as it was also desired to secure division stages of other Conjugatae, which grew in great abundance in the locality and have been reported by investigators as dividing more actively at night. Of these, three species of Spirogyra and two of Mesocarpus will furnish the material for a later contribution. As nearly all the literature upon the cytology of the Conjugatae relates to forms of Spirogyra, its consideration will be deferred until the completion of further studies in the nuclear division of the group. It is hoped then to bring into accord all the observations as to the character of chromatin and nucleoli. Filaments of Zygnema treated with the combination stain of safranin and gentian violet, were found upon examination to have retained the violet only in the cell sheath, while the nuclear structures and pyrenoids retained the safranin. Various results were obtained with those treated with Heidenhain’s haematoxylin in combination with iron alum and eosin. As the same length of exposure to the stain did not suffice for Spirogyra and Mesocarpus growing entangled with the Zygnema, the material was allowed to remain in the staining fluids for a shorter or longer time. Filaments show pyrenoids stained black by the haematoxylin, the nuclear structures retaining the eosin; or the pyrenoids may be stained red by the eosin, and the 43] [Botanical Gazette, vol. 4x 44 BOTANICAL GAZETTE [JANUARY nuclear structures black by the haematoxylin; or finally both may appear stained red by eosin. Such differences are shown in the drawings from the different preparations; the parts shaded in black represent portions stained by the haematoxylin, as in jig. 33, those in gray the portions stained by the eosin, as in jig. 13. Within a quiescent nucleus situated between the two pyrenoids thus stained, there can be seen a central body stained somewhat redder or blacker, as the case may be, than the peripheral network of granules. This network of granules, ordinarily scarcely distin- guishable from the cytoplasmic reticulum, was found in some cases to be quite conspicuous. If an examination is made of a nucleus in process of reconstruc- tion from the telophase, within the forming membrane can be seen a conglomerate mass of substance, very evidently non-homogeneous both in surface view and as seen in outline, figs. 1, 39. Around this smaller bodies can be seen in the meshes of a delicate network. The staining capacity of the larger mass and the small bodies varies in the different preparations; in some instances they are sharply defined from one another, at other times they retain the same kind and amount of stain. It cannot be denied, however, after a careful examination of stages preceding the appearance of these bodies, that the substances in both came from the chromosomes of the metaphase. Bearing in mind, then, that the large mass and the smaller granules have the same origin, it would hardly seem correct to discriminate between the two, terming the one nucleolus and the others chromatin granules. Neither method of staining nor study of their history yields evidence other than that they are of similar substance, differing only in position ._ and aggregation. It is as if in the revolutions going on within the cell some of the chromatin granules had been drawn to the center, there incompletely cohering, while others were left at the periphery. In describing, then, the quiescent nucleus of Zygnema it seems prefer- able to say that the larger portion of the chromatin granules cohere to form a central body analogous in its position to the nucleolus of higher plants. The division of the nucleus is presaged by granules collecting in the region where the cell wall will form. The activity of these vibrat- 19c6] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 45 ing granules in the living cell renders the nuclei about to divide easily distinguishable from the remainder in the filaments. Owing to the activity of these granules, changes going on within the living nucleus could not be easily followed, but changes in the form and position of the nucleus together with those of the pyrenoids were followed throughout division. Accordingly the history of changes in the chromatin is all deduced from comparison of dividing nuclei stained by haematoxylin or safranin as outlined above. If haematoxylin in combination with iron alum could be considered as an infallible criterion for distinguishing chromatic from achro- matic material, and stages could be selected from material stained by one of the methods only, it would be an easy matter to trace the history of this central body originating from the chromosomes of the metaphase. Often, as in fig. 2b, numerous deeply stained bodies are to be seen lying in the space surrounded by a membrane, with no trace of chromatin bodies without. In the nucleus represented in fig. 5, in place of the central body several smaller bodies can be seen marked off from the eosin-stained bodies by the blackness of the stain. Passing to fig. 9, where the beginnings of an intranuclear spindle are manifest, and where there are several more deeply stained bodies, and then to jig. 12, where six discrete bodies distinctly form an equatorial plate, the natural conclusion, based wholly upon simi- lar staining properties, would be that the central mass of chromatin alone furnishes the chromosomes for the equatorial plate. Such was the conclusion reached during the first year of this investigation, but further study of the material shows it to have been premature, or, if applicable at all, only to a few cases. The conviction that _ difference in staining of nuclear structures is more often a matter of manipulation than of chemical reaction, and that difference in the shade produced by the stain is merely due to the density of the body and time given for penetration, renders necessary in interpretation a great degree of caution. The following account is derived from a comparison of parallel Stages in all the preparations. As the nuclei pass from the quiescent to the active state, the cen- trally lying mass disintegrates into small bodies (figs. 2, 3); at the same time the granules lying at the periphery increase in size. The 46 BOTANICAL GAZETTE [JANUARY space within the nucleus becomes gradually clearer (jig. 5), the nuclear sap probably reinforcing the substance of the granules. As the result of the disintegration of the central body and the growth of the other granules, there may be seen lying within the nucleus twenty or more granules (jigs. 4, 5, 6). In a few cases these bodies may slightly cohere, but in the majority of cases they lie free. No cases were found here or in later stages of the formation of a spirem. In many instances all the bodies within the nucleus retained merely the eosin stain (fig. 6), and hence were entirely undifferentiated from each other. In a few cases, like jigs. 5, 7, 8, some of the bodies retained only the black stain from the hematoxylin. In one instance (jig. 11), a faintly stained larger body, with one or two smaller ones of similar shade, can be seen lying within the nuclear space, sur- rounded by numerous more deeply stained granules. If the other stages mentioned had not been observed, the latter faintly stained body might have been interpreted as a nucleolus like those in higher plants, now in the act of becoming dissolved in the cytoplasm. Extended comparison, however, of parallel stages justifies the view that this body is only a portion of the central mass of the quiescent nucleus, about to undergo still further disintegration into chromo- somes. The many chromosomes thus resulting approach one another (jigs. 6, 9), presenting in many cases an appearance analogous to the synapsis stage described as occurring in the higher plants. Finally they become arranged in a circle concentric with the short axis of the cell. In one case (fig. 10), such an arrangement was observed before the nuclear membrane became dissolved. Fig. 14 shows this massing of granules in the equatorial plane after the dissolution of the nuclear membrane. The chromosomes in this cell were all stained black, but some were drawn in lighter tint to show that they were lying in three different planes. Fig. 15 also represents a similar stage, and fig. 18 one somewhat further advanced. The chromosomes now appear to be denser than in previous stages, an interpretation based upon the circumstance that the hematoxylin stain does not as readily become washed out. After having formed the ring they appear to be drawn inward, becoming denser and undergoing a process of fusion. By this draw- f ' 1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 47 ing-in process they come to lie in two closely adjoining parallel rows. As no case of a single row of isolated granules in the same plane was found, there is no evidence that such double row was produced by the division of a single row. Fig. 25 represents two rows of chromo- somes lying in the same plane. In fig. 12 fusion has taken place to such an extent that only three chromosomes are present in each row. Many of the chromosomes presented a tetrahedral appearance, - as in jigs. 16 and 20, thus pointing to the conclusion that the fusion of the condensing granules may take place in fours. In some cases the fusion has gone so far as to result in only four groups of tetrads (figs. 22, 23, 21). Fig. 24, of a more highly magnified group, shows especially distinctly this grouping of the granules. Careful focusing on this stage indicated the presence of another underlying group. As many cases were found of such grouping of the chromosomes in fours, it does not seem that it could have been purely accidental. When the maximum amount of fusion and condensation is reached, the limit apparently varying in different cells, each half of a group becomes dissociated from its adjoining members and gradually draws away, as in jig. 27. In process of separation each group becomes broken up into smaller groups, in the meantime all becoming again arranged in two rings concentric with the short axis of the cell (fig. 26). Thus numerous chromosomes are arranged in a circle in stages preceding and immediately following the stage of the equatorial plate, in which commonly four to six chromosome groups may be seen. It seems difficult to believe that six chromosomes (jig. 12) could have resulted directly from condensation and fusion of thirty or more chromosomes (fig. 14). A comparison of chromosomes as to size and staining qualities in the two adjoining cells (figs. 12, 13), drawn with Abbé camera, would certainly indicate that each chromosome must suffer a loss of its more liquid substance in the process of being drawn into the equatorial plate, or that a few must be entirely dis- solved. Whether all condense to form a few, or whether only a few are chosen to transmit the chromatin to daughter nuclei, the remain- der becoming dissolved in the cytoplasm, cannot be stated with certainty, as the staining process does not solve the problem as to the fate of the individual granules. When all the preparations are 48 BOTANICAL GAZETTE [JANUARY examined and not a selected few, there seems to be more evidence of the first being the true account of events. It was thought at first that this difference in number of chromosomes might be due to difference of species, as none of the Zygnema exam- ined had zygospores, and hence it is possible that two or more species might have been growing together. The discovery of cells like those _ in figs. 6, 12, 13, 15, 23, 22, in the same filament is indisputable evi- dence that in the same individuals the number of chromosomes de- creases from thirty or more down to six or eight, and then increases to thirty or more. This change in number occurs in a few moments, as determined in living cells by the changes in the position of the nucleus. All the filaments were examined in surface view, so it cannot be maintained that the number of chromosomes had been increased by sectioning. As the rings of chromosomes approach the chromatophores, the cytoplasm is condensed on the side nearest the chromatophore. The explanation of this might be that a large part of the cytoplasm which is not diverted to the region of the formation of the cell plate was streaming in toward the center, as in jigs. 14-18, while in figs. 20-30 it was streaming out towards the chromatophores; that the chromosomes are forced together by the inflowing streams and in the vortex of opposing currents become dissociated. The word “dissociated” is used in preference to the word “splitting,” as there appears to be no evidence of splitting and hence of equal distribution of homogeneous bodies. The chromosomes being heavier than the cytoplasm, the condensation appears on the side nearest the chro- matophore (figs. 28, 29). | It is to be regretted that in the living cells chromosomes could not be distinguished from actively vibrating granules in the cytoplasm. Nothing could be discovered which in any way resembled spindle fibers, although streams of granules and the alternating space of nuclear activity was easily traced. The number of chromosomes finally arriving at the chromatophore _ may be fifteen to twenty in each ring, as in fig. 30. The cytoplasm, being somewhat arrested in its flow by the chromatophore, causes a change in the position of the chromosomes. The majority, as they undergo still further dissociation, are drawn to the center, incom- 1906} MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 49 pletely cohering, while a few appear lying in delicate strands about them (jig. 31). In some cases all the chromosomes may cohere to form the central body. The nuclear membrane now emerges from the condensation of cytoplasm (fig. 32). As the chromosomes are now shut off from the influence of currents in the cytoplasm they generally remain unchanged in position, fusing either to form one mass (fig. 33), or three or more smaller masses (fig. 34), or rarely (jig. 35) all the chromatic material may be diffused in the nuclear plasm, forming numerous more or less tetrahedral granules. It is to be noted that not until the chromatic rings have separated and have approached the chromatophores do the pyrenoids ordinarily show any evidence of division. Tviis observation was easily con- firmed from the study of living cells. Fig. 29 represents the only one seen, out of many filaments examined, in which the pyrenoids divided before the formation of the nuclear membrane. As the newly divided nuclei approach their respective chromatophores, one or both plastids begins to show a constriction. This deepens until when the nuclei come to lie directly over, only a narrow band of less dense substance resembling linin connects the two daughter pyrenoids (figs. 37, 38). This becomes gradually reduced until it appears only as a thread (fig. 39). Later the nucleus sinks down and the separation is complete. The constriction of the plastids forming the center of the two pyrenoids takes place synchronously, as is the case with the stages in the daughter nuclei. One instance only was observed in which one plastid suffered division when other plastids had just begun to elongate (jig. 32). Although division of the pyrenoid may be influenced by division of the nucleus, that it is not wholly dependent upon it was demon- strated by leaving actively dividing filaments of Zygnema for one hour in a watch crystal containing 10°° of water to which two drops of chloroform were added. There were but few visible signs of plas- molysis in filaments killed and stained as in other material, but while a majority of the nuclei had ceased to divide, a majority of the pyre- noids were dividing as in normal filaments. That this division was not merely fragmentation was shown by sequence of stages and the presence of the band connecting the plastids. Fragmentation of the pyrenoids took place in filaments in stronger solutions of water Mo. Got.Garacn 1906 50 BOTANICAL GAZETTE [JANUARY and chloroform in which plasmolysis occurred to a much greater extent. Hence, cytoplasmic streams, nuclear structures, chromatophores, and pyrenoids take an active part in the division of cells in Zygnema. The streams of granules, collecting where the cell plate is to form, marks the beginning; the nuclear changes then proceed, followed by division in chromatophores and pyrenoids, while all are correlative with the formation of the cell plate. It cannot then be said that division of the nucleus, the chromato- phores, and the pyrenoids are synchronous. Rather is it true that the center of activities of the cell shifts, and with this shifting division of the bodies lying in the vicinity occurs. As regards the nuclear structures in Zygnema it is apparent that there are no bodies analo- gous to the nucleoli found in the higher plants. A large portion of the chromatin, or in a few cases possibly all, fuses in the anaphase to form one or more bodies corresponding in appearance and position to that of nucleoli of higher plants. Instead of waste products of chromatin condensing to form one or more bodies in the nucleus, the waste products are not separated from the chromosomes, but retained in them until after the nuclear membrane disappears in the next division. The substances which make up chromosomes and nuclear waste products, if such we may rightly regard the nucleoli of higher plants to be, are in Zygnema morphologically indistinguish- ble. The history of chromatin before the formation of the equatorial plate may be summarized as consisting of growth, association, and condensation of chromatin bodies in groups. These groups may be partially coherent, but in no case forma spireme. After equatorial plate formation, dissociation into groups follows, continuing until the chromosomes reach the chromatophores. Although the term chromosome has been used in this account, researches as yet incomplete make it exceedingly doubtful whether the chromatin bodies in any of the Conjugatae are to be rega as at all homologous with chromosomes of higher plants. If we restrict the term chromosomes to segments of the tubular spirem,* *See MERRIMAN, Vegetative cell division in Allium, Bot. GazETTE 37:178-207- pls. 11-13. March 1904. 1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 51 then the chromatin bodies seen in jigs. 14 and 15 of Zygnema cells are homologous not with the chromosomes of Allium but with the granules seen in the earliest stage of the spirem, while groups in figs. 16 to 23 are directly comparable with the groups or rings of tetrads, which in Allium fused to form the tubular chromosomes. Zygnema possesses a mechanism of nuclear division less elabo- rated than that of the higher plants, inasmuch as dissociation of chromatin bodies occurs immediately after their association into primary groups without the intervention of a spirem. From this point of view appearances observed in Zygnema support the inter- pretation suggested in my account of nuclear division in Allium, namely, that the chromosomes are formed by fusion of bodies in groups, and that when a longitudinal splitting appears it is not to be considered a true splitting of a homogeneous substance but rather a dissociation of bodies which from the first were discrete. If this be true, then doubts may reasonably be entertained as to the validity of the conception held by Roux, and successively by many other investigators, that the complex apparatus for indirect division of the nucleus exists for the purpose of enabling each chromatin body to furnish its quota to the daughter nuclei. The essential feature of indirect division, and therein its advan- tage over direct division, appears to be the dissolution of the nuclear membrane. Thus is made possible a free interchange of nuclear and cytoplasmic substances and a renewal of the vitality of the cell. Zygnema, then, may be considered as furnishing additional evi- dence of interchangeability of nucleoli and chromatin bodies, of variability in their number, and negatively as furnishing no evidence that equal distribution of chromatin is effected by either transverse or longitudinal splitting of homogeneous bodies. Nuclear structures, cytoplasm, pyrenoids, and chromatophores are trans- ferred in equal amounts to the daughter nuclei and by a process differing not fundamentally in the result from that which would _ have been attained by direct division. NORTHFIELD, Mass. - 52 : BOTANICAL GAZETTE [JANUARY EXPLANATION OF PLATES III AND IV. The figures were drawn with the aid of an Abbé camera. PLATE III. Fic. 1. Daughter nucleus from a cell -where the cell plate is not yet com- pleted. The nuclear structures in this cell retained the eosin stain, the pyrenoids black from haematoxylin. x 1750. Fic. 2a. Nucleus preparing to divide, showing growth of bodies in the periph- eral network before breaking up of the central body. Pyrenoids and nuclear structures in this cell retained only the eosin stain. 1750. uclear material stained black by the haematoxylin, all the chro- matic material being apparently condensed in the ee occupied by the central body. X 1750. . Fic. 3. Nucleus showing the breaking up of chromatin body and increase in size of the peripheral bodies. The pyrenoids retained the eosin stain; all the nuclear structures are stained black, several of them somewhat darker than the others. X 1750. IG. 4. Nucleus showing the beginning of the massing of the chromosomes, the nuclear membrane as yet undissolved, the granules in the region of the cell plate formation being conspicuous. Chromosomes black, pyrenoids red Fic. 5. Later stage, showing the clearing of the nuclear interior, recalling the synapsis stage described in higher plants. Pyrenoids red, several chromosomes black, remainder red. X 1 Fic. 6. Similar stage, very frequent; chromosomes numerous, massed together, all stained red. 1750. Fics. 7, 8. Similar stages where there is no massing of the chromosomes. In 7, a chromosomes were stained black, others red. In 8 those stained black are grouped in one corner of the nucleus, those red are scattered. 1750. Fic 9. A stage where distinct lines of granules connect chromosomes with nuclear membrane. Four chromosomes black, others red. X 1750. Fic. to. A rare stage with numerous chromosomes arranged in circle within the sitenk before the nuclear membrane becomes dissolved. All chromosomes lack. X 1750. _ Fic. 11. Another rare stage; nuclear membrane dissolved, remains of central body still in the cytoplasm, retaining a lighter eosin stain than the other chro- mosomes. X 1750. Fics. 12, 13. Two adjoining cells in same filament showing disparity in size and number of chromosomes. Pyrenoids red, in fig. 12 chromosomes stained sere by haematoxylin; in fig. 13 nuclear structures stained red. The line of granules marking the region of cell plate formation shown in both figures. x1 Fic. 14. All chromosomes black, but some drawn lighter to indicate that they were lying in three different planes. x 2440. Fic. 15. Chromosomes black, showing indefinite arrangement as hey are being drawn to the center. 2440. ta Fa ea aalicialtn ani Y ee oe 4 os poems . +s PLATE. Tif Fa ——e Bes MERRIMAN on ZYGNEMA BOTANICAL GAZETTE, XLI E.IV PLAT. BOTANICAL GAZETTE, XLI MERRIMAN on ZYGNEMA ip 1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 53 Fics. 16, 17, 19, 20, 21, 22, 23 show successive stages in condensation of chromosomes. Chromosomes all black. X 1750. Fics. 18, 24, 25, 26. Chromosomes and pyrenoids black. X 2440. Fics. 27, 28. Chromosomes becoming dissociated into smaller groups. Chromosomes black. x 1750. An ‘unusual case of division of-pyrenoids before formation of FIG. 29 Chromosomes black, both in central mass membranes of daughter nuclei. and in the periphery, pyrenoids red. X 1750. Fics. 30, 31, 32. Chromosomes black, pyrenoids red. 1750. Fics. 33. Pyrenoids and central body of nucleus black, peripheral bodies red. X1750. Fic. 34. Pyrenoids red, all the nuclear bodies black. < 1750. Fic. 35. Nuclear bodies red, pyrenoids black. 1750. Fics. 36, 37, 38. All nuclear bodies red, pyrenoids black. 1 . Nuclear bodies red, pyrenoids black, showing vestiges of connect- ing substance. X 1750. EFFECT OF CERTAIN SOLIDS UPON THE GROWTH OF SEEDLINGS IN WATER CULTURES.' J. F. BREAZEALE. (WITH FOUR FIGURES) In certain investigations on the growth of wheat seedlings in aqueous extracts of soil, it was observed that the growth of these plants was greatly accelerated by the presence in the medium of | undissolved calcium carbonate. That the observed acceleration was not due to an increase in dissolved calcium was apparent from the fact that the presence of other slightly soluble salts of this element failed to produce any response. It appeared possible that the effect of calcium carbonate might be due to its taking up some injurious substance present in the extract. This was suggested by NAGELI’s well-known discovery? that water, which is toxic to algae because of minute traces of metals, can be improved by placing in it such insoluble bodies as graphite, paraffin shavings, or torn filter paper. It was determined to try other slightly soluble compounds which might remove from solution small amounts of solutes, either by chemical action or mechanically. The results of this investigation make up the present paper. The Russian variety of wheat known as “Chul,” obtained from Arizona, was used in most of these experiments. The seedlings were germinated in sand and then grown in water cultures in large-mouthed black bottles of about 60°° capacity. They were fixed in cork stop- pers, four in a bottle, in the manner described by WHITNEY and CaMERON: for cultures of this kind, so that the roots were submerged in the solution while the seeds were just above its surface. The solutions were always aerated by violent and repeated shaking before the cultures were started. During the growth of the plants the bottles were weighed in groups of three at intervals of three or four days, and the water lost was replaced with distilled water. The manner I Published by permission of the Secretary of Agricu 2 Nageli, C. von, Ueber ip SPL E Sitar ees in lebenden Zellen. Denkschr. Schweiz - Naturforsch. segs 33:2: 3 Wuitney, M., and Cameron, F. K. geese in soil fertility. U. S. Dept Agric., fina of Seite Bull. sa sand Botanical Gazette, vol. 41] : [54 Tas 1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 55 of fixing the seedlings practically prevents water loss, except through the transpiration of the plants. The work of Lrvincston? indicates that total loss by transpiration for a period of two or three weeks furnishes a fair criterion for comparison of the growth of different cultures of wheat grown in this manner. The transpiration figures are used in this way in the experiments here given. The work was carried on in a greenhouse with a temperature of 15 to 25° C. For Experiments I to III a soil extract from poor Leonardtown loam, collected near Leonardtown, Md., was used. It was prepared by stirring the soil for three minutes with water in amount equal to twice its air dry weight, allowing it to stand twenty minutes, and then filtering through a clean Pasteur-Chamberland filter tube, in the man- ner described by WHITNEY and CAMERON.5 In Experiment I the solids used were calcium carbonate, tri- calcium phosphate, ferric hydrate and aluminum hydrate. Ferric hydrate was prepared by precipitation from the chlorid with ammonia, followed by thorough washing with hot water. It was transferred moist to the culture media. Aluminum hydrate was prepared in a similar way from the sulfate. Data for this and the three following experiments are given in Table J. The percentage increase in transpi- ration for each solid is computed by considering the transpiration from the untreated extract as unity. All of the solids accelerated growth, as is shown by the transpira- tion figures. But in the case of the ferric hydrate the root growth was accelerated to a much greater extent than that of the tops. The roots of the culture with this substance were much longer than those of the other cultures. It is evident here that root growth was acceler- ated without a corresponding increase in transpiraticn. Experiment II comprised, besides calcium carbonate and ferric hydrate, carbon black (prepared from burning petroleum, and thoroughly washed), magnesium carbonate, and barium carbonate. The small amounts of water transpired are due to cloudy weather. The plants of this series are shown in fig. 1, the numbers in the figure corresponding to the culture numbers given in parentheses 4 Lrvincston, B. E., Relation of transpiration to growthin wheat. Bort. GAZETTE 40:178~-195. 1905. s Witney, M., and Cameron, F. K., The chemistry of the soil as related to crop production. U.S. Dept. Agric., Bureau of Soils, Bull. 22:16 ff. 1903. 56 BOTANICAL GAZETTE [JANUARY in the table. It will be noticed that carbon black shows the same tendency to produce abnormal root growth as does ferric hydrate, but to a less marked degree. Experiment III included very finely pulverized quartz flour, as well as ferric hydrate and carbon black. The two last named bodies showed here the same abnormal acceleration of root growth as was previously observed, but quartz flour, although it improved the general growth of the plants, produced no such effect. iG. 1.—Experiment II; 24 wheat plants grown 19 days. 1, Extract of Leonard- town foals 2, the same with calcium carbonate; 3, the same with ferric hydrate; 4, the same with carbon black; 5, the same with magnesium carbonate; 6, the same with barium carbonate. The experiment with ferric hydrate and carbon black has been repeated many times with extract of Leonardtown loam, as well as that of other soils, and always with the same result. In some cases acceleration of root growth is more marked with carbon black than with ferric hydrate, but usually the reverse is true. Experiment IV was carried out with an aqueous extract, prepared as above, from Miami silt loam collected at the Rhode Island Experi- ment Staticn, at Kingston, R. I. This soil had been in hoed crops 1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 57 for ten years without fertilizer, and was acid to litmus paper. Te make absolutely sure that the effect of carbon black was not due to any substance added with it, the distilled water for the soil extract was shaken with the solid carbon black and filtered through a Fic. 2.—Experiment IV; 36 wheat plants grown 13 days. 1, extract of Miami silt loam; 2, the same with ferric hydrate; 3, the same with carbon black; 4, the Same with ferric hydrate, carbon black, and calcium carbonate. Pasteur-Chamberland tube before being used. Ferric hydrate, car- bon black, and a mixture of these two bodies, together with calcium carbonate, were used in this case. The plants are shown in jig. 2. From this experiment and others of similar nature it seems clear 58 BOTANICAL GAZETTE [JANUARY that the effect of the carbon black is not due to any stimulating sub- stance which it carries into the medium. Other experiments have shown that no acceleration of growth is obtained with the addition TABLE I DATA FOR EXPERIMENTS I TO Iv. ExPerIMENT I | Expermwent II | Experiment III) Experment IV 24 PLANTS 24 PLANTS PLANTS a GROWN I9 DAYS | GROWN 19 DAYS | GROWN 17 DAYS | GROWN 13 DAYS MEDIUM Total Total Per | Total Per Bit Per Total Per trans- cent. trans- cent. cent. trans- cent. piration| increase| piration| increase dav er z increase| piration| increase cultur’s) anresire Bon extract:.% |. 84:1. cc's ;. }{2) 33 tne 51 Preis, o.+tri-calc. phosphate..} 182 | 18.1 i tele ieee : + ates spe tans Decicaiiis carbonate..} 191 | 24.0 (2) 8 TAG Al hes corns eee Do.+ ferric hydrate. ..... 184 | 19.5 Bae 233.3 | 187 |266.6 |(2) 176} 158.8 Do. +-sitminwin Wydiate..6 2 487: eta ian be | ek Pe op eee ee ’ Do.+carbon black ...... ..ee |... | (4) 69/109.0 | 154 [201.9 | (3)139] 104-4 Do.+magnes. carbonate..| .... | ....-|(5) 73|121.2 ced i Baers ey ones er Do.+ barium carbonate...| .... | .... | (6) 78/136.3 aie wee Do.+ quartz flour........ pa ee ear es heel Sete 87 Fors Do.+ ferric hydrate, car- crt and calcium etic Ap a ors ae See Wee Py crag cre tk mr en deren es bP pee of iron salts to the extract of poor Leonardtown loam. Therefore, a slight increase in dissolved iron cannot be the cause of the accelera- tion noted in the case of this hydrate. It seemed probable that ferric hydrate and carbon black had their effect through an active removal, perhaps by mere adsorption, of some injurious substance occurring in the culture medium. Such a substance might have been in the soil extract originally, or might be produced by the plant roots, or both suppositions might be true together. The third alternative proved to be the correct one. To obtain evidence in this regard, Experiment V was carried out. Four different soil extracts were shaken with carbon black, filtered, and then used as culture media, comparison being made with controls in untreated extracts. The four soils-were of two types, a good and a poor soil of the Cecil clay type, and a good and a poor of the Leonard- town loam type, the former from Statesville, N. C., and the latter from Leonardtown, Md. Chemical analysis both of the aqueous extracts and of the solution obtained by digestion with hydrochloric 1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 59 acid, fails to show any material difference between the good and the poor varieties of these soils, although they are agriculturally quite different. These cultures were grown under the direction of Mr. F. D. GARDNER, in charge of the Division of Soil Management, of this Bureau. They consisted of forty-eight plants and were grown for 1, extract of Cecil .—Experiment V; 48 wheat plants grown 15 days. clay, ey 2, the same, filtered from carbon black; 3, extract of Cecil clay, poor; 4, the same, filtered from carbon blac fifteen days. The results are given in Table II; percentage increases are given for each treated extract compared with the same extract untreated considered as unity. It will be seen from this table that the extract of good Cecil clay 60 BOTANICAL GAZETTE [JANUARY EI. DATA FOR EXPERIMENT V. Culture no. Meatum ao) aos on ek er : aapedel st = mi sie POO cr. ne ote ewe ASG ian a |! —codnape ee Mise sachs wean pete Do. ROC oe tee ae 471 = 200 Bcad mops tae Extract 3 Cet clay, POOR sx ccsta vec P srs WOT scan Wy eed ieees AD cess ae ney th Sica] Oe UE DORN LEE OTOO 5. ooo. ssa Jonnie oh aces Sesh maee 477 “ize? [Ser eS eee Extract of Leonardtown loam, good.. a! ee eee ee irae, ogee Ceara SN TRMOTEE S Scw re emo ope eee 407 +13.4 ype eres Ea ract of ie te loam, poor.. DRL ea «tte average Buea sae Do carbon filtered si. i465 sien ees 349 + 28.7 was not improved by treatment with carbon where the latter is filtered out. All the others were im- proved, although the improve- ment in the case of good Leon- ardtown loam was not as marked as that in case of the poor. The leaf development and _ general appearance of the plants were essentially in proportion to their transpiration. The roots showed __ the same acceleration with carbon ~ treatment and filtering that had been observed in experiments in which the solid was left in the solution. Fig. 3 shows the roots of cultures nos. 1, 2, 3 and 4. es part at least ‘of the injurious matter which is removed by car- bon black is in the soil extract at the start. : A number of experiments were : glee pede igi ee grown carried out, using water twice Ae ee be ed water; 2,thesame, redistilled, first from potassium dichromate and sulfuric acid, then from alkaline eens permanganate, both in glass boilers, con- 1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 61 densation being carried on in a platinum tube. Upon this water the effects of carbon black and ferric hydrate were tested, the solids remaining in the water during the growth of the plants. The former gives little or no increase in transpiration, the latter a moderate increase, but both solids produce’a marked acceleration in root growth. Twelve seedlings grown fifteen days in redistilled water with and without ferric hydrate are shown in fig..4. No. 1 shows those from the untreated water, no. 2 those with the solid. The water here used probably did not contain injurious substances, and therefore the effect of the solids is most probably due to the removal of some injurious exudation arising from the plants. Further, distilled water in which plants have been grown for a number of days is found to give less growth upon replanting than does unused water, and the injurious effect of the used water is corrected by shaking with carbon black or ferric hydrate and filtering off the solid. Thus it seems that wheat seedlings do give off bodies from their roots which are toxic to them- selves. When the work so far recorded was practically completed, the appearance of TRUE and OGLEVEE’s paper® on somewhat similar experiments made it seem advisable to withhold publication until some further tests suggested by that paper could be made. These authors find that by the introduction of sand, filter paper, parafiin, or potato starch into solutions cf copper sulfate in which seedlings of Lupinus albus are growing, the toxicity of the solute is remarkably decreased. By this means the killing concentration of the salt may _ be effectively reduced, according to the amount of the insoluble body present, either to a stimulating concentration or to one in which the physiological effect is not apparent. They. reasoned that the solid absorbed the salt from the solution and in this way produced an effect closely paralleling that of simple dilution. It was determined to test the effect of solids in solutions of sulfuric acid. Maize was here used instead of wheat. First, the strength of this acid necessary to prevent the growth of maize seedlings was 6 True, R. H. and Octever, C. C., The effect of the presence of insoluble sub- stances on the toxic action of poisons. Science N. S. 19:421-424... 1904. Bot. GAZETTE 39:1-21. 1905. In this connection see also DANDENO, J. B., The relation of mass action and physical affinity to toxicity, etc. Am. Jour. Sci. 17: 437- 458. 1904. 62 BOTANICAL GAZETTE [JANUARY determined as lying between 2/2750 and /3250.7. Then a series of varying concentrations of this acid on either side of the toxic limit was carried out, placing clear sand, quartz flour, filter paper, and paraffin shavings in the solutions. In no case was the apparent death limit modified by the presence of these substances. The death limit was also determined for a solution of sulfuric acid saturated with calcium sulfate, and for the same solution with an excess of calcium sulfate, but the solid again had no apparent effect. With copper sulfate solutions carbon black was found to decrease the toxic effect, just as the authors above cited found to be true for the solids with which they worked. In order to bring out the effect of the carbon black, copper sulfate in the proportions of one and five parts per million of copper was added to the nutrient solution above described; a portion of the solution thus prepared was shaken with carbon black and then filtered, and wheat seedlings were grown in the treated and untreated solution. This series constitutes Experi- ment VI. Twenty-four plants composed a culture and the experiment lasted twelve days. The results are given in Table III. ‘TABLE: Tf. DATA FOR EXPERIMENT VI. Culture no. Medium — eff eign 2 aoe Be vp te wee Nutrient solution........... 220-2 PEGs Lees Do.+1 p. an vas Tenaya ee 67. Lege seen Decks pms Cu. Si... 41 per emner sey eater As 2, a pdt Siig TE 140.2 sl i a ce toate As 3, but carbon-treated. . T7875 The growth of the plants was proportional to their transpiration. It is evident that the carbon removed sufficient copper to render carbon-treated solutions much less toxic than the untreated ones. This is a direct corroboration of the work of TRUE and OGLEVEE, but with another solid, and in this experiment the filtering out of the solid removes any possibility of its having any effect directly upon the roots. The explanation of these authors seems to be correct, as far as copper sulfate is concerned. The failure in the present 7 This or aeromine had been made previously, but was repeated for the present work. See Cameron, F. K., and BRreazEAate, J. F., Toxic action of acids and salts on seedlings. wee aie aia: 8:1-13. 1904. i 1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 63 instance to get the same effect with sulfuric acid may be due to the fact that to be injurious this substance must be at a much higher con- centration than is needed in case of the copper salt, while the relative amount of solute absorbed by solids is much greater in dilute solutions than in more concentrated: ones. TRruE and OGLEVEE suggest this explanation for their failure to get marked improvement by the use of sand in solutions of phenol and resorcinol. On account of the presence of toxic substances in distilled water as ordinarily prepared, from copper boiler and tin condenser, most workers with toxicity problems have used water redistilled in glass. In these laboratories the distilled water is often quite toxic to wheat seedlings, but its injurious effect is prevented if it is first shaken with carbon black or ferric hydrate and the solid filtered off. It is found that water so treated produces as good a growth of seedlings as does the most carefully prepared redistilled water. It appears from these experiments (1) that extracts of certain soils are toxic to wheat seedlings in water culture, and that this toxicity is removed wholly or in part by carbon black, calcium carbonate, ferric hydrate, and other solids; (2) that the toxic substances of ordinary distilled water may be removed by ferric hydrate or carbon black; (3) that the roots of wheat seedlings give off substances which are toxic to themselves and that these substances can be made inactive by the presence of the last named solids in the culture medium; (4) that the presence of ferric hydrate and carbon black in the solution seemingly accelerates to a marked degree the development of roots, causing them to surpass the tops in growth. The work here reported was done chiefly in the laboratories of the U. S. Department of Agriculture, Bureau of Soils, Washington, D.C. It was finished at the Rhode Island Agricultural Experiment Station, Kingston, R. I. I am indebted to Dr. F. K. CAMERON, and to Dr. B. E. Lrvrncston, of the Bureau of Soils, for much valuable suggestion and advice. U. S. DEPARTMENT OF AGRICULTURE, Bureau of Soils, Washington, D. C. PRrehe ek AR LICLES. NOTES ON NORTH AMERICAN GRASSES. V. SOME TRINIUS PANICUM TYPES. BELOw are given a few notes upon certain species of Panicum described by Trinius from American material. The herbarium of Trrntus, the father of modern agrostology, is deposited in the St. Petersburg Academy of Sciences. The curator of the herbarium, Dr. D. I. Lrrwrnow, very kindly sent, for deposit in the National Herbarium, portions of the speci- mens of the species mentioned in this article. It will be observed that three of the seven species had been described by earlier authors, while four have been recently described as new species. It must be said, however, that it would be impossible to identify any of these four species from the description alone. This is true of many of the earlier descriptions of the group Dichotoma of the genus Panicum, and on account of the difficulty of consulting the types of these species, it is in a measure excusable, on the ground of convenience, to describe unidentified species as new, without determining the application of the older names. Such painstaking and laborious comparison should not, however, be shirked by a monographer. P. CHAMAELONCHE Trin. 1826, Gram. Pan. 242. are et minus. Panicula (semipollicari) lucidissima; Spiculis pusillis, obovato-oblongis, obtusiusculis, glabris; pedicellis hispidulis; Gluma inferiore flosculis triplo breviori 1-, superiore eosdem aequante 5-nervi; Hermaphrodito elliptico, mucronulato, laevi, neutrum aequante. V. spp. Am. bor. (TRATTINICK, ex. coll. Enslini). Plantula caespitosa, stricta, culmo tenuissimo, ramuloso. Folia lanceolata 1. lineari-lanceolata, strictiuscula, semipollicaria, glabra; culmea et juniora subinvoluta, acicularia, breviora. Panicula pauciflora, tenera, prodit e summa vagina elongata, cujus folium eandem fere aequat. Flosculus neuter univalvis ? Hermaphroditus albescens. Label accompanying type specimen: ‘‘In Am. bor. s. dat. sine nom. ex. hb’° Enslini cl. Trattinick.” Specimen =P. Baldwinii Nutt. 1898. Scribn. Div. Agros. Bull. 11:43- P. pemissuM Trin. Spec. Gram. 3:319. oe The type specimen is from Rio Janeiro. I have not seen this species from North America. It is mentioned here because the name occurs occasionally in the literature of North American grasses, and has been doubtfully applied to certain of our species. Botanical Gazette, vol 41] [64 1906} BRIEFER ARTICLES 65 P. Ensiint Trin. 1826, Gram. Pan. 2 Pedale. Panicula (subdigitali) ude lucida; Spiculis subultra- linealibus, ellipticis, acutiusculis, pilosulis: pedicellis glabris; Gluma inferiore flosculis sub 4-plo breviori 1-, superiore eosdem vix ae 7-nervi; Her- maphrodito oblongo, acutiusculo, laevi, neutrum fere aequa An Panicum tenue Miihlenb. (quod Pan. liton ne Mant 2. 'p. 250) quaerit ill. N. AB Esenp. in litt. V. spp. Am. bor. (TRATTINICK, e collect. Enslini.) Culmus basi ramosus, ad paniculam usque vaginatus: vaginis arctis, fissura pubescentibus. Folia glabra: radicalia lanceolata-oblonga 1. ovata, sesqui- pollicaria, lineas 4-5 lata; superiora lineari-lanceolata, duplolongiora, patentia. Panicula e summa vagina prodit radiis subcapillaribus, parum patulis. Gluma inferior epilis, superior mucronulata cum flosculi neutrius valvula inferiore pilis adspersa. Hermaphroditus albescens. Label accompanying type specimen: “(an Pan, tenue Muhlb. quaerit NEEs AB Es.) ab Enslino in Am. bor. ]. dt. sine nom. cf. Trattinick Wiennae 1820.” Specimen =P. equilaterale Scribn. 1898, Div. Agros. Bull. 11:42. Characterized by having the spikelets of P. commutatum but the leaves elongated and widely spreading. P. FLORIDANUM Trin. 1835, Mem. Acad. Petersb. VI. 37: 248. The type specimen from Georgia is Paspalum racemosum Nutt.=P. bifidum (A. Bertol.) Nash, as has been generally recognized. P. JEJUNUM Trin. 1836. Bull. Acad. Petersb. 1:76. The type specimen from Louisiana sent by Hooker in 1835 is Panicum hians Ell. =Steinchisma hians (Ell.) Nash. P. LANCEARIUM Trin. 1822, Clavis Agros. 234. 1179. Gr. miliaceum americanum, minus, panicula parva. Pluk, Phyt. p. 176. Tab. 92 f. 6. Mor. p. 197. no. 15. Panicum lancearium m. (de quo alio loco). Label accompanying specimen: ‘“Plukn. Tb. 92 {. 6.2? In Am. bor. ab Enslino I. dt. cl. Trattinick.” Specimen=P. Nashianum Scribn. 1897, Div. Agros. Bull. 7:79. The specimen matches Curtiss 4029 from Florida, the first specimen cited in the original description of P. Nashianum. Both have glabrous spikelets. The second specimen cited by Scripner, Nash 466, from Florida (the type on account of the specific name), has pubescent spikelets. Since Trinius gives a binomial to a plant described by PLUKENET and by Morrison under a polynomial designation, PLUKENET’s plant is the type. In Kew Index P. ancearium is cited as Agrost. Bras. 246. The name 66 BOTANICAL GAZETTE [JANUARY is mentioned in that work (NEES, 1829, Agrost. Bras. 226) in a note appended to P. parvifolium Lam., where it is referred to P. angustijolium Ell. It is quite distinct from ELLiort’s species, however. Trintus himself describes the species | later as follows (1826, Gram. Pan. 22 Spit beaes maeum. | Panicula (vix pollicari) lucidissima; Spiculis parvis, obovatis, glabris: pedicellis scabriusculis; Gluma inferiore foarailia triplo breviori enervi, superiore eosdem aequante 7-nervi; Hermaphrodito oblongo, acutiusculo, laevi,*neutrum aequante. V. spp. Am. bor. (TRATTINICK ex hbi°. Enslini). Ima basi in ramos discedit simplices, tenues attamen firmos, satis multi- (sex-)folios. Folia lanceolata, pollicaria, sensim breviora, praesertim basin versus ciliatula. Panicula e summa vagin ae satis pauciflora, lineari- oblonga. Flosculus neuter bivalis. Hermaphro Since PLUKENET’s figures of this and the next cannot be identified, TRINIUS’ spectmens should be taken as the substitute type of these two species. P, LEUCOBLEPHARIS Trin. 1822. Clavis Agrost. 234. 1177. Gr. miliaceum americanum, majus, panicula minore. Pluk. Phytogr. p. 176. Tab. 92. f. 7. Mant. p. 95. (excl. Syn. Sloan. ut. ipse Sloaneus monet). Citatur a Gronowio (Virg. p. 12.) ad Pan. paniculatum floribus muticis; sed quid illud? Figura bene convenit cum Panico quodam herb. notsr. ex Amer. bor. (Pan. leucoblepharis m.) praeter cilia foliorum elegantissima, rigidiuscula.— Synon. Recchit ap. Pluk. admodum dubium Label ereenenyte specimens: ‘“‘ab Enslino in Am. bor. 1. dt. cl. Trattinick.” Specimen =P. ciliatum Ell. 1816. Sketch 1:126. Like P. lancearium the name is founded upon a figure in ‘’PLUKENET and is further described by Trintus in Gram. Pan. 219. 1826. Spithamaeum et minus. Panicula (ultrapollicari) lucida; Spiculis sub- rvis, obovato-oblongis, pilosulis: pedicellis scabriusculis; Gluma_ inferiore flosculis plus duplo breviore 3-5, superiore eosdem aequante 7-nervi; Her- maphrodito oblongo, obtusiusculo, scrobiculato-punctato, neutrum aequante. V er. bor. (TRATTINICK e plantis Enslini). asi ramosum. Folia cordato-lanceolata, amplexicaulia, pl. min. pollicaria, pallide viridia (plerumque elegantissime), pectinato-ciliata. Vaginae superiores elongatae. Axis pilosus. Gluma inferior epilis. Flosculus neuter bivalvis. P. ciliatum is characterized by the ciliate but otherwise glabrous leaves and pubescent spikelets a little less than 2 ™™ long P. UNCIPHYLLUM Trin. 1826, Gram. Pan. 242. : Spithamaeum-pedale. Panicula (1-sesqui-pollicari) lucidula; Spiculis mini- 1906] BRIEFER ARTICLES 67 mis, oblongis, pilosis: pedicellis glabriusculis; Gluma inferiore flosculis triplo breviori 1-, superiore eosdem aequante 7-nervi; Hermaphrodito elliptico, laevi neutrum aequante. Panicum laxiflorum Spreng. in Mém. de St. Pétersb. II. p. 291. Panicum heterophyllum Miihlenb. teste N. ab Es. V. spp. Am. bor. (TRATTINICK). Culmus tenuis, adscendens, basi ramosus. Folia, quorum plura basi plerum- lata: superiora angustiora, dissita. Panicula ovata, axis radiisque glabris. Flosculus neuter bivalvis. Hermaphroditus albescens. Label accompanying type specimen: “Pan. heterophyllum Muhl. (Test. Nees) an Pluckn. Tab. 92 f. 8? ex herb Enslini, spmna Am. bor. Trattinick.” Specimen =P. columbianum Scribn. 1897. Div. Agros. Bull. 7:78. In recent works this name has been applied to a species of the Janu- gimosum group having rather stiff foliage and the leaf blades hirsute on _ both surfaces. The true P. unciphyllum is easily recognized by the short crisp pubescence and the very short ligule, characters not mentioned in the original description.—A. S. Hirccock, U. S. Dept. Agric., Wash- ington, D. C SPOROGENESIS IN PALLAVICINIA. Tue August number of the BoTANicaAL GAZETTE contains a paper by Mr. A. C. Moore on Sporogenesis in Pallavicinia. 1 regret again to ask for space on this matter, but Mr. Moore has so completely (though of course inadvertently) misrepresented my own position with regard to- the nature and the significance of the quadripolar spindle in the Junger- mannieae, and further, the grounds on which he founds his criticism appear to me to be so open to objection, that I venture to ask for an oppor- tunity of replying to his strictures. Firstly, then, as to the significance attached to the quadripolar spindle in 1894-5. From Mr. Moore’s account it would seem that I regarded, as the Most essential feature of its importance, the simultaneous distribution of the chromosomes of the dividing nucleus of the mother-cell to the four spores that are finally produced. I certainly believed that in Pallavicinia decipiens such a distribution occurred, and that it resulted from the suppression of the period of rest normally intervening between the first and second maiotic divisions. In this I may be right, or further investigations may show that, in the species In question, I missed the binucleate stage, But this is really not the 68 BOTANICAL GAZETTE [JANUARY essential matter at all. The result of my work published in 1895 went to show that in most forms there are two consecutive mitoses, the second, following more or less rapidly on the first, and I believed that in P. decipiens the brief interval might be so shortened as to have become practically obliterated. But the circumstance that quadripolar spindles were shown by me to be plainly visible in properly fixed material of forms in which no such extreme telescoping of the normal sequence of events takes place, clearly proves that, whatever the significance of the quadripolar spindle may be, it certainly is not essentially related to a simultaneous distribution of the chromosomes amongst four daughter nuclei, and I never thought it was. What I believed in 1895 (and I have seen no reason to materially alter my view), was expressed as follows: ‘“‘The quadripolar spindle, then, is only a special case of ordinary karyokinetic phenomena; instead of two relatively large masses of protoplasm there are four distinct aggre- gations, one in every lobe, each exercising an independent strain, and the direction of the strains may continue separate to the very end of the process or not, according to the form and special circumstances of the cell.”* I may perhaps add, that the principal importance of the phe- nomenon, in my view, lay in its bearing on the permanence of the centrosomes, at that time a widely accepted doctrine. In the second place, Mr. Moore seems to think that his observations on P. Lyellii vitiate the conclusions based on a study of P. decipiens. I venture to think they do nothing of the sort. It is clear that the two species differ in the form of their spore mother-cells to a marked degree, and also that this difference is exactly of a nature to account for the unequal persistence of the peculiarities of the spindle in the two cases. For the lobing of the spore mother-cell is so much less in P. Lyellii than in the other species, that it would be a matter for surprise if the quadripolar character of its spindle were so long retained. I confess, however, that I should have expected centrospheres to be present at the stages represented in pl. III, figs. 1-3 of Mr. Moore’s paper. They are so obviously demonstrable in Ameura pinguis and in Fossombronia pusilla, the spore mother-cells of which resemble in their lobing those of Mr. Moore’s plant. One feels a little difficulty in repressing a suspicion as to the successful fixation of his material, a suspicion not dispelled by the further contem- plation of figs. r2 and 13. They so faithfully depict preparations I have t Annals of Botany 9: 508. . 7 1906] BRIEFER ARTICLES 69 myself very often obtained when the fixation had been imperfect. It is, of course, easy in these plants to secure admirable preparations of the stages preceding and following on the maiotic divisions, but I am sure Mr. Moore will agree with me as to the great difficulty encountered in successfully fixing the cell contents at this critical period. Personally, I have not found chromacetic acid (the fixative used by him) very suc- cessful, but obtained far better results with Flemming’s solution and, if due precautions are taken, with acetic alcohol. The latter, in par- ticular, has yielded results of especial excellence, owing partly, no doubt, to the relative rapidity with which it traverses the somewhat impervious cell wall—J. B. Farmer, Royal College oj Science, London. REPLY. PROFESSOR FARMER acknowledges that in 1894 he believed in the simultaneous distribution of the chromosomes to the four spores in Pal- lavicinia decipiens. His description stands as the only account of a pro- cess without parallel in the plant kingdom, and he must have realized its exceptional nature. The account became all the more remarkable when Professor FARMER’S own studies on a number of liverworts, published in the following year, showed two successive mitoses in the spore mother- cells as in other groups of plants. He acknowledges now that he may have missed the binucleate stage. This is precisely what I believe he did, but since I have not investigated P. decipiens I cannot assert that he did so. Now he states that this simultaneous distribution is really not the _ essential matter at all. Apparently the essential matter to him is his observation that several liverworts conform to the normal sequence of nuclear division during sporogenesis. Yet these conclusions, bearing as they do on Pallavicinia decipiens, served to emphasize the peculiarities of that account, and I feel confident that most, if not all, cytologists would pick out the description of a simultaneous distribution of chromosomes as - the most essential feature of his paper of 1894. I venture to think that botanists are not so much interested in the explanations which Professor FARMER may make of what he did or did not believe in 1894 and 1895 relative to the quadripolar spindle (which opinions they can form for themselves), as in the facts of sporogenesis in the liverworts. My study of Pallavicinia Lyellii is plainly a challenge of his account of P. decipiens, and together with Professor Davis’s work on Pellia, leads us to believe that the ‘‘quadripolar spindle” in all liverworts is a phenomenon of prophase followed by spindles of two successive mitoses, in essential agreement with the events of sporogenesis in other plants. vic) : BOTANICAL GAZETTE [JANUARY The reader must judge for himself whether it is at all likely that two species in the same genus should differ from one another so fundament- ally as would appear from Professor FARMER’s description of sporogenesis in Pallavicinia decipiens and my own account of P. Lyellii. Respecting the fixation of my material, I may say that I have no reason to think the penetration was not sufficiently rapid to fix the cell contents. Even with imperfectly fixed material my main conclusion is easily demon- strable, viz., that in P. Lyellii there are two successive mitoses in the spore mother-cell. Let us not lose sight of the main point at issue— ANDREW C. Moore, South Carolina College, Columbia. CURREN T-LELERA TURE. BOOK REVIEWS. The algal vegetation of the Faeréese coasts. BORGESEN’s extremely interesting account of the algal associations on the coasts of the Faerée Islands,‘ is one of the most important contributions to the cS eran ll «Sey ecological side of marine botany. The work is a description of conspicuous algal associations along a varied rocky coast line, particularly favorable to algal vege- tation, and is illustrated by more than thirty very excellent plates and figures from photographs. The factors affecting the algal vegetation are discussed; such as temperature and salinity of the water, tides and currents, wave action, temperature and humidity of the air, and light. The littoral and sublittoral floras are described, both for exposed and sheltered coasts, and also the floras of tide-pools and caves. A great many algal associations and formations may be clearly recognized in the Faerdes, some of them very conspicuous, as the Chloro- phyceae formation, the Porphyra association, Fucaceae formations, Laminariaceae formation, and Alaria association. A particularly interesting chart plots the position of these associations in their position above and below the mean sea level. It is extremely interesting to note that the cave flora is composed of forms of the sublittoral flora, which in the dim light are able to grow near the surface, or they are types which have the habit of growing in shaded situations outside. Littoral forms which grow in the brightest light are only found near the entrance of the caves. On entering a cave a condensed picture is obtained of the vertical : distribution of algae from above downward. The forms in the deepest shadows f are all red algae and some of them species which are usually found at great depths in the open sea. It is clear that light is the most important factor affecting the position of algal associations along a coast. There is a detailed comparison of the algal flora of the Faerée Islands with neighboring countries, Scotland, the Orkney and Shetland Islands, Norway, and Iceland, preliminary to a discussion of its origin. The flora had its origin from a mixture of Atlantic and Arctic species, which wandered northward with the retreat of the ice. Some of the arctic forms remained, adjusting themselves to the warmer waters, but there are many peculiarities of the algal flora which demand special explanations. BORGESEN does not believe that there were post- glacial bridges of land which made possible the migration of forms, but holds that factors now operative might have brought to the islands many algae from neighboring countries. * BORGESEN, F., The pee peta of Faerdese coasts. Im 834. pls. 13-24. Meads en: . Thiele. 1905. [Reprinted ible ‘Rtens a the Faerdes. See Bor. GAZETTE ty 392. 1903-] 1906] | 71 ye BOTANICAL GAZETTE [JANUARY Sea currents are regarded as of greatest importance. The pronounced cur- rents from the nearest land do not bathe the islands, but experiments have proved that heavy winds and storms will drive floating objects out of the main currents, and BORGESEN believes the general conditions to be favorable to the introduction of algae from the west and north coasts of Ireland, the west coast of Scotland, and the Hebrides, while the currents from east Iceland run straight to the Faerées. It is also possible that algae may be introduced from the west coast of Norway. - Fragments of the algae may drift for many days, especially such as are provided with bladder-like floats, or their spores may be so carried, and floating pieces of timber covered with algal growths are known to travel long distances. Smaller algae of the littoral flora are very likely to be introduced with mud upon the feet and bodies of birds. Finally BORGESEN believes that algae may be introduced through the shipping which visits the islands. These are merely some of the most striking conclusions in an account that is full of interesting observations on the life conditions and habits of marine algae.—B. M. Plant diseases. FREEMAN has produced a finely illustrated volume on plant diseases,? the first part of which is devoted to a discussion of fungi in general, while the second special part treats of specific fungous diseases of plants. The object of this book, as set forth in the preface, is “rather educational than immediately practical.” It is an attempt to give a general account of the nature of fungi, for the purpose the work becomes rather broader than would be indicated by the title, Minnesota plant diseases. t part comprises a discussion of the morphology, physiology, and pee of fungi; but, while this part contains much excellent material, the arrangement lacks the logical sequence of first importance in a book of an edu- cational character. It consists rather of a series of interesting pictures without due regard to pedagogical principles. This is likely to leave the mind of the reader confused. The sub-headings of the first chapter on nutrition are as fol- lows: What the fungi are; The fungus method of obtaining nutrition; How the nutritive method is expressed in structure; Parasitism and saprophytism; Storage organs; Fungus shoestrings or strands; Physiology of the mycelium. Then, in chapter III, Fungus life methods, we have as the first subhead, again, Parasitism and saprophytism, the rest of the chapter dealing with habits or rather habitats of different aati 00 great an effort is made to avoid scientific pass Thus we have such “‘spore-like swimming-spore-cases,”” “‘Sac-spore-capsule.” It would seem that the reader who can comprehend the allusions to the phylogenetic relationships between fungi and algae would not find it too difficult to comprehend a few scientific terms. ? FREEMAN, E. M., Minnesota plant diseases. Imp. 8vo. pp. xxiii+ 432- figs, 211. St. Paul: Rape of the Survey. Bot. Ser. V. 1905. 2 ey ciel apn ert age mar ani muratety 2m Cero eo eT a 1906] CURRENT LITERATURE 73 The second part of the book is devoted to descriptions of special diseases. These are classified according to the nature of the crops on which they occur, as follows: Timber and shade trees; Field and forage crops; Garden net Orchards and vineyards; Greenhouse and ornamental plants; Wild pla Under those heads the groups of fungi, as rusts, smuts, mildews, etc., are together —H. HaAssELBRING. Regeneration. WITH THE TITLE Studies in regeneration NEMEC? has published in rather voluminous form the results of his investigations on the regeneration of root-tips. The general conclusions may be briefly summarized as follows. Cutting a trans- verse section just at the tip results in the regeneration of a new tip in a radial manner. The dermatogen-and outer part of the periblem takes no part in this, the new tissue arising from the inner part of the cortex and the plerome. There is first of all the formation of a callus of hypertrophied cells, between which and the meristem arises the group of initials by which the new root-tip is organized. This group is radial from the beginning, the majority of its cells arising from the plerome, only the peripheral ones coming from the periblem. Proceeding back from the tip, the se tesane for regeneration diminishes from the periphery inwards, soon disappearing from the periblem. As long as the central cells of the plerome still possess this capacity the regeneration is radial. Farther back it is confined arises. When the capacity of the inner cells of the periblem and the outer cells of the plerome to take part in regeneration is lost, the replacement of the removed root-tip occurs only through the origin of lateral roots, which arise in the peri- cambium. When the root is cut through obliquely, the regeneration of the new root-tip occurs at the part of the cut surface nearest the tip. When the tip is slit lengthwise each half re-forms a new tip. If a tip is slit lengthwise for about 1™™, and then one of the halves is removed by a transverse incision, the remaining half regene- rates a new half, and also, at the surface formed by the transverse cut, a new tip is developed. Lateral incisions to produce new roots must go at least half way through the plerome. Unless such an incision is made just back of the tip a new tip is soon organized immediately above the cut. The original tip is pushed to one side and finally is displaced entirely. When the incisions are made on two opposite sides of the root at different levels, new root primordia arise at each place, but only the one nearest the original tip continues to develop. If two incisions are made, on opposite sides and at the same level, a new root arises at each, but one is soon suppressed, while the other develops and finally replaces the original tip. About forty-eight hours after the wounding, starch usually appears in the cells of the periblem just above the cut. The grains are not yet 3 NEmec, B., Studien tiber die Regeneration. Imp. 8vo. pp. 387. figs. 180. Berlin: Gebriider Borntraeger. 1905. M 9.50. 2 74 BOTANICAL GAZETTE [JANUARY mobile, and are aggregated about the nucleus. In about twenty-four hours more, however, they become statoliths and fall to the bottom of the cells. During this time the original tip has been losing its starch, and there is a period of from forty- eight to seventy-two hours in which the old tip has lost its starch and the new tip has none in a movable form. During this period the roots are ageotropic. In ferns the root-tips do not regenerate. Tips cut off transversely just back of the apical cell are unable to organize a new one, though they may continue growing for several weeks. As the statolithic starch is in the root-cap, and this does not regenerate, such roots remain ageotropic. Besides the discussion of the experiments, a number of chapters are devoted to a discussion of such topics as the influence of external conditions on regener- ation, polarity and regeneration, growth and regeneration, purposefulness of regeneration, relation between geotropism and the presence of statocytes, and other interesting topics connected with regeneration. As the root-tip regenerates from so many kinds of injuries that could never occur in nature Némec considers that at least in the great majority of cases the capacity could not have arisen because of its utility. The immediate stimulus, he thinks, does not lie among nutritive changes, or arise from the wound, but is a phenomenon of correlation, due to the breaking of the connection between the vegetative tip and the root meristem.—W. B. McCattium. Plant histology. CHAMBERLAIN has revised and rewritten much of his Methods in plant histology,+ adding several new chapters, elaborating and in many instances shortening the processes. Several new formulae are given for killing and fixing. e paraffin method has been notably improved and the celloidin method has been treated at greater length. A method for embedding in soap is also given. The new chapters deal with microchemical tests, free-hand sectioning, special methods, the use of the microscope, and micrometric methods involving the use of the camera lucida. A very important new chapter deals with methods of staining filamentous algae and fungi and mounting them in Venetian turpentine. An abstract of the methods of PFEIFFER and WELLHEIM is given, together with such modifications as have been found to give successful preparations. Delicate forms like Vaucheria can be carried through the stains and finally mounted in Venetian turpentine without showing the least trace of plasmolysis, and even if slight plasmolysis should occur it can be corrected by manipulation of the mount- ing medium. Preparations made by this method are exceedingly brilliant and show a wealth of detail not possible with other methods. For example, the two nuclei in zygospores of Spirogyra can be readily seen with a low magnification. The Venetian turpentine method, which gives preparations requiring no sealing and as hard and durable as balsam mounts, should almost entirely replace the glycerin method. 4 CHAMBERLAIN, CHARLES J., Methods in plant histology. pp. x+262. 7gs- 88. Chicago: The University of Chicago Press. 1905. Net $2.25; postpaid, $2-39- 1906] CURRENT LITERATURE 75 Much attention is given to collecting and keeping material alive in the lab- oratory. K1EBs’s method of securing reproductive phases in algae and fungi is presented in a practical manner. Specific directions are given for making such preparations as are needed by teachers and by those who wish to get a compre- hensive view of the plant kingdom from the lowest to the highest forms. The book will be useful to those who wish to keep in touch with modern microtech- nique.—W. J. G. LAND. Bibliographical index of North American fungi. THE compilation of a bibliographical index of North American fungi by FARLOWS is one of the most serviceable tasks ever undertaken in the interests of American systematic mycology, and the publication of it by the Carnegie Institu- tion one of its best contributions to the promotion of botany. The work is the ' outgrowth of an effort to bring together references to all North American species in the form of a card catalogue. This was begun in 1874, at a time when there was no complete record of the species known from North America. Within a few years of its inception Mr. A. B. SeyMour was entrusted with the details of this herculean labor, under Dr. Farlow’s direction, and his painstaking fidelity is worthy of recognition. It is the aim of the work to include all references having any bearing on the taxonomy of fungi occurring in countries north of the Isthmus of Panama, the scope of the original plan (which was restricted to the region north of Mexico) having been greatly extended, on account of the close connection of species from our southern border with those of Mexico, Central America, and the East Indies. References to works of purely morphological, cytological, and sui interest have been excluded; so have purely popular accounts, unless Ww of use in giving divination of the species or in furnishing good inns. In nomenclature the work is conservative. The principle of adopting the oldest specific name has been generally followed. Where the vagueness of older descrip- tions has made it uncertain to what stn they applied the writers have had no scruples in rejecting the older n The index itself is arranged ‘aiphaetically The names are printed in bold- face type, synonyms and cross references being in italics. The citations, arranged in chronological order under each name, follow the form adopted by the Madison Botanical Congress in 1893 and by Section G, A.A.A.S. in 1894. In many cases of confused synonomy, critical examinations were made of authentic specimens and the related literature. Notes of interest obtained thus are added under the Species in question. The present part, which is part I of the first volume, includes names from Abrothallus to Badhamia.—H. HASSELBRING. S Fartow, W. G., Bibliographical index of North American fungi. Vol. J, part 1. 8vo. pp. xxxv+312. Washington: Carnegie Institution. 1905. 76 BOTANICAL GAZETTE [JANUARY MINOR NOTICES. Japanese vegetation —Professor Mryosut, of the University of Tokyo, has begun the publication of photogravures of Japanese vegetation,® to represent wild and cultivated plants and plant societies. Each picture is on a separate sheet of cardboard 20.5 27°™, the size of the print being 16X23°". Accom- panying the illustrations is a descriptive text in both English and Japanese. The author has not yet determined the number of plates to be issued. So far, two parts have appeared, part I containing eight plates of cultivated and semi- cultivated an and part II containing eight illustrations of the vegetation of the island o The illustrations are well chosen and well made. Among the most effective and characteristic are the long avenues of giant mountain cherry trees, gorgeous . with their spring blossoms, the graceful bamboos bending beneath their burden of winter snow, and the forest vegetation around the Hannya waterfall. The descriptive text is precise, and interspersed by interesting remarks which show that the ae has an eye for color and setting. e hoped that the series may be continued to give us many more Testes of the flora of this interesting country.—F. C. NEWCOMBE. A botanical cyclopedia—-An illustrated German dictionary of botanical terms has appeared under the editorship of Camitto K. SCHNEIDER,’ with the assistance of a number of other German botanists. This volume of almost 700 pages presents much more than a list of definitions, for there are illustrated descriptions of the morphology and minute structures of organs, of the sort one would expect to find in a cyclopedia. The terms, of course, are those em- ployed in the German language, and the work will not take the place, for the English or American botanist, of Jackson’s excellent Glossary of botanic terms. —B. M. Davis NOTES FOR STUDENTS. Chemotaxis of spermatozoids.—The chemotaxis of the spermatozoids of Isoetes has been studied by Surpata.8 In Jsoetes japonica, which was used for the study, the sporangia ripen in autumn. Microspores, sown in tap water in Perti dishes late in November, begin to germinate about the middle of January. The duration of the swarming movements of the spermatozoids is shorter than in the ferns, vigorous movements lasting only about five minutes; some movement of 6 Miyosur, M., Atlas of Japanese vegetation. With explanatory text. Tokyo: Maruzen Kabushiki Kaisha. 1905 7 SCHNEIDER, C. K., Illustriertes Handwoérterbuch der Botanik. Imp. 8vo. pp- 690. figs. 341. Leipzig: Wilhelm Engelmann. 1905. M 16. 8 SHIBATA, K., Studien tiber die Chemotaxis der Isoetes-Spermatozoiden. Jahrb. Wiss. Bot. 41:561-610. 1905. SE ae a ee a y a ‘ 1906] CURRENT LITERATURE 77 the spermatozoid, however, may continue for ten or fifteen minutes, and of the cilia for five minutes longer. PFEFFER’s capillary method was used in the experi- ments. The principal headings are: position chemotaxis, relation between the intensity of the stimulus and extent of the reaction, repulsion by free acids and alkalis, negative chemotaxis with the ions of heavy metals, repulsive effect of alkali salts, behavior with osmotically acting substances, repulsive effect of ions of certain organic acids, the action of narcotics, theoretical, and review. Malic acid acts as a strong topochemotactic stimulus and may be regarded as the specific stimulant for the spermatozoids of Isoetes, although certain other substances also exert some topochemotactic influence. Free malic acid in weak solutions exerts a positive chemotactic influence, but in stronger solutions a nega- tive one. The salts of various metals act as negatively chemotactic stimuli, as do also the anions of di- and tribasic organic acids, including malic acid. The positive chemotaxis with malic acid is of a typically topotactic nature. The reaction consists in a turning of the body axis of the spermatozoid and a movement toward the source of stimulation. Whether the structure for the perception of chemotactic stimuli consists of the whole body of the spermatozoid or only of localized portions of it is not yet determined.—C. J. CHAMBERLAIN. Tuberization.—The causes of tuberization still furnish a field for study. BERNARD first supposed that Fusarium Solani was the endophytic fungus of the potato; this has since been disproved by GALLAUD and by BERNARD himself, but the identity of the fungus is still undetermined. H. Jumette® has been con- ducting experiments on Solanum Commersoni, a tuber-bearing species related to the potato, but as yet his results are largely negative. The chief interest attached to his studies are occasioned by the fact that S. Commersoni has small slowly developing tubers placed on long stolons; these are the very characters which the potato is said to have had when first introduced into Europe, before the endo- phytic fungus became sufficiently abundant. S. Commersoni was infected by fungi from S. tuberosum, but, as stated above, with negative results. JUMELLE thinks that with suitable infection, it may be possible to secure tubers like those of the potato, and further experiments are in progress. It should be said that GALLAuD thinks that BERNARD has not yet isolated the true tuber-forming fungus.—H. C. Cow es. Two parasitic fungi—KirBaHNn'? has worked out the life histories of two common species of the so-called Imperfecti group. The first of these is the common elm fungus, Phleospora Ulmi (Fr.) Wallr. This is connected with an ascomycetous form, which appears on the infected dead leaves during the winter and ripens in spring, when the spores are ejected and infect the young 9 JuMELLE, H., Del’influence des endophytes sur la tubérisation des Solanum. Rev. Gén. Bot. 17: 49-59. 1905. to KLEBABN, H., Untersuchungen iiber einige Fungi im perjecti unddie ea Aivtalapeles Rais I. u. Il. Jahrb. Wiss. Bot. 41: 485-560. figs. 75. ™ 78 BOTANICAL GAZETTE [JANUARY elm leaves. The form is named Mycosphaerella Ulmi Klebahnt'. Both conidia _and ascospores produced identical mycelia in cultures. The Phleospora was produced by sowing ascospores on the under side of elm leaves. No infec- tion took place from spores sown on the upper surface. The study of Gloeo- sporium nervisequum (Fckl.) Sacc. revealed a rather complicated series of forms belonging to this fungus. An ascogenous stage develops on the dead leaves, asin Phleospora. This is Gnomonia Veneta Klebahn. Beside the usual conidial form and the ascogenous form, the fungus assumes thc form of a Myxosporium on the young branches, and there produces the twig wilt always noticed on sycamore trees affected with the Gloeosporium. A fourth form develops on the dead leaves. This is a conidial form of the Fusicoccum type. Proof of the connection of all these forms rests mainly in the similarity of the mycelia produced in pure cultures from the various spore forms. Infections could not be pro- duced readily, but a few cases of inoculation with ascospores were successful. The various spore forms have been described under different names, which are given in the synonymy.—H. HAssELBRING. Shore formations in Denmark.—WarmIN«, in collaboration with WESENBERG- Lunp and others in an interesting paper, has correlated the work of plants and animals in the shore formations of western Denmark.t? A “vad” is a shallow coastal lagoon, cut off from the sea proper by a line of islands, and bare at low tide; the bottom may be of sand or clay, the latter type prevailing in the more tranquil places. The sandworm, Arenicola marina, is the most characteristic animal of the sandy “‘vader”’ or shallows, and the excrements of this worm are found there in great abundance. Hence it has commonly been thought that these animals have a soil function, similar to that of the earthworm, and that they help to build up the shallow into a marsh. The authors, however, find that Arenicola is very sedentary in its habits, swallows only from surface layers, and that it retards rather than furthers soil enrichment. The waves wash any fine particles landward, leaving the Arenicola shallow as sandy as before and hence well adapted for the continued prosperity of Arenicola. In shallower water closer to the shore, where the bottom material is finer, the amphipod, Corophium grossipes, dominates; here the bottom is characteristically red-brown in colo1 and presents a riddled surface appearance. e Corophium shallows teem with animal life, and here the influence of the animals is such as to convert the area somewhat rapidly into a marsh. To this work blue green algae contribute some- thing, but animals are much more important land builders than plants in the Corophium shallows. Large areas of land have been gained from the sea in this way in Denmark, and one case is cited where a fertile meadow has been developed from a barren sandy shallow within two hundred years. In the argillaceous tt Preliminary note in Zeitschr. Pfl. krankh. 12:257. 1902. 12 WARMING, E., Bidrag til Vadernes, Sandenes, og Marskens Naturhistorie. In collaboration with Wesbrnies: LuND, OsTRUP , et al., with French résumé. Ko ngl. Dansk. Vid. Selsk. Skrift. VII. beak 1904. , 1906] CURRENT LITERATURE 79 shallows, in contrast to the above, plants are the more efficient land-builders; the developmental processes in such places are well known and need not be recounted here. ARMING also speaks of sandy plains that are subject to occa- sional inundation. Here algae play a great part in soil-making; it is common for a layer of Phycochromaceae to penetrate for three to five centimeters into the sand, cementing the grains together, and giving a greenish appearance to the ground. Many of the diatoms characteristic of such places are listed by habitats. The peculiar depressions of salt marshes, called ‘‘pans”’ by OLIVER and TANSLEY, are thought by WARMING to be formed where heaps of decaying vegetation have lain; the consequent destruction of the vegetation makes it easy for the waters to wash out the soil in such places.—H. C. Cow es. Periodicity of sexual organs in Dictyota.—WILtAqs, in the third contribution to his series of Studies in the Dictyotaceae,'3 discusses the remarkable periodicity in the formation of the sexual cells in Dictyota. The sexual organs are produced during the fruiting season in fortnightly crops, synchronous with the spring Sa and a general liberation of the gametes takes place on a particular day, at a interval after the highest spring tide, varying, however, in different localities Of the factors (temperature, pressure, aeration, etc.) which fluctuate with the alternation of neap and spring tides, the one which seems to account most satis- factorily for the facts of periodicity is the increased illumination of the plant during the low water of spring tides. The times of initiation and liberation may be slightly accelerated or retarded by exceptional meteorological conditions, as when wind causes a difference of two or three feet in the height of water, or a rise of one inch in the barometer accompanies a depression of six or seven inches in the tide—B. M. Davis. Brown pigment of algae.—The generally SE view that the color of the chromatophores of the brown algae and diatoms res the presence of a brown pigment, phycophaein, in addition to " Madipbyll is challenged by Mottscn,'+ who believes that the phycophaein garcons from these algae is a post mortem product. He holds that the brown pigment is a + tie substance, phaeophyll, which passes readily over to chlorophyll ates poabole with hot water, alcohol, and other fluids. A similar brown pigment is found in the orchid, Neottia nidus-avis. Beside the phaeophyll, 20 brown pan “ag diatoms contain carotin and a bluish-green pigment, leucocyan racts of Bennettites —LicNier's from a reexamination a his preparations of che involucral bracts of Bennettites Morierei concludes that in all the sterile scales, superficial or otherwise, which enter into the composition of the strobilus, the terminal enlargement is due to hypertrophy and does not result from a reduc- tion of the bract.—W. J. G. Lanp. *3 Witiiams, K. L., Studies in the — tik; The origami of the sexual cells in Dict yota dichotic’. Ann. Botany 19:531-560. figs. 6. 1905. ™4 Motiscn, H., Ueber der braunen behead der Phaeophyceen und Diatomeen. Bot. Zeit. San. sone pl. O- 1905: 1SLIGNIER, O., Notes aaamacet ines sur la structure du Bennettites Morierei. Bull. Soc. Linn. Norwaidie V. 8:(pp. 7.) 1904- NEWS. PROFESSOR EDUARD STRASBURGER traveled in Egypt during part of December and January. Dr. OskaR BREFELD, professor of botany at Breslau, has retired owing to failing eyesight. Mr. WALTER FISCHER has resigned his position as Assistant in Botany at the Ohio State University, and has taken up work in the United States Department of Agriculture, Bureau of Plant Industry. Dr. A. F. BLAKESLEE, who is spending the winter in investigations at Halle, was awarded the Bowdoin prize of Harvard University ($200) last spring for his work on sexual reproduction in black molds. In November and December Dr. Joan W. HARSHBERGER, of the University of Pennsylvania, delivered a series of five lectures on “Weird and. marvelous plants” in the Ludwick Institute courses of free lectures, Philadelphia. Dr. Ernest A. Bessey has been transferred from Washington to the Sub- tropical Laboratory of the U. S. Department of Agriculture at Miami, Florida, where he may be addressed in future. Professor P. H. Rotrs, formerly of the Sub-tropical Laboratory has accepted the directorship of the Agricultural Experi- ment Station of Florida. A RECENT circular gives an account of the Royal Hungarian Central Institute of Viticulture, the buildings of which were completed in 1904. This institute was initiated in 1896 by a law decreeing the establishment near Budapest of an institution for studying the problems of viticulture and wine-making, and giving scientific and practical instruction in these subjects. In 1898 Dr. IstvANFFI, then professor at the University of Kolozsvar, but perhaps better known from his association with BREFELD during part of the latter’s extensive investigations, was called to organize and direct the new institution, whose first work was done in quarters rented until the completion of the new ones. The present buildings are five, each of three stories. The main hall, 66%21™ contains the library, museum, offices, and lecture room. The four smaller halls, all similar in con- struction, are devoted respectively to the four sections: (1) physiology and path- ology, (2) chemistry, (3) zymology, and (4) practical viticulture and oenology. The institution is excellently equipped for carrying on these different branches of work. Its primary object is that of an experiment station whose field is restricted to the wine-growing interests alone; secondarily, provision is made for instruction to advanced students in the practical aspects of viticulture and oenology. [1906 80 Oe NERVOUS DISORDERS The nerves need a constant supply of ae ary to keep them steady and A Soieriec f of the phosphates a ms a lowering of nervous tone, indi- cated by exhanstion, slonioes eat: ache or insomnia Horsford’s Acid Phosphate (Non- Alcoholic.) furnishes the phosphates in a pure and abundant form. It supplies the nerve with near giving life force Bi Sea ste, restores the strength a nd indue restful ee without the use of danger- r- edhe drugs. 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Y. ee ee MEDICAL OPINIONS OF “All the Argument Necessary.” The International Journal of Surgery, August, 1905, under the heading “Cystitis,’’ says: ‘‘In the treatment of Cystitis, water is the great aid to all forms of is the ideal form in which to administer medication. Moreover, BUFFALO LITHIA WATER it to the Cystitic patient, as it is not only a pure solvent, but has the additional virtue of containing substantial quantities of the Alkaline Lithates. Patients should be encouraged to take two quarts per day, if they can, and the relief they will obtain will be all the cocci necessary after the first day or so. “The Results mcr Me of Its megEay Value.” the Bladder and Unitiere: vir females. The results satisfy me of its pfs rae value in a large class of cases usually most difficult to treat.’ “I Have Witnessed Decided Beneficial Results from Its Use.” Wm To owles, M.D., formerly Professor of Anatomy and Materia Medica of the University of Vir- are marked in causing a disap- ginia: ‘‘ The effects of pearance of Albumin from the urine, and in shea eer of Bright’s Disease | have witnessed decided beneficial results from its u “Results, to Say the Least. Very Favorable.” T. Griswold Comstock, A. M., M. D., St. Zouis, Mo., says: ‘I have sce or BUFFALO LYTHIA WATER £2,°7<<2.°Esanic conditions, with results to say the least, very favorable.” Additional medical testimony on request. For sale by the general drug and mineral water trade. PROPRIETOR BUFFALO LITHIA SPRINCS, VIRCINIA- Holds America’s Highest Prize Consumption 1 Pneumonia -4. 9 Walter Baker & Co.'s are preventable! | Breakfast Cocoa Finest in the Just a O° ‘chp ices nl little! Y In every living spe keep an open or. Platt’s Q vessel containin ter and Chlorides © P 9 AG chet you? Platt's atts nothing by 9 Chlorides ; Q AWARDS IN sickness. 4 oa is surcche ae Sold in a and de Cans WEIGC WALTER BAKER & CO. Limited Established 1730 DORCHESTER, MASS, THE DAINTIEST SOAP MADE is HAND SAPOLIO for toilet and bath. Other soaps chemically dissolve the dirt—HAND SAPOLIO removes it. It contains no animal fats, but is made from the most healthful of the vegetable oils. It opens the pores, liberates their activities, but works no chemical change in those delicate juices that go to make up the charm and bloom of a perfect complexion, Test it yourself. THE FAME OF SAPOLIO has reached far and wide. Everywhere in millions of homes there is a regard for it which cannot be shaken. Sapolio has done much for your home, but now for yourself—have you ever tried HAND SAPOLIO, for toilet and bath? It is related to Sapolio only because it is made by the same company, but it is delicate, smooth, dainty, soothing, and healing to the most tender skin. It pleases every one- ITS USE IS A FINE HABIT—ITS COST BUT A TRIFLE a ARS. By our system? L/1C GO PIANOS38 23222 | Vol. XLI A Se BOTANICAL GAZETTE | February, 1906 | THE /00 ) Editors: JOHN M. COULTER and CHARLES R. BARNES A gl _ CONTENTS Chemotropism of Fungi Harry R. Fulton a eee The Embryology and Development of Riccia lutescens and Ricca crystallina (with plates V-IX) Charles E. Lewis Briefer Articles Note on the Relation between Growth of Roots and of Tops in Wheat Burton Edward Livingston Current Literature iad News The University of Chicago Press CHICAGO and NEW YORE William Wesley and Son, London Purity and Pears - The best of Pears’ is purity; freedom from everything adulterant or injurious, and no free alkali—TThat is how Pears’ refreshes 2m invigorates the skin, enabling it to be healthy and pure, that complexion which, like the snow, is matchless — creating — OF ALL SCENTED SO. pee ost meee “All rights secured.” ai APS nin red OTTO OF ROSE IS THE BEST. — Che Botanical Gazette A Montbly Fournal Embracing all Departments of Botanical Science Edited by Joun M. CouLter and CHartes R. BARNES, with the assistance of other members of the botanical staff of the University of Chica Vol. XLI, No. 2 Issued March 3, 1906 CONTENTS CHEMOTROPISM OF FUNGI. Harry R. Fulton -— - es et gine eae 81 THE EMBRYOLOGY AND DEVELOPMENT OF RICCIA LUTESCENS AND RICCIA CRYSTALLINA (wirH PLATEs V-Ix). Charles E. Lewis - - 110 BRIEFER ARTICLES. OTES ON THE RELATION BETWEEN GROWTH OF ROOTS AND OF TOPS IN WHEAT. CON- TRIBUTIONS FROM THE Hutt Botanica, LAporaAtory. (LXXXI WITH FIVE ston FIGURES). Edward Burton Livings n : : . - | 9 CURRENT LITERATURE. BOOK REVIEWS eee ; i icneuk te tc Si SY LOSS 8 Gs THE SWISS MOORS. REPRODUCTION OF MILDEWS, MINOR NOTICES - - - - - - - - - - - tl < - 146 NOTES FOR STUDENTS - - - - - - - . - - - ~ - 149 NEWS s Be : . “s : : yh sé “ ‘ a : ue ‘ =; “ESD acne for the Editors should be addressed to them at the University of Chicago, Chicago, III. Contributors are requested to write scientific and td names with icrtictdlar Tr care, eh use the he System of weights and measures, and in citations to follow the form shown in the pages of the BoTaAN AZETTE Separates, if desired, ee be ordered in advance of publication. 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In all cases the memoirs are to be based on on a considerable body of original and blished work, accom- eid iby a a general view of a pe ae of the ro hi hh, i? 4 Any thing in the mem ring the essay caps, competitio f the identity of the author shall be considered as debar- fad Preference will b 54 i I f being t d Pp I made direct] ly in competition for the prizes Each memoir 1st of on year for which the prize is offere: The ‘o one borne by ‘the cpt Bade sie must res in ecg y bere' SF . Sacer on or before Prser- d, ociety assumes no en gear for publication of tamer submitted, BJECTS FOR 1 me and superscribed w x. An experimental re “study i in riba tig 2, Acontribution to ig petition in plants, 3- A physi cap cal life h history f a single sp plant 4. Phylog f il organi tre” study i in stratigraph 6. A research in minera Et 7. A study of entectics in pak penta, 8, A study in river capture SUBJECTS FOR 1907: = s, Ss} tid “J Pe ra S| dé le le le BS) BS [gael gS SPECIES Eacl S| | 3 FaaclZacl“ac| “nel Sas] J |. les a} O |O hse Fa|.8 Mo a\e8 g au 4 SSIS [g lg Bs) & geige o ae < a BAW l< Sterigmatocystis nigra.......... Fire . Po a ee ps r Mucor mucedos. Yoav eeu: jvc) Qe fest 4 ae lect YF | Bel ae) Ax] Bat Oa) Se Botrytis vulgaris... cic. es cok fla [ocebessdeks r ily te SEAS Bi nee banca lear ae ee : Pe RSS ae x bigest uted a a eI Te fee IB ine hy, S He ne REN é i 1906] FULTON—CHEMOTROPISM OF FUNGI 89 affected. The repellent effect was much less marked, never affecting more than one-fourth of the hyphae, and even in the most marked cases some of the hyphae would grow toward the tubes and enter them. Table I gives the most marked instances of attraction and repulsion noted in the series; the symbols have been explained above. In Table II is given a comparison of the effects of a number of representative compounds as observed by Miyosnr and in the present investigation. Leaving out of consideration Saprolegnia jerax, the extreme effects recorded in Mryosut’s table were obtained TABLE It Aspexott- |__ STERIG- Mucor MUCEDO LUs NIGER | MATOC¥STIS REAGENT NIGRA I i iil IV onobasic sree phot, 2 Joss. a3 ae ar ar Monobasic sodium phosphate, 2 %....... ae a2? ed bi utral ammonium oles phate, 2%...... ay az a2 as al ammonium phosphate, 0.2% ee ax ee ar Neutral ammonium phosphate, 0.1 %.. ay os ciate Pirate, By eee es eae G a3 az? az r Potassiuin nitrate, ¢ 0... 2... 2 es seca r ay r _ Calcium nitrate, 29%... 60 0c eee e e ar Tr . Magnesium sulfate, 2%.........++...+- r fc) r r € ice Mike eo se we se ew eeee a3 ° ar ° Potassium chlorid, 2%... 02.02.0000 r ° r 2 odi OM Bek usivi'g on aees os r ° r ° um-potassium tartrate, Oe Nee eee r a2? r ° Cane sugar, 2 A epee avccste sey Garg wks ta ae ay ° ar ° 2a ree tetera we cae a eae a — = ee Epeloe Sat ee bs eee ae ° ae ° us iy Boy Pape ee ae eg a3 ar a2 ar? ad = pe Paine © pe ere ay *e - Lactose, 2 2% Oe ence es a, ay? vee Oey kk MESON chivas ESET ok week ay ay? Malice’ 2 %, Rohe Wire hw eee e Wie ay ete ay ° see ER 2 Sear ee Ey tee ro fs) ax? Dextrose, $0 Geet ee a, aa ae ies OR een gr ea er ee ° ay a 2 %, WOKS Ste Big ce ee a ae eee Bae ag a3 5h 1% ee ee a3 r ? ay ° Ethyl alobol, Wists paSiy penne aces z a2? r ay? — i eee ele ao a kee a eee we we ay 3° ay ? ay Fi a SEE SS SRUREN, Manian eae tae pe ag a2? ax ar? 2 oF EEE SEE acer pene eas te Sees ° ay? ° ay? . Peptone, re eae ° aot az? J M. - % Peer w ere e sere ress eerteonese a3 a2 mae leat extract, ta ee RES re ere ay ag ag ? 1% a3 ar ag ay es eS aries ay ° as af Rs ae eee ay ° ° ar? go BOTANICAL GAZETTE [FEBRUARY with Mucor mucedo and Aspergillus niger; the values given by him for these forms are to be found in columns I and III below. It was impracticable to use Asperillgus niger in the present study, and ' Sterigmatocystis nigra was substituted with the understanding that the two forms are not always distinguished. The present results for Mucor mucedo and Sterigmatocystis nigra are given below in columns II and IV respectively. ' In the control cultures, where distilled water was used in the tubes, the effect was the same as for the majority of chemical sub- stances; that is, 10-30 per cent. of the hyphae turned toward the tubes. The same amount of positive turning was observed in the case of all the strongly toxic compounds used. Even with a 0.05 per cent. solution of mercuric chlorid and a 1 per cent. solution of copper sulfate, which completely inhibited germination within a radius of eight to twelve tube diameters from the openings, the hyphae not only grew across the diffusion areas, but 10-30 per cent. of those approaching the openings turned toward them and grew for a con- siderable distance into the tubes. Although four concentrations of cane sugar ranging from 20 to 0.1 per cent. and four concentrations of meat extract ranging from 10-0.01 per cent. were used, no definite relation between the strength of stimulus and that of response was apparent. Two corresponding series were made with ten representative compounds; in one series sugar-beet agar was the culture medium; in the other, distilled-water agar. No difference in the behavior of the hyphae due to a difference in the media in which they grew could be observed. Tests with mica plates —Thin sheets of mica were cut into pieces about 25X16™™; these were perforated with a needle, the holes being 0.1-0.15™™ in diameter, and about 2™™ apart. Covers of suitable size were cut from glass 1™™ thick. A layer of gelatin or agar was placed on a cover, a mica plate was placed on this just before it hardened, and a second layer was placed above the plate. The chemical to be tested was made up in double strength solution in redistilled water, and one volume of this was added to one volume of gelatin or agar, also made up double strength in redistilled water. It was usually found convenient to have the layer containing the ice Ce Seen: ce . 1906] FULTON—CHEMOTROPISM OF FUNGI QI chemical next to the cover, the spores being distributed in the outer layer. The cover was inverted over a stender dish of 36™™ diameter containing distilled water, and was sealed with a coating of vaseline around the rim of the vessel. This method seemed to possess distinct advantages over the one with capillary tubes: the numerous perforations made it possible to make a large number of observations from a single preparation; TABLE III 5 s S g 2) E 8 4 & z g z : Ba<| & 2 Ae |p E 4 i] no ae ea = i = a im Neutral potassium phosphate, 2 2% ecesas 12 19 24 22 33 ee A a s3 20 18 25 36 25 Monobasic potassium phosphate, 2 2 fe =} ag 22 24 26 36 23 Ds| 20 26 23 20 25 21 Dibasic ammonium eS hate, 2 % 13 20 2I 20 15 23 ane ee 2% ie saped eweaes 49” | 41 34 31 35 Bd Phosphorie acid, 6.9 9.06054 less cn ces 48’ | 30 35 39 23 3° Malic acid, r 1% AP Serra ali, le te ee x x 24 x 38 x e me | day aie et teh tare Oe 21 21 26 21 17 x Tartaric acid, 170 per ne Re eae Ne ae pe * 20 43 22 19 18 5 7 eicaie ee el 25 27 34 25 24 22 Oxalic acid, 1 es eet er aan x 17 23 x 17 0 Rcainintt em aaianad Ges 23 22 23 34 II 40 Cane sugar, 5 $ Yor. iver sSa mew ea eas waa nee 38 32 24 31 a0 48 BE el. ca pene oe 30 37 20 30 20 55 . Sp ee ee ere eee ra ee 30 35 24 a2 23 Glucose, 5 o. Rea Segre Grape igs Dog oa are a 39 39 20 34 30 37 et Gnic aGere sete ee ce un 40 24 23 39 20 3° POPRONE BT Senne 14 28 21 13 20 19 : eee ais ws Son cs Wee eas ous 26 23 20 21 13 22 Pepsin active Nig ae ae SSE 24 20 19 42 23 30 te ; 3 Sey MCLE as pee 25 30 22 27 31 22 Potced Sopa 2 %.. eee isn neers a6 t.. 37 ah 33) 4 | ee Aspa: dy OR analaeee eireeterer acer tty 19 23 30 23 30 15 be ey ot as 15 23 30 20 30 24 es sulfate, 0.00 % Nees ae wae aie 9 21 21 25 34 31 Dibasic sodium phosphate, 2 2% ors uaa? 35 25 26 26 15 20 mew wees 38 Ki) 27 20 26 a3 Starch vag Mek. WORE ROT OE Tee 24 20 21 27 25 te a see ge tree cee . 27 22 20 26 15 Maltose, 2 2%, Sree ee earn Reena 33 30 22 21 25 14 pee eect aire e sss 36 29 28 31 30 28 Control (lestilbad wate @elatin) ©... 5 60s 28 30 29 24 22 18 Note: gana X indicates that germination was inhibited. Uromyces caryophyllinus was tested with the majority of these substances, in the few instances in which growth was sufficiently great for a determination, the — was practically the same as for the other fungi. g2 BOTANICAL GAZETTE [FEBRUARY the medium containing the chemical and that containing the spores could be more nearly equalized in amount and in consistency; fewer hyphae would take a course that would lead them through the open- ings without apparent turning; better sterilization could be secured; and there was less difficulty in making up the preparations. In making the counts, hyphae within a radius of one opening diameter from the margin of each opening were considered; the hyphae within such an area were classed in the counts as those turning toward the openings, those turning away from the openings, and those apparently indifferent. After an examination of the entire preparation in each case, those holes were selected for the counts which represented the average condition. In calculating the percentages from the counts, the difference between those attracted and those repelled was made the dividend, and the total number within the observed area was made the divisor. The results are shown in Table ITI. If the percentages of turning toward distilled-water gelatin as determined by the control experiments, be deducted from the per- centages of turning toward the various chemical compounds, it will be found that in only three instances would the difference, which would be supposed to indicate the amount of turning due to chemical influences, approximate 25 per cent., or about the average of the lowest of the several degrees of positive chemotropism recognized by Mryosui. Most of the values, even for supposedly highly attract- ive substances, are very near the controls. Evidently, the results thus far have not been favorable to the theory of chemotropism. But it was thought that the fungi, all of which grew perfectly normally in gelatin made up in distilled water, might turn more strongly from some other medium in which there was less of available nutriment, to one having an abundance. In agar made up with distilled water, the fungi germinated after a greater length of time and grew more slowly; but the turning from this non-nutrient agar toward a nutrient salt solution favorable for fungous growth was no more marked than from agar containing the same proportions of the nutrient salts to the nutrient salt solution; nor did the nutrient solution seem to attract from either medium more strongly than ‘did distilled water. This test was made with 1906] FULTON—CHEMOTROPISM OF FUNGI 93 Sterigmatocystis nigra, Mucor mucedo, and Mucor stolonijer by the capillary tube method. | It is possible, although there is ‘evidence against it in some of the previous experiments, that the diffusion of the solutes might be rapid enough to bring about practical uniformity in the media before the germination of the spores. To determine whether the time at which the stimulus is applied has an influence upon the reaction, series were made up with two of the more slowly growing species, Botrytis vulgaris and Monilia fructigena, the spores being distributed in non-nutrient gelatin placed above the mica plates. Four duplicate series were arranged. In one the layers of gelatin containing the substances to be tested were placed below the plates at the time of making up the cultures; in another series these layers were added just as the spores began to germinate; in a third, after the most vigorous hyphae had attained a length of 15 to 20 spore- diameters; in a fourth when the length equaled 40 to 50 spore- diameters. The final counts were made a little later, when the growth was about 75 spore-diameters. The results are given in Table IV for the five compounds tested and the control. The numbers in columns I indicate the average number of hyphae in the area around each hole; those in columns II, the percentages of turning toward the holes. TABLE IV Non- Hi Ox KH,PO (NH,)NO sve Beat] Paar | SSG [mua | MEO | ON | come Cres | Appli- cation ‘ i | ue | I Ir | I | n | I | It I | II I It 8 of} ar | 35 | xz 38 | 6.3 4-9} 37 | 5-5 | 36 | 9-2 | 34 a3 £0. 9.6). 38° 1 Gio 1238 Te. | 36.1 §4.) 30 | 72} 36-1 7-61 34 e6| 13 a) ge fee Tae tat 95.1 6.4.) 3t $14: 33.4 9.21 38 AE! x6 || 3.3| 30 | 7-6| 34 | 5-5] 3015-8] 35 | 5-0| 32 | 5-6 | 36 < Sa0.0 UG.g1 384-260-140 1 tr] 98 4 3.7) -3t.| 8:2 | 34 | 4-4 [06 Be 10 9-5 | 36 1 3x ao 4 Es ao. }.3,7 | 38--1.6.8 | 25) 21 -| 32 S8| ts | 13 | 33 |7-s| 27 | 10 | 32 | 4-3| 30 | 5-4] 27 | 33 | 27 i} 20 fT 32 ay 16.47 8615-5 1. 20 1 8-1] 26 | 8-3 | go It is evident that the time of application has little or no effect upon the amount of turning. It will also be seen by comparing these 94 BOTANICAL GAZETTE [FEBRUARY results with former ones, that it is immaterial, as far as the per- centage of turning is concerned, whether the spore-containing layer is above or below the one containing the test substance. Other culture media were used, such as 10 per cent. and 25 per cent. glycerin, gelatin made up to contain 10 per cent. of glycerin and to contain 5 per cent. of cane sugar. With none of these were there more distinct indications of chemotropism than in former tests, in which non-nutrient gelatin was the culture medium. This would indicate that the available nutriment and the concentration of the medium have no effect. The final test along this line was made with silica jelly, a medium free from organic matter and of suitable consistency. ‘The method of preparation was that used by Moore (23), except that dialysis was accomplished satisfactorily with parchment paper. In order to secure the proper coagulation of the medium, it was necessary to add mineral salts to all media used. A nutrient salt solution, containing 18“of ammonium nitrate, 0.5%" of monobasic potas- sium phosphate, 0.25%" of magnesium sulfate, a trace of ferric chlorid, and 58" of glucose, in a volume of 1o0°*, was made the basis of the work. It would seem that if fungi show chemotropism under any conditions it would be by turning from a medium lacking some one or more of the elements necessary for full development, toward the diffusion centers of compounds supplying the missing element or elements, or toward a full nutrient solution. In the tests each compound in turn, except ferric chlorid, was omitted from the silica jelly containing the spores, and in each case jelly containing the omitted compound in proper proportion on the one hand, and full nutrient jelly on the other were used on the opposite side of the mica plate from the above-mentioned spore-containing layer. Control tests were carried on with jelly lacking one and the same compound on each side of the plate, and also with full nutrient jelly on each side of the plate. To reduce evaporation, the lower layer was covered with an unperforated mica plate. The results are given in Table V, in percentage of response by turning from _ the first-named medium in which the spores were sown to the con- trasted one. There is a very striking uniformity in the percentages, and this 1906] FULTON—CHEMOTROPISM OF FUNGI 95 TABLE V ne =< a Ho < & B 5 e a| < F B : E CONTRASTED CULTURES IN SILICA JELLY “u& Bog 4B ao #65 opegen waa: 2 at ER Oe |aea Ze ae Ze pa 3 on 5 om a A a a Full nutrient : full nutrient .....2...7........ 35 32 32 33 20 acking glucose : aesiie pete ce Poe 32 36 37 3° 33 Lacking glucose : containing glucose........... 27 3° 34 oO 26 Lacking glucose : full nutrient................ 3 36 33 3° 3° Lacking KH,PO, : lacking KH,PO,.......... 92.09 40 Sy On ae Ae Lackin aPO, de pein me PO, eran 36 36 31 30 28 Lacking KH,PO, x fall watrient. toca: ewer 32 a0" 3g 36 | 28 Lacking (NH,)NO, : lackin ae “CNET Ale Saree as ee a ae Ge we 6 | 28 Lacking (NH,)NO, : ati aes )NO, eto a 32 32 33 29 Lacking (NH,)NO, e full nutiaent is. escras ere. 34 33 33 She 3? Lacking MgSO, : lackin ng MgSO er ee 33 ar 33 a3 27 Lacking MgSO, : containing MgsO, setae ade is 39 34 33 33 29 Lacking MgSO, : full nutrient................ 29 33 32 35 3° under conditions that would be presumed to be most favorable for chemotropic reaction. Tests with epidermis and cabuan jilms.—To test the effect of physically different perforated sheets as well as effectually to repeat the methods used by former investigators, use was made of celloidin films which had been perforated, and of strips of epidermis of Yucca aloifolia. This gave, with reference to physical properties, a range from the wholly impermeable mica plates on the one hand to the semipermeable celloidin films on the other. The tests with epidermis were made with Monilia fructigena, Sierigmatocystis nigra, Botrytis vulgaris, Sphaeropsis malorum, and Mucor stolonijer, the spores of which were distributed in non-nutrient 8 per cent. gelatin above the epidermis in its final position; gelatin layers containing 5 per cent. cane sugar, 4.5 per cent. dextrose, 0.01 per cent. copper sulfate, 0.1 per cent. oxalic acid, 0.2 per cent. phosphoric acid, and non-nutrient gelatin, were spread below the epidermis. Under these conditions the penetration of the stomates or turning toward the stomates was practically zero. When no culture medium was used, the spores being merely spread on the under surface of the epidermis, hyphal growéh was good. A few hyphae of each species grew through the stomates; but there was no evident turning toward them, and in no case was there pene- tration of more than one or two per cent. of the stomates. 96 BOTANICAL GAZETTE [FEBRUARY In similar series with celloidin films, the turning from one gelatin layer to another was about equal to that obtained with mica plates. When the spores were spread on the film without a culture medium, very few of the hyphae grew through the holes, the percentage of turning being negligible. The hyphae in these cases were sur- rounded by a distinct film of condensed moisture. TESTS FOR OTHER FACTORS. The tests thus far have failed to give evidence of the existence of any marked chemotropism. There has been at the same time a considerable and fairly constant turning of the hyphae from a medium containing spores to a sterile medium, when these were separated the one from the other by any one of several partitions. Since this turning is apparently unaffected by the chemical relationships of the media employed, the cause of the turning must be sought in other factors. Two possibilities at first present themselves; the mechanical partitions may have a thigmotropic or other influence, or the germinating spores themselves may affect the direction of owth. Tests without mechanical partitions —A slight modification of the method employed by CiarK (4) was used. A large drop of agar, 8™™ in diameter, was placed in the center of a sufficiently large square of glass; this drop was surrounded by four drops of about 5™™ diameter, equidistant from the first, and with a space of about 3™™ between each smaller drop and the larger one. Non- nutrient agar and to per cent. sugar-beet agar were used for the drops, and were arranged in four combinations: the central drop was of nutrient agar and two small drops diagonally opposite each other were of nutrient agar, the other two being non-nutrient; the central drop was of nutrient agar, two small drops adjacent to each other of nutrient agar, and the other two of non-nutrient agar; two similarly arranged combinations had central drops of non-nutrient agar. The fungi used were Monilia sitophila, Mucor stolonijer, and Botrytis vulgaris. A few spores were sown with the platinum ‘needle at the center of the large drop in each preparation; the cover was inverted over a Stender dish containing distilled water and was sealed to its rim. The growth was watched until the hyphae had grown about two-thirds of the distance from the center to the margin 1906] FULTON—CHEMOTROPISM OF FUNGI 97 of a large drop; the preparations were then opened, and with a sterile needle the small drops were pushed up until their edges came in contact with the larger drops. Later observations showed that the hyphae of the three fungi grew readily from either medium into a similar or a dissimilar medium, and with the same percentage of turning. An equal amount of turning toward the agar drops was observed in the case of those hyphae which had grown beyond the bounds of the larger drops on the moist glass; whether the agar was nutrient or non-nutrient seemed immaterial. The turning was apparent at a considerable distance from the surface film in so large a percentage of cases as to negative the supposition that the physical condition of the film has an influence. A further test without mechanical separation was made by placing small crystals of cane sugar, copper sulfate, oxalic acid, monobasic potassium phosphate, and ammonium nitrate in the center of layers of non-nutrient gelatin on cover glasses. Spores of Monilia jructi- gena, Botrytis vulgaris, Sterigmatocystis nigra, Mucor stolonijer, and Monilia sitophila were used; in some instances they were evenly distributed in the gelatin, in other instances the gelatin was inocu- lated by being touched with the needle at several points varying in distance from the crystal. In no case was there any distinct turning toward or away from the diffusion center. The effect of hyphae upon the direction of growth—CLARK (4) explains his results by supposing that the fungus secretes some substance to which the growing hyphae are negatively chemotropic. While this hypothesis would very well explain his results, he seems not to have made it the subject of experimental study. It may be reasonably assumed that if a fungus is negatively chemo- tropic to its own secretion, the stimulus to turn away from an area containing the fungus would, in early stages of growth, be in some degree proportional to the amount of mycelium in that area. To determine whether the amount of mycelium has an effect, inoculations were made with differing numbers of spores; the growth was from non-nutrient gelatin and gelatin containing M/4 solution of dextrose, and was toward similar as well as different media as indicated in Table VI, where the results are given. The direction of growth is from the first-named medium to the second. Columns 98 BOTANICAL GAZETTE [FEBRUARY I give the average number of spores per hole, and columns II give the percentages of turning toward the holes. TABLE VI S x Mucor uatoorssis| MO Bomaceens-| “yea |: eee CoNnTRASTED CULTURES IN GELATIN NIGRA ee ee GENA LONIFER 2 : : AVE] S4. [15 (Ol 24 | O.8) 27 || 5.0) 207) Essig Non-nutrient : M/4 dextrose..... re hoe 26 | ae | we | er [5 gh ee eae : $.5] 24.| 4.2| 25 | 8.3] 29 | 4.8] 26 | 5.0) 25 M/4 dextrose : M/a dextrose..... ae | ae | 2g \-au.} 2 | 47. | a | 96 | oe M/4 dextrose : non-nutrient...... 8.0] 25 | 7.7| 26 | 6.7) 28 | 6.0] 28 | 6.3) 31 BY 35 | 7-1 36 33 37 | 1 ae xt ; . e ; 3-01 33° | 5:0) 30 oO 3h | 4: 22 I Non-nutrient : non-nutrient...... Bel got c6 | ad | x8 | aot ep | oe] ee Soe 3.01 30:|. 464] 27-|-6.3| 20 |-7.0|-28-| 27.729 M/ 4 glycerin : M/4 dextrose..... ag ely oe er ers re ee 3% pf i Kine: Ss 0rad t-§. 3) 28%) 6.3) 28° | 5.0) 27) 83 Non-nutrient : 0.01% CuSO,.... sod an bas ae|ar | ga} 2) oe ee There is indication that the number of hyphae in a given area and the amount of turning from that area are correlated. It may be said of this, as of succeeding tests, that it is at best only relative. It is manifestly impossible to eliminate growing hyphae from the experiments, and their effect is the very factor to be tested. A test was made by comparing preparations in which only one layer of gelatin was inoculated, with preparations in which both layers were inoculated. Two parallel series were prepared: in one the spores in the lower layer were about twice as numerous as in the other series. An examination of Table VII, which gives the results, shows that the percentage of turning toward a layer containing hyphae is less than toward a sterile layer; and in either case the abundance of mycelium in the layer influences directly the turning TABLE VII Montnta STERIGMATOCYSTIS | Mtcor sToLo- NiORA ahies Mucor MUCEDO I II Til I II Itt I I Ii I II Ill I Il ul 6.5 | 0. |: 26:4 6.6) ot 92 beet Ot ee) are oh ge o | 28 taj 0: | 35. .45 | @y go | tA} oO 1 ae 8 4540) oe) ee te 4-5} 7 | 9615.01) 8 | os 14.51: > 4 36-1 47 | 45-1} 98 @ | 10 4 me 9 G4 oa) 19 8 ae be tS 24-0. 30.135 1.40 fF 29 1906] FULTON—CHEMOTROPISM OF FUNGI 99 of the hyphae from that layer. Column I gives the average number of hyphae per hole in the layer from which the turning takes place; column II, the average number of hyphae per hole in the layer toward which turning takes place; column III, the percentage of turning. To determine whether this negative turning is due largely to chemical changes in the culture medium, beet decoction in which a fungus had grown was tested with that fungus. The sugar-beet decoction was diluted to one-third the strength of the stock solution. When a good amount of mycelium had been formed, the liquid was filtered under sterile conditions at room temperature. The filtrate was replaced in the incubator in the case of those species which had fruited, and was allowed to remain for twenty-four hours in order that any spores that had passed through the filter might germinate; it was then refiltered. In this way a practically sterile medium was obtained. One volume of this decoction was added to one volume of double-strength gelatin, and tests were made with this TABLE VIII * a B GrowTH TOWARD GELA-} GROWTH TOWARD GELA- tin Mape UP witH Tin Mane Up witH Bret DecocTIon IN _| FrEsH Beet DECOCTION. Wuicu Founeus Hap RO Grown I Il I It 1, Doteytls vulgadis. 6.5002 si: 6.2 20 5-3 33 ; 15 3% 15 35 II. Sterigmatocystis nigra............ 10 29 g-o 34 19 27 17 37 IiI. Penicillium MAUCUE ys 23a heen 8.0 23 12 3r 18 22 28 23 IV. Monilia sitophila........0.2+...- 8.0 22 9-0 26 : 23 25 20 33 ¥.. Mucor-stolonifer. >... 2... 0. 33s 19 -32 22 38 VI enricus fabaceus.. < 5c ys neces 49 24 50 32 VIt. Coprinus micaceus.............- 31 24 43 30 VIII. Daedalia quercina............+-- wo 468 35 31 in comparison with control beet decoction in which there had been no fungous growth. The results are shown in Table VIII. Column I gives the average number of hyphae per hole; column II, the percentage of turning toward the holes. . The percentages for all of the fungi in this table, except Mucor 100 BOTANICAL GAZETTE [FEBRUARY stolonifer, have been computed from four distinct series. In this way erratic results have been nullified, and the difference in effect, although not marked, may be regarded as constant. There is certainly a lessened attractive influence in the case of the decoction in which the fungi have grown; this might be due to the mere abstrac- tion from the decoction of nutrient substances, or to the conversion of compounds occurring in the decoction into compounds which are repellent in their effect, or to the secretion of products by the fungus which have a repellent effect. That the first is probably not the case is to be inferred from previous tests, in which these fungi have been found to grow as readily toward distilled water and other non- nutrients as toward nutrients; a mere decrease, therefore, in the amount of available nutriment could hardly have a pronounced effect. We must conclude, therefore, that a medium in which a fungus has grown may become less attractive, or more repellent, to the fungus through the agency of some undetermined substance or substances, which are secreted or otherwise formed by the growing fungus; this reaction would -be a special kind of negative chemo- tropism. Miss FERGUSON (13) found that germinating spores of Agaricus campestris, or bits of older growing mycelia, have a very marked effect in causing the germination of spores of this species; at the same time there seems to be a retarding effect upon the growth of the protruded germ-mycelium. Mycelium that is not growing, masses of ungerminated spores, or growing mycelia of other fungi do not have the same influence over germination. Her observations lead her to suppose that oxygen or carbon dioxid is not the deter- mining factor, but that some secretion is formed which stimulates or makes possible the emission of the germ-tubes. Other observa- tions relative to the influence of germinating spores upon the growth of fungi have been made by KrHLMAN (16) and REINHARDT (28); to these reference has already been made. : Numerous instances have been recorded of the influence by various plant cells upon the direction of movement and of growth of other cells of the same or a different kind, and the general terms cytotaxis and cytotropism have been applied to this peculiar sort of chemical influence. In the cases enumerated by PFEFFER (27, # 1906] FULTON—CHEMOTROPISM OF FUNGI IOI sec. 155), the effects seem to be due either to the excretion of a hypo- thetical specific substance which furnishes the stimulus, or to changes in the relative proportions of oxygen and carbon dioxid through respiratory or photosynthetic activities. Since these phenomena seem to be analogous in a general way to the above-described turning of fungous hyphae, and since the term cytotropism indicates nothing as to the exact nature of the ultimate stimulus-substance, this term might be found a convenient one for designating the present cases. The effect of moisture-—The circumstance that a number of preparations in which the culture medium had become evidently dry, gave large percentages of turning, suggested that the hyphae might react to a hydrotropic stimulus. Layers of agar containing spores of several fungi were spread on cover glasses, and sterile strips of filter paper were placed in contact with the agar and allowed _ to dip into the water of the Van Tieghem cell. The average per- centage of turning toward the strips for those spores within a distance of 0.35™™ from their edges was 4o per cent. for Penicillium glaucum; 50 per cent. for Mucor stolonijer, Monilia sitophila, and Sterigmato- cystis nigra; 55 per cent. for Mucor mucedo; 57 per cent. for Botrytis vulgaris; and 60 per cent. for Monilia jructigena and Sphaeropsis malorum. The percentages given are less than they would be if a smaller area about each strip had been considered; this may be due in part to the circumstance that the spores were rather thickly sown, and the hyphae from those nearest the strip, being numerous, exerted an effective repellent influence on those more distant, causing them to grow away from the strip. Notwithstanding this, the evidence of positive hydrotropism for these fungi was quite conclusive. As a further test, mica plates were cut to fit Van Tieghem cells and were perforated; a drop of non-nutrient gelatin was placed on each and covered with a perforated plate small enough to fit inside the cell; upon this was placed another layer of non-nutrient jelly containing the spores; another perforated plate was added, and a third layer of gelatin, sterile like the first; the lower surface of this was left uncovered. The mica cover was inverted over a cell con- ’ taining water, during the time required for the proper growth of the fungi, evaporation took place from the now uppermost layer to s 102 BOTANICAL GAZETTE [FEBRUARY the surrounding atmosphere, as was apparent from the dry condition of the gelatin around the holes in the uppermost plate; water diffused from the lower layers to supply the deficiency. In this way it came about that the middle layer, which contained the spores, was moister than the uppermost layer, but drier than the lowermost. There was observed a very decided turning of the hyphae toward and through the openings in the third plate, which separated the middle and lowermost layers, while comparatively few grew toward the upper- most layer. The estimated percentages are shown in Table IX. TABLE IX = < 4s = a “| a < n 2 S8/S2/$2)Ee| 88/68) 38| 8 ge DrrEcTIONs PO|Bo|s4|/eea1f5/2#/8e|82)| 28 z Ze|°o iH < blagipYipdl ge S0).09)3n/80) 82) a2 seisei se AB\SE/BR/RB| Ae/Be |" 4/28 | BA & = ae > a a a) ey From middle layer to lowermost layer..| 43 | 70 | 80 | 55 | 68 | 65 | 73 | 63 | 65 From middle layer to uppermost layer..| 20 | 20 | 20 | 30 | 18 | 15 | 18 | 1 18 Experiments were set up in which very firm gelatin (16 per cent.) containing the spores was covered with mica plates having a few perforations. The plates were sealed to the covers by an application of vaseline around their margins.. The covers were then inverted over stender dishes level full of sterile distilled water. In this way the water came in contact with the gelatin only through the perfora- tions, and diffused from these through the gelatin layer. Hyphae of Mucor stolonifer grown under these conditions showed a tendency to grow toward the openings from a distance of 1.5™™, but on coming within 0.5™™ of the openings, the course was changed, and the hyphae circled the openings in lines more or less concentric with their margins. The majority of those nearer the openings than 0.5™™ grew in a radial direction away from them. Ina few instances hyphae grew into the water. Mucor mucedo showed a quite decided turning toward the holes, and about 65 per cent of the hyphae within a radius of three hole-diameters turned through them and grew into the water. With Botrytis vulgaris about 40 per cent., and with Penicillium glaucum 85 per cent. of the hyphae within a correspond- ing area were affected positively. In every case the growth —EEO 1906] FULTON—CHEMOTROPISM OF FUNGI - 103 in the water was in all directions, directly downward, as well as radially in a horizontal plane. The value of the control cultures, which were duplicates in all respects excepting that the dishes were only partly filled with water, was vitiated by the accumulation of condensed moisture in comparatively large drops about the openings in the plates. This caused an unmistakable turning toward the holes, which was not so decided, however, as in the test cultures. It is evident from these results that all of the fungi tested in this regard are, under the conditions of experiment, positively hydro- tropic; but Mucor stolonifer may under certain conditions show a negative hydrotropism. This response to a hydrotropic stimulus probably accounts in large measure for the constant turning toward protected layers from those more exposed, which latter may have become drier through evaporation. A sharp distinction between bedwieeneis and Pacareries on the one hand, or between hydrotropism and osmotropism on the ’ other, cannot in all cases be made, although these phenomena in typical cases are quite distinct. The phenomena here reported are probably due primarily to differences in the moisture content of the layers, and not to water currents, either molar or molecular. For this reason the term hydrotropism has been applied, which is not in agreement, however, with the current view (PFEFFER, 27, p. 592), that in the case of the fungous mycelia heretofore studied, osmo- tropism and rheotropism, but not hydrotropism, have been estab- lished. It is further recognized that chemical rather than other properties of water furnish the effective stimulus, in which event hydrotropism would properly be regarded as a special kind of chemo- tropism. : Aerotropism.—Under the conditions prevailing in some of the experiments above described, there was doubtless an inadequate supply of oxygen, as when a medium poor in oxygen was enclosed between impervious plates. There was then a very decided tendency for the hyphae to turn toward the edges of the plates. The obser- vation of this phenomenon from time to time suggested that the fungi might show an aerotropic sensibility, either as a positive reaction to oxygen, or as a negative reaction to carbon dioxid. In order definitely to test the matter, experiments were arranged 104 BOTANICAL GAZETTE [FEBRUARY in which the growth toward holes in mica plates could be observed when the plates separated normal non-nutrient gelatin from non- nutrient gelatin saturated with carbon dioxid on the one hand, and from normal non-nutrient gelatin on the other. The fungi used were Penicillium glaucum, Sterigmatocystis nigra, Mucor mucedo, Botrytis vulgaris, Monilia fructigena, Monilia sitophila, and Phy- comyces nitens. In no case was the percentage of turning toward the carbon dioxid gelatin greatly different from that toward the _control gelatin. As a further test, a layer of gelatin containing spores was placed below a perforated mica cover for a Van Tieghem cell, and a per- forated mica plate, small enough to fit inside the cell, was placed below the gelatin. A layer of normal gelatin was spread below this last plate, and a layer of carbon dioxid gelatin above the cover. This preparation was sealed to the cell rim, and the whole placed * under a bell jar practically filled with carbon dioxid and kept at a room temperature of 21-24° C. Efforts were made to have the moisture conditions equal within and without the cells; and the exposed gelatin layers, which served very well as indicators, showed no difference in this respect until after the observations on the majority of the preparations had been made, although there was drying of the outer gelatin layer by the time the more slowly growing fungi had reached the proper stage. The same fungi were used in this experiment as in the preceding one, with the addition of Mucor stolonifer. In most instances the turning toward the gelatin con- taining carbon dioxid and exposed to an atmosphere of carbon dioxid, was as great as toward normal gelatin; the growth, however, was less vigorous in the former case. In those preparations in which there was less turning toward the carbon dioxid gelatin, this gelatin had become evidently rather dry. It is to be concluded, therefore, that the observed turning toward the edges of preparations is not due primarily to aerotropic sensi- bility. The experiments also negative the supposition that the observed repellent influence of growing hyphae may be due to the | consumption of oxygen or to the production of carbon dioxid by the fungus, or to both. oO Z rah Al ck & tt RE Fe Te he g the fact that osmotropism is 1906] FULTON—CHEMOTROPISM OF FUNGI 105 intimately associated with chemotropism, and that many of the tests for the latter are in equal measure tests for the former, direct tests were made by growing the fungi in media of higher osmotic pressure and of lower osmotic pressure than the test media, as well as in an isosmotic medium; glycerin, a good nutrient substance, and yet a substance reported by Mryosut to be neutral in its chemotropic effect, was used to give the desired concentration to the culture media. The series failed to show that the concentration of the culture medium has an effect upon the amount of turning. How- ever, no excessively high concentrations of mineral salts were used. Other tropic phenomena.—Under conditions that would favor a manifestation of geotropism and of thigmotropism, there was no indication that these are concerned in determining the direction of growth of the fungi. The effects of light and of heat were in no way tested, but they probably do not enter as factors. Biological significance—The conclusions reached in these studies may be found to have a somewhat important bearing upon the biological problem of infection by parasitic fungi. In the absence of any experimental investigation, nothing definite can now be said. It would seem, however, that the drying of dew and other surface moisture in which spores had gerimnated, might be a condition favoring the hydrotropic turning of the germ-tubes toward the stomates, especially if the cells within are over-distended with water, which has frequently been observed to be a condition favorable for infection; if the germ-tubes are numerous in the vicinity of a stomate, the repellent influence of these upon one another would Cause some to seek the unoccupied region within the stomate. At all events the phenomenon of the entrance of germ-tubes, whether by way of the stoma or through the cuticle, is a complex one, of which many factors remain undetermined. That mere entrance is probably not due to specific peculiarities, either of host or parasite, is evidenced by the recent work of Miss Grsson which has been mentioned by MarsHatt Warp (33). Miss Grsson found that Spores of various members of the Uredineae sent their germ-tubes readily into the stomates of plants widely different from their hosts, and which they were unable to infect. MARSHALL WARD met with numerous instances of the same phenomenon. This would indicate 106 BOTANICAL GAZETTE [FEBRUARY that the entrance of a hypha into a stoma is merely a preliminary act, distinct from infection proper, and controlled. by general con- ditions, while the fate of the hypha after its entrance is determined by complex reactions between parasite and host, which are largely specific in their nature. In the light, then, of known facts, no simple explanation, such as the theory of chemotropism due to the presence of specific chemical compounds, is adequate. Chemotropism may possibly be one factor in the complex phenomenon, but it is certainly not the predominant factor. CONCLUSIONS. Various tests upon a number of fungi failed to indicate the exist- ence of any definite chemotropic sensibility to nutrient substances or other chemical compounds in solution. If positive chemotropism exists, it is less prominent than other tropic phenomena involved, and was obscured by them. Those substances which furnished nutriment to the fungi caused a decided growth, often with thickening of the hyphae and an increased branching; but they did not cause a more marked turning of the hyphae toward the diffusion centers than did non-nutrient and toxic substances. All of the fungi tested show a tendency to turn from a region in which hyphae of the same kind are growing toward one destitute of hyphae, or in which the hyphae are less abundant. The turning toward a medium in which mycelium has grown, but from which the mycelium has been removed, is less marked than that toward a medium in which no mycelium has grown. This may be regarded as a negative reaction to stimuli from chemical substances, which owe their origin in some way to the growing fungus. Various fungi show a positive hydrotropism; but an over- abundance of moisture may cause a negative reaction in certain fungi. The changing of the direction of growth of fungous hyphae is a complex phenomenon in which at least two factors, cytotropism and hydrotropism, are concerned. Since the complete elimination of neither of these factors is possible, all tests must be relative, and to that extent unsatisfactory. It would seem that the reactions of mycelium to various stimuli 1906] FULTON—CHEMOTROPISM OF FUNGI 107 are not necessarily the same as the reactions, under similar conditions, of sporangiophores, gametophores, and other specialized parts. The writer wishes to acknowledge his indebtedness to Dr. B. M. Duccar for his very helpful suggestion and cirticism, and to Dr. Witt1am TRELEASE for the opportunity to consult the library of the Missouri Botanical Gardens. BoranicaL LABORATORY, University of Missouri. LITERATURE CITED. - BREFELD, O., Botanische Untersuchungen iiber Hefenpilze. 1883. Buscen, M., Ueber einige Eigenschaften der Keimlinge parasitischer Pilze. Bot. Zeit. 58: 59. 1893. CrarK, J. F., On the toxic effect of deleterious agents on the germination and developmen of certain filamentous fungi. Bot. GazETTE 28: 293. Now " , On the toxic properties of some copper compounds with special reference to Bordeaux mixture. Bor. GAZETTE 33:45. 1902. De Bary, A., Beitr. zur Morph. u. Phys. d. Pilze. 4:85. 1881. , Comp. Morph. and Biol. of Fungi, etc., Engl. Ed. 1887. 366. 1884. Dietz, S., Beitrige zur Kenntniss der Substratrichtung der Pflanzen. eee Unters. Bot. Inst. Tiib. 2:4 88. Duccar, B. M., Geceeipcieg a spores. Bor. GAZETTE 31:44. 1901. , The culiivaition of mushrooms. U. S. Dept. Agric., Farm. Bull. no. ee I 10. Errera, L., Die grosse ee ee bei den Fruchttragern von Piscine Bot. Zeit. 42 , On the cause of kee action at a distance. Ann. of Bot. 6: cas ; 12. Fatcx, R., Die Bedingungen und die Bedeutung ics a bei Sporodinia grandis. Cohn’s Beit. Biol. Pflan. 8:213. 190 13. FERGuson, Marcaret C., A preliminary study of ‘he BEST i of the spores of Agaricus campestris and other basidiomycetous fungi. Bur. Pl. Indus. U. S. Dept. of Agric. Bull. 16:26. 1902. 14. HormelsTer, Die Pflanzenzelle 286. 1867. 15. Jonsson, B., Der richtende Einfluss ge ene auf wachsende Pflanzen und Pflanzetheile. Ber. Deutsch. Gesells. 1:512. 1883. 16. Kratmann, O., Zur gency Sones — A ae Acta Soc. Sc. Fennicae 13:12. 17. Kress, G., Zur Pipidoge der Fortpflanzung einiger Pilze. I. Sporodinia grandis. Falah. Wiss. Bot. 32:55. 1898. BOTANICAL GAZETTE [FEBRUARY Kwny, L., Ueber den Einfluss ausserer Krafte auf Anlegung von Sprossungen thalléger Gebilde. Sitz. Bot. Ver. Brandenburg 23:8. 1881. . Masser, G., On the origin of parasitism in — Abstract in Proc. Roy. Soc. Cpt B73:118. 190 de 9 . Mryosut, M., Ueber elon copiasins der Pilze. Bot. Zeit. 52:1. 1894. , Die Darchboheuhe von Membranen durch Pilzfaiden. Jahrb. Wiss. Bot. 53:28. 1895. - Moutscu, H., Untersuchungen iiber den Hydrotropismus. Sitz. Akad. Wiss. Wien 88: 847. 1883. - Moore, G. T., Methods for growing pure cultures of algae. Jour. App. Micr. 6:2312. 1903. . NorpDHAUSEN, M., Beitrige zur Biologie parasitirer Pilze. Jahrb. Wiss. Bot. 33:1. 1898. . Prerrer, W., Locomotorische Richtungsbewegungen durch chemische Reize. ied: Deutsch. Bot. Gesells. 1:532. 1883. eber chemotactische Bewegungen von Bakterien. Flagellaten, und Vorvoonead: nters. Bot. Inst. Tiib. 2:582. , Pflanzenphysiologie 2. 1904. Rania: M. O., Das Wachsthum der Pilzhyphen. Jahrb. Wiss. Bot. 23:47. 289 STRANGE, B., Ueber chemotactische Reizbewegungen. Bot. Zeit. 48: 140. "1890 . SrevYER, Reizkriimmungen bei Phycomyces nitens. Leip. Diss. 1901. . SWINGLE, W. T., Bordeaux mixture, its chemistry, physical properties, and toxic effects on = and algae. site Veg. Phys. and Path. U. S. Dept. Agric. Bull. 9: . Warp, H. diesec A lily disease. Ann. of Bot. 2:319. 1888. , Recent researches on the parasitism of fungi. Ann. of Bot. 19:1. 1995. Worontn, M., Ueber die Sklerotienkrankheit viet Vaccinieen-Beeren. Mém. Acad. St. Pétersbourg VII. 36:1 (no. 6). WortMany, J., Ein Beitrag zur Biologie der wi oe Bot. Zeit. 39: 368. 1881. , Zur Kenntniss der Reizbewegungen. Bot. Zeit. 45:812. 1887. Ba Sea ee ee ee ee BED ang a dr gS ET at Sel iar SE THE EMBRYOLOGY AND DEVELOPMENT OF RICCIA LUTESCENS AND RICCIA CRYSTALLINA.' CHARLES E. LEwis. (WITH PLATES V-IX) In June 1903, while collecting liverworts in the vicinity of Ithaca, N. Y., an abundance of material of Riccia lutescens was found growing around the edges of dried-up ponds. In some cases the plants formed beautiful rosettes, but usually they grew in irregular clusters, often being so closely crowded together as to cover the ground for several square centimeters. The individual plants vary greatly in shape and size. The younger light green plants consist of a narrow, thin, ribbon-shaped thallus which has a longitudinal median groove. In the older plants the fore part of the thallus is thickened, very large air cavities being formed: The thallus is attached to the soil by numerous rhizoids from the older part, the apical end being free. On the under side are numerous colorless lamellae. As the fruiting plant is unknown there is doubt as to the rela- tionship of this species, authorities differing widely as to its status. LINDBERG (21) claimed that it was merely a sterile terrestrial form of Ricciocarpus natans. UNpdERWwooD (30) says of it: “approaches certain terrestrial forms of Ricciocarpus natans, and possibly derived from that species, but better kept distinct.”” SrEPHANI (28) states that it is probably not a Riccia but a sterile marchantiaceous hepatic. For the purpose of determining the true relationship of the species, Professor ATKINSON suggested the desirability of following the development of the plant through the summer and autumn, and of securing fruiting specimens if possible. He had found young antheridia in plants collected several years before, but had not traced the development further. It also seemed desirable to study the embryology and cytology of the plant if material could be obtained, because comparatively little has been done on these phases of the life history of Riccia. * Contribution no. 106, from the Department of Botany, Cornell University. 109] [Botanical Gazette, vol. 4: 110 BOTANICAL GAZETTE [FEBRUARY BIsCHOFF (2) investigated a number of species and settled beyond a doubt the function of the sexual organs. His work was followed the next year by LINDENBERG’S monograph (20) which added little that was new. - The study of the development of Riccia really begins with Hor- MEISTER (15), who gave an account of the development of the thallus, sexual organs, and fruit of Riccia glauca. Kwy (19) made a careful study of the apical cells and the method of growth of the thallus. He did not secure plants developing from spores but used delicate thalli which had grown crowded together and did not bear sexual organs in the younger parts, so that the regular order of cells was not disturbed. He discovered the origin and manner of growth of the ventral scales and described the develop- ment of the sexual organs. Although HormeEIsTER believed that young antheridia and archegonia could not be distinguished, Kny points out that they are distinct after the first walls are formed. LEITGEB (22) gives a complete account of the method of growth of the thallus in the Ricciaceae. His study of the sexual organs and fruit was in many cases incomplete on account of insufficient material. BIOLOGY OF RICCIA LUTESCENS. The account of the biology of Riccia lutescens given here is based on field observations extending through two years, together with experiments and observations upon plants kept growing under favorable conditions in the greenhouse and laboratory. The first observations were made late in June. At that time the plants were growing-upon the mud around the edges of ponds. Some of the thalli were very small and delicate, appearing merely as green specks on the mud, while others, which seemed to be older, had the ribbon-shaped form and thickened apical end already described (figs. 1-3). Material was collected and examined from time to time during the summer and autumn, with the expectation of finding plants bearing the sexual organs, because the statement is usually made that the species of Riccia fruit in summer and autumn when growing on the soil. The plants continued to grow well vegetatively through- out the summer, when they were in such a location that they were 1906] LEWIS—DEVELOPMENT OF RICCIA T1y supplied with sufficient moisture. In some cases the mud became so dry and hard that the plants were killed, but whenever they were sheltered by a stone or other object, or were growing on the sides of holes, such as cattle tracks, they grew well. In October all but the youngest and most crowded plants showed the typical Riccia lutescens form. At this time young antheridia were found. Material was now collected and fixed from time to time for the purpose of studying the development of the sexual organs. In very few cases were archegonia found in plants collected in autumn. A few young stages were found in plants collected late in November, at which time the older antheridia were almost mature. No further development took place out of doors until spring, because the plants became covered with snow, or with water by the filling up of the ponds, and remained so until April. A quantity of the plants were kept growing on the soil in the green- house through the winter, and developed mature sexual organs long before spring. Plants taken from under water in March, just as the ice was going out of the ponds, showed exactly the same form as in November, and little or no further development had taken place. So it seems that the development depends to some extent on temperature, and might be expected to vary with different con- ditions of climate. A warm winter, in which some growth might take place, would in all probability hasten the development of the sexual organs. Another point of interest is that the submerged plants did not seem to have been injured. A quantity of material still attached to the soil was taken from under water late in March, and was kept growing in shallow pans in the laboratory so that it could be kept supplied with a sufficient quantity of water for growth but not enough to flood the plants. This was done in order to determine whether the plants would con- tinue the development of sexual organs and fruit in the same way when supplied with a limited amount of water and growing on the soil, as when supplied with a large amount of water which would tend to cause them to break loose and float. It was found that the’ Plants growing on the soil did produce fruit abundantly and at the same time as those growing under natural conditions. The archegonia begin to develop in April in the same thalli which have hz BOTANICAL GAZETTE [FEBRUARY produced antheridia, and all stages are found by May 1. About this time fertilization takes place, and by May 25 all stages of sporo- phyte are found. The arrangement of the sexual organs in the thallus is shown by figs. 7-11. The vegetative growth is very rapid during April and May, the thallus becoming broad and branched by the increase in the number of growing points. The narrow older part by which the thallus is attached decays, and the younger part bearing the sexual organs and sporophytes is set free and floats upon the water. When the plants are supplied with a large amount of water changes take place in the lamellae. They grow to great size and become purple. In the floating thallus decay of the older part continues; the part bearing the antheridia first disappears, then the part bearing the sporophytes, and finally the growing points may be separated, one thallus thus giving rise to several new individuals. In most cases observed the decay of the older parts in floating plants did not advance so far. The plants were carried up around the edge of the pond by the waves, and as the water went down were left stranded upon the mud. When the thalli settle down upon the mud, the large ventral plates wither, and rhizoids are put forth which in a few days attach the thallus to the soil. Growth now continues at the growing points, so that new branches are produced which form rosettes. When the thallus is injured at this time new plants are imme- diately produced from the cells of the apical region. This was first observed in plants injured by being covered with mud, in which case slender delicate outgrowths were produced (fig. 4). Other plants injured by snails soon developed long slender plants (fig. 36)- Thalli were cut into pieces to determine whether other cells would show the same plasticity, but new plants were produced only from cells near the growing point. V6cHTING (33) found in Lunularia that regeneration takes place from cells in various parts of the thallus, but this does not seem to be true of Riccia natans under the conditions in which I have studied it. Large numbers of the plants which were left upon the mud when the water went down were injured by cattle coming down to the ponds to drink. Later in the season the cattle tracks were lined with young, green, ribbon-shaped plants 1906] - LEWIS—DEVELOPMENT OF RICCIA It3 which were outgrowths from the growing points of the older plants. The cattle tracks serve a good purpose, as the young delicate plants are shaded and protected to some extent during the dry season. Two forms of the thallus are produced by the different methods of propagation. In the one case the thallus after it becomes attached to the soil continues its growth, branches and forms a rosette, while in the other case the thallus is injured, and very delicate forms are produced. When large numbers of the floating thalli are deposited near together and are then injured, we find the irregular clusters of plants which have been described in the first paragraph. The thallus of this plant during the floating period bears such a striking resemblance to Ricciocarpus natans that one is led to the conclusion that Riccia lutescens is only a ground form of Ricciocar pus natans. Since the beginning of this study and after it was well under way, a paper was published by GARBER (11) which dealt. ‘with the life history of Ricciocarpus natans. Several points in the biology of the plant as given by GARBER differ from those found to obtain at Ithaca, and since the structure of the thallus as well as the embryology is conclusive proof that the two forms are the Same species, it seems proper to call attention to these differences and then to give briefly the embryology before taking up the other phases of the study. The greatest difference in our observations lies in the relation of the supply of water to sexual reproduction. GARBER states that Ricciocarpus natans as it grows at Chicago spends its entire life, from the germination of the spore to the production of spores, in the floating state, and that the occasional fruiting plants found upon the soil in summer are plants in which the sexual organs devel- oped and the sporophytes began their development while the plants were floating. He observed no case in which sexual organs were produced on plants growing upon the soil and states that Ricciocar pus natans has not yet acquired the power to reproduce sexually when growing upon the soil. The sexual organs develop in April. The plants at Ithaca, however, spend the greater part of their life upon the soil and only float upon the water for a few weeks at the fruiting period. The sexual organs begin to develop in autumn while the plants are on the soil and plants kept on the soil and sup- 114 BOTANICAL GAZETTE [FEBRUARY plied with a limited amount of water developed fruit. At the time when the antheridia begin to develop the gametophyte is under favorable conditions for vegetative growth, but is not supplied with an abundance of water. The soil is moist and the conditions are such as would favor the growth of a terrestrial form like Marchantia. The plants seem especially adapted to spend the winter submerged and do not perish under such conditions. The fore part of the thallus contains very large air cavities and thus the tissue is aerated. It is well known that certain higher plants which grow in wet situa- tions have large air spaces in the tissue, and GANONG (12) calls attention to the fact that those marsh plants which are submerged for a portion of the year are able to survive on account of their capacity for air storage. About May 1 the older part of the thallus, which is narrow and thin, has decayed, and the younger parts, bearing the sexual organs, is set free and floats. GARBER points out that when land forms are placed upon the water only a small portion - of the apical end remains above the surface, while the older part of the thallus extends into the water and decays. This is true of plants taken from the soil in summer, but in the spring when the free part of the thallus is thick and contains large air cavities, it floats readily. The length of the floating period depends of course upon the conditions of the pond. In some cases the plants may very soon be carried up around the edge of the pond and deposited on the mud, but floating forms are usually found until the ponds are almost dry. In the case of ponds which do not become dry in summer, both forms would be found. The floating period affords an excellent means for distribution. When the plants grow upon the soil and are not protected during the winter by a covering of snow or water, they are usually killed by freezing, but in some cases plants which were brown and seemed to be dead produced new thalli from the growing point. The young delicate thalli are well adapted to tide over the dry season, because they can live with a less supply of water than would be needed by the older plants. RELATIONSHIP OF THE SPECIES. The form usually described as Riccia lutescens should be regarded as a ground form of Ricciocarpus natans. Both in the field and in 1906] LEWIS—DEVELOPMENT OF RICCIA ars cultures in the laboratory the forms have been observed showing the transition. There can be no doubt that the plant which I have described is the true Ricciocarpus natans, and the description of the ground form as a distinct species came about naturally from the conditions of its growth. In such ponds as have been observed here, the water is high in April and May, so that the floating plants are carried up around the edge and left on the soil. In June or July the water has entirely disappeared from the pond and the only plants found are the slender ribbon-shaped ones which have developed from the floating form. In my first summer’s collecting, when the ponds were dry by the last of June, I saw not a vestige of the old Ricciocarpus natans, and felt sure that the plants collected were Riccia lutescens. It seems possible that the plant was first described as a distinct species from material collected under similar conditions, because it is said to occur in dried up ponds and ditches. If in summer and autumn some water were present, so that some of the typical Ricciocarpus natans would be found floating, the origin of the ground form might readily be seen, but in such a case there might be failure to associate the ground form with Riccia lutescens. Only by following the development and observing the transition of one form into the other under different conditions of growth can the true relationship be determined. My observations have con- vinced me that Riccia lutescens is only a ground form of Ricciocar pus natans and should not be regarded as a distinct species.’ The plant now known as Ricciocarpus natans was formerly ‘regarded as a Riccia. In the structure of the thallus Ricciocarpus is more complicated than the species of Riccia. The most important taxonomic characters, however, have been the arrangement of the sexual organs and structure of the sporophyte. Hooker first found fruiting plants in dry specimens sent to him by Torrey from New York in 1824. BrscHorr found fruiting plants in the autumn of 1829 near Heidelberg, and describes anthe- ridial plants, but his figure of the antheridium is not very convincing, ? Having determined the ground form as Riccia lutescens, specimens were sent to Professor A. W. EVANS in October 1904. He considered that we were right in refer- ring the plants to that species, but stated the views of different authorities in regard to the status of the species. 116 . BOTANICAL GAZETTE [FEBRUARY as it looks more like the mass of tissue which projects up as a ridge into the median groove, the cells being quite too large for those of an antheridium. Although Hooker considered that the plant should remain in the genus Riccia, Corpa placed it in a new genus, Ricciocarpus, on the basis of Hooxker’s description and figures which were taken from dried material. Corpa’s figures are copies of HOOKER’S. BiscHorrF held that there was no real basis for the change, as the mature sporophyte does not differ from that of other Riccias, the separation being based on the mistaken notion that the capsule walls disappear entirely at maturity, and that the genus Riccia should not be divided on account of differences in the thallus brought about by the different conditions under which the plant grows, since the method of fruiting is the same in all the species. LEITGEB regarded Ricciocarpus as a distinct genus, on account of the more complex structure of the thallus and the grouping of the sexual organs. He thought that the antheridia were collected into groups similar to those in the Marchantiaceae, but GARBER’S results and my own show that LerrcEB was not correct, and that the antheridia actually form only one group. The archegonia are also arranged in a definite part of the plant in one group. The question now arises whether this is a more advanced con- dition of development than is found in species of Riccia. In the lower species of Riccia, the sexual organs are said to be indiscrimi- nately scattered over the surface of the thallus, while in Riccia fluitans a regular alternation of single antheridia and archegonia is described.” CAMPBELL, in discussing the arrangement of sex organs in Riccia, says that in the two forms which he studied, Riccia hirta and Riccia glauca, he found as a rule that several of one sort or the other would be formed in succession. I have observed the same in Riccia crystal- lina, although the older sporophytes appear scattered in the thallus. LINDENBERG described the fruit of Riccia crystallina as scattered, but the antheridia are described and figured as being in a group along the middle part of the thallus. He described and figured the fruit in Riccia glauca as being sometimes in rows and sometimes scattered. Most of the figures show them in more or less perfect rows along the longitudinal axis. 1906] | LEWIS—DEVELOPMENT OF RICCIA 1i7 In Riccia minima, LINDENBERG (20, p. 429) describes and in pl. 20 figures the antheridia as arranged in two rows, one on each side of a median groove. In Riccia bulbosa the antheridia are along the median groove for its entire length, sometimes in pairs and sometimes far apart. Riccia Bischoffiit has the antheridia in two or three rows in the thallus. It seems highly probable that a careful study of a large number of species of Riccia by modern methods would show that in many of them there are produced groups of antheridia and archegonia in distinct parts of the thallus. Since the characters upon which the genus Ricciocarpus has _ been based, with the single exception of the structure of the thallus, have been found wanting, it seems to me that there is not sufficient reason for retaining the genus. The thallus varies in form according to the supply of water, and when growing on the soil has been called a species of Riccia. Many plants assume quite different forms when growing under different conditions, but the different forms are not regarded as species. | We should then write: RICcIA NATANS L. Syst. Veget. 956. 1774.—Bischoff, Nova Acta Acad. Caes. Leop. Carol. 17: 2. 1835.—Lindenberg, Nova Acta Acad. Caes. Leop. Carol. 18: ease —Sullivant, Gray’s Manual 2ed. 1856. Ricciocarpus natans Corda, Opiz Naturalischentausch. 1829.—Leitgeb, Die Riccien, Unters. Lebermoose 4:1879.—Lindberg, Revue Bryol. 9:82. 1882. (Includes Riccia natans L. and Riccia lutescens Schw.)—Schiffner, Engler and Prantl. 1893.—Campbell, Mosses and Ferns. 1895.—Underwood, Systematic Botany of oe America. Hepaticae. 1895.—Garber, Bot. GAZETTE 37:101- 177. pls. 9-10. 1904. Riccia dso Schw. Specimen Fl. Amer. Sept. Crypt. 26. 1821.—Linden- berg, Nova Acta Acad. Caes. Leop. Carol. 18: pl. 26. 1836.—Sullivant, Mem. Amer. Acad. II. 4: pl. 4. 1849.—Sullivant, 2d ed. Gray’s Manual 684. 1856. —Underwood, Systematic Botany of North America, Hepaticae. 1895. Riccia velutina Hooker (in part) Ic. Pl. pl. 149: founded on sterlile thalli of Riccia lutescens and fertile thalli of Riccia crystalline, —s to Sullivant, Gray’s Manual, 1856. EMBRYOLOGY. Material for study was collected during the autumn and spring, and fixed very satisfactorily in 1 per cent. chromacetic acid or in chromosmacetic. 118 BOTANICAL GAZETTE [FEBRUARY The large air cavities prevent the penetration of the fixing fluid, to overcome which the pieces were submerged by means of cotton plugs. After dehydration the material was passed through chloroform — into paraffin. Sections were stained with the triple stain of Flem- ming or with Heidenhain’s iron-alum haematoxylin. SEXUAL ORGANS. Young antheridia were found in October. They begin to develop while the plants are young and growing on soil not supplied with a large quantity of water, although the conditions for vegetative growth are good. At this time the thallus is ribbon-shaped, with a thick- ened apical end and a longitudinal median groove, the thallus in cross-section having about the shape of an inverted Y with a ridge of tissue between the arms (fig. 9). Very few plants are found which do not produce antheridia. The archegonia develop later in the same thallus. At first there seemed to be in this a distinction between Riccia lutescens and Ricciocarpus natans, because Ricctocar- pus natans has been described by SCHIFFNER, LEITGEB, and CAMpP- BELL as being strictly dioecious, but the work of GARBER shows conclusively that it is monoecious. The earlier observers state that Ricciocarpus fruits in autumn, so it seems probable that their material was collected after the older portion of the plant had decayed, leaving only the portion bearing sporophytes. The antheridia are produced in acropetal succession in three to five rows (figs. 10, IT). The antheridium develops as has been described for other species of Riccia. A superficial cell on the floor of the dorsal furrow just back of an apical cell protrudes above the surface and is cut off by a horizontal wall. The outer cell increases in size, and is divided by three or four cross walls, then a longitudinal wall is formed divid- ing the young antheridium into two equal parts: this is followed by a second longitudinal wall perpendicular to the first. Then periclinal walls are formed which cut off the single layer of cells which form the wall of the. antheridium. The cells in the center now undergo repeated divisions until a very large number of cells is formed. Each of these cells is almost. cubical in form and in Riccia has been described as producing a single spermatozoid, 1906] LEWIS—DEVELOPMENT OF RICCIA 119 Kwy (19). The mature antheridium’ is a short stalked oval body with a conical apex. As the antheridium develops, the vegetative tissue grows up and surrounds it so that it is enclosed in a cavity which opens into the dorsal furrow. This cavity is formed in the same way as the air spaces of the thallus. The apex of the antheridium is a little below the floor of the dorsal furrow and the sperms escape through the neck formed by the surrounding tissue. Although the antheridia begin to develop in autumn, they are not mature until the following spring, because the growth is checked by the cold. Plants kept in a warm place produced mature antheridia during the winter. A series of archegonia is developed which is a continuation of the series of antheridia (fig. 7). The archegonium is at first super- ficial on the floor of the dorsal furrow. Later it becomes enclosed in a cavity by the upward growth of the vegetative tissue as in the case of the antheridium except that the neck of the mature arche- gonium protrudes above the bottom of the furrow. The origin of the archegonia side by side at the bottom of the dorsal groove is Shown in figure 9. In this way three to five rows are formed and later a large number of sporophytes are found in each thallus. The archegonium develops in general as has been described by JANCZEWSKI (17). My observations confirm the account given by GARBER for Ricciocarpus natans, as a comparison of the figures will show, so it is unnecessary to describe the development here. About the time when the archegonia are mature, cross-sections of the thallus show numerous, delicate, almost hyaline, club-shaped hairs extending up from the floor of the median groove. Each hair consists of a stalk of two or three short, narrow cells with a much larger cell at the free end. These hairs bear a striking resem- blance to paraphyses(jig. 78). LEITGEB (22, p.31) describes ‘‘papillae’”’ which grow up from the bottom of the groove and regards it as highly probable that it was the dried remnants of these which LINDEN- BERG observed when he wrote: “Sporangium vor aussen mit kleinen unregelmissigen braunen Schuppen bedeckt ist, die Fragmente einer zersprengten friiheren Hiille zu sein scheinen.”” As the hairs become older they become brown and break down so that they 120 BOTANICAL GAZETTE [FEBRUARY would give much the appearance described by LINDENBERG. We know now that the sporophyte has no Hiille or sheath. SPOROPHYTE. The development of the sporophyte agrees with the account given by CAMPBELL (3) for Riccia and by GARBER for Ricciocarpus natans. The first division is usually transverse but may be oblique (jig. 21). The next wall may be perpendicular to the first so as to form a quad- rant (fig. 22), or parallel to it, producing a row of cells. Divisions take place in all directions after this until an almost spherical mass containing several cells is formed. Then the amphithecium becomes distinct. as a single layer of cells enclosing the spore producing cells. The growth takes place rapidly but the divisions of the cells are not simultaneous, usually only a few dividing cells being found in a sporophyte. 7 The sporophyte continues its growth until a solid mass of three to four hundred cells is produced. Then the calyptra and amphi- thecium expand and the spore mother-cells becoming free separate from one another and become rounded. From the surrounding cells, which are richly stored with food, there is secreted a large amount of nutritive material which fills the space around the mother- cells, giving them favorable conditions for growth (figs. 25, 206). The spore mother-cells increase rapidly in size and again fill the cavity. That part of the nutritive material not absorbed by the spore mother-cells is pressed into thin plates between them. This material takes .a deep blue stain with gentian violet. A fuller dis- cussion of the spore mother-cells and of their division to produce the spores will be given in another place. Before the spores are mature the inner layer of the calyptra collapses. The amphithecium is distinguishable until the spores are almost mature. The outer layer persists but the cells are usually shrunken. The contents of these cells is no doubt used up to supply the growing spores with nourishment. All of the spore mother- cells produce spores, there being no sterile tissue in the form of elaters. In discussing the simple form of sporophyte of Riccia, GARBER considers that the absence of sterile tissue is to be associated with the habit of the plants; since there is not much chance for the attach- ment of an independent sporophyte, there is no sterile tissue in 1906] LEWIS—DEVELOPMENT OF RICCIA the form of a foot. When we consider the fact that some other Hepaticae which have the foot well developed grow on very wet -soil and require as much moisture for their development as do some of the species of Riccia, this theory does not seem entirely convincing. The sporophyte develops during May and June. A given sporo- phyte requires about three weeks for its development. SPOROGENESIS. Usually the most favorable cells for the study of cytological details are the spore mother-cells. Their large size, abundant contents and active growth at the time when divisions are taking place, permit good results in fixation. Riccia crystallina has furnished the most satisfactory material. In July, 1903, an abundance of fruiting Riccia crystallina was found growing on the mud on the bottom of a dried up pond not far from the ponds where the form known as Riccia lutescens was growing. This species had never been collected in this region before. Having so determined the plant, I referred specimens to Prof. A. W. Evans who confirmed my determination. He says: “Apparently this species represents an addition to the hepatic flora of New York. I find no mention of it in local lists of New York plants and there are no specimens of it from your state in my herbarium.” » | These plants had been growing under favorable conditions, as the pond had not contained much water at any time during the spring. The thalli formed rosettes growing so close together as almost to cover the ground. The number of fruiting plants was very striking, as it seemed impossible to find a single sterile plant. All Stages in the development of the sporogonium and spores were easily obtained, and some stages in the development of the sexual organs, but changes were taking place very rapidly and the younger Stages were of comparatively rare occurrence. The development of the sexual organs and fruit agrees with that of other species of Riccia. Each thallus produces several sporophytes which are easily recognized when mature as small black spherical bodies imbedded In the tissue. These plants continued.to develop and produce sporophytes for only a short time after they were discovered. The month of July 522 BOTANICAL GAZETTE [FEBRUARY was the most favorable time for the collection of material showing karyokinesis in the spore mother-cells. During August, the spores became mature and the thalli broke down. No good specimens could be collected after August 25. This differs from what has been observed for some other species of Riccia, which are described as withstanding long periods of drought, the thalli continuing their growth again when supplied with moisture. (CAMPBELL, 3.) During the following winter this pond became filled with water and did not become dry until late in the summer, so that only a few plants were found as compared with the large number of the pre- ceding year. This made a difference in the time of fruiting. In September the sporophytes were in about the same stage of develop- ment as in July of the preceding year. This may explain why different authors give different seasons for the fruiting of Riccia. It seems that conditions of temperature and water supply exert such an influence that in the same species and locality the time may vary considerably from year to year. In general, I think it may be said that good conditions for vegetative growth will hasten rather than retard the fruiting of Riccia. The thallus of Riccia crystallina is small and thin; its surface presents a series of wide depressions separated by thin lamellae; and there are no ventral scales. The fixing fluid easily penetrates and the spore mother-cells are usually well fixed. The development of the spore mother-cells agrees with the account given for Riccia natans, but there is not such a large number produced in a sporogonium. When the spore mother-cells come to lie loosely in the sporogonium, they are surrounded by nutritive material. The mature spore mother-cells are then generally spherical, but they may be elliptical or so angular by crowding as to look like a tissue. The contents of the spore mother-cells of Riccia has been described as granular by CAMPBELL (3) but the structure of the cytoplasm in Riccia crystallina is a fine reticulum with the granules occurring usually at points of intersection of the fine threads of the network. The older spore mother-cells as well as the mature spores contain considerable oil. In the nucleus of the spore mother-cell the chromatin is scanty and is irregularly scattered on a fine linin network. No nucleolus 1906] LEWIS—DEVELOPMENT OF RICCIA 123 has ever been observed (fig. 34). When the nucleus is preparing for division, the chromatin leaves the linin network and collects into several bodies which soon move together to form one irregular mass. I regard this as the synapsis stage. Such bodies of chromatin have been found often and in cells which seemed to be well fixed so it Seems to represent a stage in the preparation for division and not to be a result of shrinkage as has been suggested by certain authors for other plant cells in which the same condition has been ‘observed. The body of chromatin occupies a position at one side of the nucleus, and the rather large nuclear cavity appears hyaline. There can be little doubt that the body described by CAMPBELL (3) as a nucleolus is really the’entire mass of chromatin in the synapsis stage. From this mass of chromatin a short thread develops which later Segments to produce the chromosomes (jig. 35). The small amount of chromatin present here makes the details very difficult to deter- mine. The four chromosomes, easily counted here as well as in the nuclear plate and on the way to the poles, are very small and appear almost spherical when on the spindle although they are short, thick, curved rods. The development of the spindle is not easily observed. Divisions take place almost simultaneously in all the cells of a sporogonium and the changes are very rapid. By far the commonest stage of division is that in which the chromosomes are in the nuclear plate (fig. 38). Neither centrosomes nor centrospheres occur in the spore mother-cells of either Riccia crystallina or Riccia natans. Around the nucleus preceding the formation of the spindle, there is an accu- mulation of material, apparently composed of fine fibres. The nucleus elongates, becoming somewhat elliptical but not sharp. pointed. The fibres about the nucleus do not give the appearance of centrospheres but are like the weft of kinoplasmic fibres described for certain pollen mother-cells (jigs. 36, 37). It has been impossible to find any nucleus which showed anything resembling a multipolar spindle. The poles of the spindle are probably determined by the elongation of the nucleus at an early stage in the spindle formation. The spindle is composed of very fine fibres, some of which extend from pole to pole, while others extend from the poles into the cytoplasm, reaching almost to the nuclear plate (fig. 38). The 124 BOTANICAL GAZETTE [FEBRUARY mature spindle has very broad poles and its formation does not seem to have been controlled by a centrosome or a centrosphere, as a comparison of the spindles of the spore mother-cells with those of the cells of the antheridium makes clear. The minute chromosomes separate, four going to each pole, after which a cell plate is formed in the usual way (figs. go, 41). The daughter nuclei do not come to a true resting stage. The chromatin is scattered in almost spherical bodies in the hyaline cavity of the nucleus, which do not represent the individual chromosomes, as their number and size vary considerably (jig. 42). The second division takes place in much the same manner as the first. The spindles are arranged with their long axes parallel to the first cell plate, so that the cell plates formed in these spindles are almost perpendicular to that formed in the first division (figs. 43-47). The latter does not disappear during the second division but remains and the walls separating the spores are laid down here (fig. 47). The walls separating the cells of the young tetrad are thin and delicate, but in the mature spore the outer layer of the wall becomes thickened and folded. The mature spore is almost black, and its contents are largely oil. When carried through chloro- form into paraffin and sectioned, the spores seem to have only scanty . granular contents, due to the fact that the oil has been removed in the process. The nucleus is very small. During the winter and spring following their development, unsuc- cessful attempts were made to germinate the spores. It may be that they had been allowed to remain dry too long before they were moistened, for in nature they would not be dry very long even in tiding over a dry season. ; The spore mother-cells of Riccia natans do not furnish such satisfactory material for study as do those of Riccia crystallina, because-the large air cavities of the thallus prevent the penetration of the fixing fluid and so the spore mother-cells often shrunk. sufficient number of good preparations was secured, however, to show that the process of division does not differ from that of Riccia crystallina. | 1906] LEWIS—DEVELOPMENT OF RICCIA 125 SPERMATOGENESIS. The development of the spermatozoids has been treated by a number of investigators, among them, CAMPBELL (4), LECLERC DU SABLON (23), GUIGNARD (14), SCHOTTLANDER (27), and StTRas- BURGER (29). It will be observed that most of these papers were published before methods of preparing material for study were so well developed as at present. The work of BELAJEFF (1)-confirmed by that of STRASBURGER (29) shows that the spermatozoid in the Hepaticae consists not only of the metamorphosed nucleus but also of the cytoplasm. IkENo (16) not only confirms the view that the spermatozoid consists of cytoplasm as well as nucleus but also discusses the develop- ment of the cilia and the homology of the blepharoplast and centro- some of Marchantia polymorpha. He finds that the body which becomes a blepharoplast in the developing spermatozoid appears in the earlier nuclear divisions of the antheridium and functions as a centrosome. It is, however, not permanent, but appears at the time of nuclear division and disappears during the process, so that it is not found in the daughter cells until about the time for the formation of the spindles of their division. After the last division which gives rise to the cells that develop into the spermatozoids, the body does not disappear but remains and becomes a blepharoplast. IKENo argues from this that the centro- Some and blepharoplast are homologous. He has good grounds for Such an argument in the case of Marchantia polymorpha, because centrosomes have been reported also in the vegetative cells of that plant, by Morrrer (24) and by VAN Hook (32). In other plants which have the blepharoplast, centrosomes are not found, and the body appears in only one or two‘divisions before the formation of the cells which produce the spermatozoids. Morrtier (26) in discussing IKENO’s Paper raises the question whether the bodies which Ikeno has figured as are in some Cases more than ordinary granules such as appear in the cytoplasm of other cells in which centrosomes are known to be absent. IKENO has pointed out, however, that the cytoplasm of these cells is very finely granular, there being no other bodies in the cell which bear any resemblance to the ones figured as centrosomes. He also calls 126 BOTANICAL GAZETTE [FEBRUARY attention to the fact that centrospheres have been described in dividing spore mother-cells of Pellia epiphylla, by FARMER (6, 7, 8, 10) and by Davis (5). The occurrence of centrospheres here has been questioned, however, by GREGOIRE (13). In a recent paper, FAR- MER (Q) reports centrospheres and occasional centrosomes in the spore mother-cells of Aneura pinguts. In order to get good results in Riccia natans it is necessary to fix the material when growing rapidly. About equally good results were secured with chromacetic acid and with Flemming’s weaker solution. The sections were stained with anilin safranin and gentian violet. It was found best to stain deeply in gentian violet and then to wash out carefully. In this way all details can be brought out clearly, although IkENo did not find it good for Marchantia. The development of the antheridium has been described. When almost mature it consists of a large central mass of cubical cells surrounded by a wall one cell in thickness (fig. 33). In preparations from plants in which some antheridia are mature, one finds several stages in the development. The nuclear divisions do not take place simultaneously throughout an antheridium but usually all the cells of one of the segments marked out by the first walls dividing the antheridium, show the nuclei in the same stage of karyokinesis. In the most favorable preparations, therefore, one may find several stages of division in the same antheridium. The cells of the young antheridium are almost cubical, with finely granular cytoplasm. The nucleus is rarely exactly spherical and has a rather thick membrane. The chromatin is in an irregular central mass, made up of a number of pieces. A nucleolus cannot be distinguished. The cavity surrounding the chromatin is large and hyaline (figs. 53, 54). In some cases a large number of small bodies of chromatin were found scattered irregularly in the nuclear cavity. The number of chromosomes is four. It seems that the nuclei in the young rapidly growing antheridium rarely come to a typical resting stage. The question of the presence or absence of centrosomes in the cells of the young antheridium was taken up carefully, because previous observations on the karyokinetic figures in the sporophyte cells and spore mother-cells have convinced me that no such body 0 1906] LEWIS—DEVELOPMENT OF RICCIA 127 appears there. On the other hand centrosome-like bodies appear in the cells of the older antheridia at the time of nuclear division. There can be no doubt that these are distinct bodies, and they cannot possibly be interpreted as accidental granules in that position. In Some of my preparations hundreds of cells showing them are found on a single slide, and they are so distinct that the preparation could easily be used for class demonstration. These bodies appear in the cells of the antheridium in early stages of its development. I have been unable to determine whether they appear in the earliest cell divisions but they appear in the antheridia which consist of only a few cells. They are not permanent, but disappear and arise anew with each division. IKENO regarded it as highly probable (though unable to state this positively) that in Marchantia these bodies were of nuclear origin. He figures a small spherical body inside the nuclear membrane, which in a later stage is found outside the membrane. This body then divides into two, which arrange themselves on opposite sides of the nucleus. If the bodies have their origin as one, which later divides as described, they act as do the ceritrosomes which have been described for other plants. In Riccia natans, nothing has been observed to indicate that the body is of nuclear origin, except that it stains in much the same Way as the mass of chromatin in the nucleus. In some of my pre- parations a single body has been observed near the nuclear membrane (fig. 53). These bodies have never seemed so distinct as the ones Which appear at the opposite ends of the nucleus and in the poles of the spindle. There is a dark central part, surrounded by a mass of cytoplasm which is more or less irregular but does not give the appearance of distinct radiations such as are described i in the centro- spheres of certain plants. When these single bodies were discovered, a careful search was made of the same preparations and of others in which the two bodies were on the opposite sides of the nucleus, in order to discover if possible the intermediate stages which it would seem should appear in such preparations. In cases in which two bodies have been observed, they have always been on opposite sides of the nucleus, ©r so nearly opposite that -the spindle developing between them 128 BOTANICAL GAZETTE [FEBRUARY might take the curved form shown in jig. 60. The origin of the two bodies is of importance in determining the homology of the centrosome and blepharoplast and will be discussed later. Starting with the stage in which the centrosome-like bodies are on opposite sides of the nucleus, the nuclear division takes place in the following manner. At first the bodies are at a little distance from the nuclear membrane, then the nucleus elongates so that the membrane closely approaches the bodies, becoming somewhat pointed. At the same time one observes that there is a collection of kinoplasm at the poles of the nucleus and extending along the nuclear membrane for some distance. At this time the bodies at the poles do not show radiations in any direction, but are very distinct (fig. 54). The spindle is formed from the kinoplasm which has been described, and when formed consists of a few thick fibres which converge at the poles, so that the centrosome-like bodies occupy the position of true centrosomes. About the time when the spindle develops, the chromosomes are formed from the central mass of the nucleus and become arranged in the nuclear plate. They are closely crowded together in this stage, and not so easily counted as when they have moved to the poles. The photograph (figs. 75, 76) shows the dense mass formed by the chromosomes when arranged in the nuclear plate. It was impossible to determine how the division takes place in the chromosomes as they move to the poles. The changes take place so rapidly that stages are rarely found in which the chromo- somes are on their way to the poles. The centrosome-like bodies disappear during the division, but it is difficult to say at just what point. Fig. 56, a cell taken from an antheridium in which only one or two more divisions will take place, shows the centrosome-like bodies quite distinctly when the chromosomes are almost at the poles, but by the time the chromosomes are at the poles and before the daughter nuclei are formed, the bodies disappear (fig. 57). These bodies are best seen in preparations which have been over- stained and washed out. In some cases my preparations were stained deeply enough to show the spindle and chromosomes well, but only an occasional spindle showed the bodies at the poles. When these slides were over-stained and carefully washed out, the bodies were brought out very distinctly in all cases. ; 1906] LEWIS—DEVELOPMENT OF RICCIA 129 After a large number of divisions has taken place the antheridium consists of nearly cubical cells, each of which has been considered by earlier investigators to produce a single spermatozoid. SrrRas- BURGER (30, p. 482) says of Marchantia polymorpha: “Die Spezial- mutterzellen der Spermatozoiden sind durch fortgesetzte, sich rechtwinklig schneidende Teilungsschnitte angelegt worden.” Camp- BELL (4) describes and figures the spermatozoid mother-cell of Pellia as producing two spermatozoids. IKENO (16) discovered that in Marchantia each of the cubical cells undergoes another division in which the spindles are arranged diagonally, in the earlier divisions the long axis of the spindle being parallel to the long axis of the cell. In this last diagonal division no cell wall is formed between the daughter cells, each of which develops into a spermatozoid. Thus each of the cubical cells produces two spermatozoids instead of one. IKENO cites several cases in which two spermatozoids are produced from a single mother-cell and thinks that this is probably general in the liverworts and mosses. JOHNSON (18) has described a diagonal division of the cubical cells of Monoclea, but he figures a wall separating the two parts of the cell and regards each three-cornered cell as the mother-cell of a spermatozoid. He does not give the details of nuclear division in the earlier stages of the antheridium nor in the formation of the Spermatozoid mother-cells. In the last division of the cells in the antheridium of Riccia natans the spindles are arranged diagonally as in Marchantia. This arrange- ment of the spindles is quite striking. They are larger than in the earlier divisions and the bodies at the poles are very distinct. In Some cases the spindles are curved (figs. 58-60). No wall is formed between the daughter cells, each of which develops into a spermatozoid. The centrosome-like bodies do not disappear after this division (fig. 6r). They remain in the cells, at first near the nuclei. The daughter cells are contracted, occupying the central part of the cell cavity (figs. 62, 63). Soon the centrosome- like body moves away from the nucleus toward the end of the cell. - Those in the two spermatids may be at the same end or at opposite ends (figs. 63-67). When the spermatid has become somewhat rounded, the centrosome-like body has taken its position in contact 130 BOTANICAL GAZETTE [FEBRUARY with the cell membrane (jig. 68). When the cilia appear they are inserted in this very small body so that it comes to function as a blepharoplast. Its small size as compared with that of the cilia of the mature sperm makes it seem probable that some of the material for the growth of the cilia must be drawn from another source than the blepharoplast itself, although it disappears to such an extent that in the mature sperm it cannot be recognized as the point of insertion of the cilia. The developing spermatozoids of Riccia natans do not remain enclosed in the mother-cells until they are mature, but at about the stage represented by figs. 70, 71 the walls break down and the young spermatozoids lie free in the cavity of the antheridium. Here they seem to undergo considerable growth. The material for this growth is probably derived from the surrounding cells as they become collapsed in old antheridia. The nucleus of the developing spermatozoid takes a position at one side of the cell and becomes homogeneous. It seems probable that other material than the chromatin of the spermatid nucleus must enter into this part of the spermatozoid, because it is very evident that the body contains more material than would be obtained from the chromatin alone. Soon the nucleus elongates, following the outline of the cell and becoming crescent-shaped (figs. 71-73): In some cases, a distinct vacuole occurs in the cytoplasm although this is not always the case (figs. 71, 72). The mature spermatozoid becomes long and slender and consists of the nucleus, the material of which seems to have increased in amount, a small amount of cytoplasm, and the cilia which are derived from the blepharoplast and in all probability from a part of the cytoplasm surrounding it. IKENO describes a spherical body which appears in the spermatids of Marchantia before the cilia begin to develop and disappears about the time that changes take place in the nucleus. It has been impos- sible to find such a body in Riccia, although it would seem, judging from IKENo’s figures, that it could easily be seen if present. The question of the homology of the blepharoplast and centro- some is one which it seems to me has not yet been settled. In Mar- chantia, where centrosomes have been reported in the vegetative cells as well as in the antheridium, the evidence seems good that the i 1906] LEWIS—DEVELOPMENT OF RICCIA 131 centrosome and the blepharoplast are homologous, and this is the conclusion of keno. In all other plants in which blepharoplasts are known to occur, centrosome-like bodies are not present in any cell divisions except those immediately preceding the formation of the sperms. That centrosomes occur in liverworts in cells outside the antheridium is open to question. The conflicting reports of those who have investigated Pellia epiphylla make it clear that no distinct body occurs there which can be regarded as a centrosome, although aggregations of kinoplasm, called centrospheres by most authors, do occur. In Riccia natans, it seems very evident that centrosomes do not occur in the divisions of the spore mother-cells. The spindle poles are broad, and there is not even a suggestion of a centrosphere such as has been described ‘for Pellia. In the cells of the sporophyte GARBER reports centrospheres but no centrosomes. I have never been able to observe them in my preparations. When the spindle is fully formed there are no fibres radiating into the surrounding cytoplasm (figs. 48-52). Although the thallus of Riccia natans does not present favorable material for cytological study, a number of cells showing nuclear division in the gametophyte have been observed near the growing point. The greatest difficulty here is the presence of numerous deeply staining granules in the cell. In some cases granules resem- bling centrosomes appear at the poles of the spindle, but they do not differ in appearance from the other granules of the cell, and it seems probable that their occurrence here is accidental. Summing up, we find that in Riccia natans centrosomes are not found in the cells of the gametophyte, sporophyte, or spore mother- cells, but that bodies occur in the dividing cells of the antheridium which seem to function as centrosomes. In Riccia and Marchantia, the blepharoplasts certainly have much more the appearance of centrosomes than in any other plants in which blepharoplasts have been described. The bodies have every appearance of centrosomes when at the poles of the elongated nucleus or at the poles of the spindle. Perhaps the strongest objection to regarding these bodies as centrosomes lies in the fact that in Riccia natans they occur only in the cells of the antheridium, while the blepharoplasts reported 132 BOTANICAL GAZETTE [FEBRUARY in other plants appear only in the last two generations of cells con- cerned in the formation of spermatozoids. Those who argue in favor of the homology of the centrosome and blepharoplast certainly find their best evidence so far in the liverworts, but it seems to me that this evidence is not conclusive when the bodies occur only in cells of the antheridium. In those plants in which centrosomes are known to occur, a single body divides to produce two, which arrange themselves on opposite sides of the nucleus (MoTTIER, 25). IKENO has reported a similar condition in Marchantia. In Riccia natans, however, the evidence seems to favor the view that the two bodies arise anew with each division, appearing on opposite sides of the nucleus at the same time. In this respect they behave more like blepharoplasts. MorrieR (26) in discussing this question has called attention to the fact that it is questionable whether we can speak of organs as homologous which, as such, are without genetic continuity. The question as to whether true centrosomes have genetic continuity has not yet been decided, but it is probable that they do not in all cases. | SUMMARY. 1. Reccia lutescens and Ricciocarpus natans are forms of the same plant, the former occurring on the ground in summer and autumn when the ponds are dry, and the latter as a floating form. Either form can be changed into the other by altering the supply of water. Therefore, Riccia Jutescens should not be regarded as a distinct species. : 2. The genus Ricciocarpus has been based largely on characters which do not exist. In my opinion, the only real basis for separating it from Riccia is the more complex structure of the thallus. BiscHOFF did not regard this as a good character for the separation of the genus. 3. The plant is monoecious, antheridia and archegonia being produced in definite groups in the same thallus. The sexual organs appear in autumn when the thalli are growing on the ground and complete their development the following April. Abundance of water is not essential to sexual reproduction, as the plants fruit when kept growing on the soil and supplied with a limited amount of a 1906] LEWIS—DEVELOPMENT OF RICCIA 133 water; therefore the ground form is not sterile, as was the opinion of LINDBERG and GARBER. 4. Plants which have been growing attached to the soil and have been submerged by the filling up of the ponds do not necessarily perish, but are adapted to spend the winter under water and then to break loose by the decay of the older part of the thallus and float upon the water in the spring. 5. The plants are propagated vegetatively by the separation of branches of the thallus, by the decay of the older part, and also by the growth of new plants from cells in the apical region. 6. The sexual organs and fruit of the two species studied agree in their development with the accounts given for the other species of Riccia. There is no rudimentary integument surrounding the archegonium or sporophyte of Riccia natans. The sporogonium of Riccia natans is larger than that of Riccia crystallina and produces a larger number of spores. The only sterile tissue in either is the amphithecium, a single layer of tabular cells surrounding the mass of spore mother-cells. 7- Centrosomes are not present in cells outside the antheridium nor would I interpret any structure observed in the cells of the 5 eee phyte or the spore mother-cells as a centrosphere. 8. Bodies which resemble centrosomes, and which are con- sidered to be true centrosomes by certain authors, occur in the cells of the antheridium. These bodies do not have genetic’ continuity, but arise de novo with each division. They do not disappear after the last division of antheridial cells but remain in the spermatids and later become blepharoplasts. 9. In the earlier divisions of cells in the antheridium, the spindle is arranged parallel to the long axis of the cell, but in the last division, the spindle is placed diagonally in the cell. No wall is formed between the two cells produced by this division, each of which becomes a spermatozoid. Thus two sperms are produced from each cuboidal cell. , 10. In the developing sperm, the blepharoplast takes a position on the membrane of the cell and the two cilia grow from it, the nucleus becomes almost homogeneous in structure and _ crescent-shaped, almost enclosing the cytoplasm. The mature sperm consists of the 134 ; BOTANICAL GAZETTE [FEBRUARY nucleus, the cytoplasm, and cilia which have received material for their growth from the blepharoplast and probably also from the material surrounding it. 11. The amount of chromatin in the nucleus is small. There is no nucleolus present unless the masses of chromatin which are found in nuclei which are undergoing repeated division be inter- preted as nucleoli. 12. The number of chromosomes is four for the gametophyte and eight for the sporophyte. 13. The cytoplasm of the spore mother-cells appears to be a fine reticulum, in which are numerous granules usually located at the point of intersection of the fibres of the reticulum. 14. The mature spore contains a large quantity of oil together with a small amount of granular matter. The nucleus of the spore is very small. In conclusion I wish to thank Professor Gro. F. ATKINSON and Dr. E. J. Duranp for valuable advice and assistance during the progress of this study. LITERATURE CITED. 1.-BELAJEFF, Ueber Bau und Entwickelung der Antherozoiden. Heft 5 Characeen. 1892 (Russian). German translation, Flora 79:1-48. 1894. 2. BriscHorr, Bemerkungen iiber die Lebermoose vorzuglich aus den Gruppen der Marchantieen und Riccieen. Nova Acta Acad. Caes. Leop. Carol. Nat. Am. 17: part 1. 1835. 3- CAmpBELL, D. H., The structure and development of mosses and ferns. jigs. 1-7. New York. 1895. , Zur Entwickelungsgeschichte der Spermatozoiden. Ber. Deutsch. Bot. Gout 5:120-127. pl. 6. 1887. 5- Davis, B. M., Nuclear stadicy: in Pellia. Annals of Botany 9:147-180, pls. 10-11. 1895. 6. Farmer, J. B., On spore formation and nuclear division in the Hepaticae. Annals of Hotaiey 9:469-523. pls. 16-18. 4. 7. , The quadripolar spindle in the spore iiotherecll of Pellia epiphylla. Anaale of Botany 15: 431-433. Igor. 8. , On the interpretation of the quadripolar spindle in the Hepaticae. Bor. ‘Gacexee 37:63-65. 1903. 9. and Moors, J. E. S., On the maiotic phase (reduction divisions) in animals and plants. Quart. Jour. Mic. Sci. 48:489-557. pls. 34-41- 1905 1906] LEWIS—DEVELOPMENT OF RICCIA 135 4 an La NI al oo and Reeves, J., On the occurrence of centrospheres in Pellia epiphylla Nees. Annals of Botany 8:219-224. pl. 14. 18 - GARBER, J. F., The life history of Ricciocarpus natans. Bot. GazETTE 37: 161-177. ‘ls. 9-10 as - Ganone, W. F., The veetation “ the Bay of Fundy.salt and diked marshes. Bor. Gazerre 36: 420-455. 903: - GrécorreE, V., La figure achromatique dans le Pellia epiphylla. La Cellule 21:193-239. pls. 1-2. 1904. GuIGNARD, L., Développement et constitution des anthérozoides. Revue Gén. Bot. 1:10-27. 1889 - Hormetster, W., Vergleichende Untersuchungen der héherer Kryptogamen. Leipzig. 1857. English translation: “The higher Cryptogamia,” Ray Society. 1862. - IkEno, S., Die Beer aes von Marchantia polymorpha. Beih. Bot. Cena 15: 65-88. 1903. - JANcCzEwskI, E. von, Vergleichende Untersuchungen iiber die Entwicke- lungsgeschichte des Archegoniums. Bot. Zeit. 30:377-393, 401-417, 440-443. 1873. - Jounson, D.S. Peer aa and relationship of Monoclea. Bot. GAZETTE 36: 185-205. pls. 16- Kwny, L., Ueber Bau a ine der Riccien. Jahrb. Wiss. Bot. ve 364-386. pls. 44-46. 1866-67. LINDENBERG, Monograph. Nova Acta Acad. Caes. Leop. Carol. Nat. m. . Exrpnese, Reds Bryol. 9:82. 1882 - Lertces, H., Die Riccien, Untersuchungen iiber die Lebermoose 4:1-101. pls. 1-9. 186. - Lecierc Du Saston, Sur la formation des anthérozoides des Hépatiques. Compt. Rend. Acad. Sci. Paris 106:876-878. 1888. - Morrier, D. M., The centrosome in cells of the gametophyte of Mar- chantia. Proc. Ind. Acad. Sci. 1898: 166-168. 1899. , Das Centrosome bei — Ber. Deutsch. Bot. Gesells. 16: 1ag-re8, jigs. 5. 1898. , The Ss of the spermatozoid in Chara. Annals of Botany 18: sae 254. pl. 17 SCHOTTLANDER, P., Beitriige zur Kenntniss des Zellkerns und der Sexual- pi bei Key peng, Beitr. Biol. Pflanzen 6:267-304. pls. ° 4-5.. aa F., Bull. Herb. Boissier 6:377. 1898. STRASBURGER, E., Schwarmsporen, Gameten, pflanzliche Spermatozoiden, und das Wesen der Befruchtung. Histol. Beitr. 4:——. 1892. ———, Botanisches Praktikum, 4 Aufl. ayes: . UNDERWwoo p, L. M., Systematic botany of North America. Hepaticae, advance sheets. ‘Sige, » 136 BOTANICAL GAZETTE [FEBRUARY 32. ig Sgr J. M., Notes on’the division of the cell and nucleus in liverworts. T. GAZETTE 30: 394-398. aa: seca H., Ueber die Regeneration gs Marchantieen. Jahrb. Wiss. Bot. 10:367-414. pls. 12-15. 1885. EXPLANATION OF PLATES V-IX. All drawings, except fig. 7, were made with camera lucida. Figs. 8-52, with Bausch & Lomb oculars and objectives, as follows: Figs. 8, 10, 11, 1 in. ocular, % objective; 9 and 25 2 in. oc., $ obj.; 19, 21-24, 26, 32, 33, 2 im. oc., wy obj.; 12-18, 20, 27-31, 34-52, I in. on as obj.; Figs. 53-73 with Zeiss oc. 18, 2™™ apochromatic objective, 1.40 apert The figures of plate VI were oe slightly more than one-half in repro- duction. All figures are of Riccia natans except the spore mother-cells (figs. 34-47) which are of Riccia crystallina. Fic. 1. Rosette of plants awhes on ae soil; a, natural size; b, enlarged. Fic. 2. Land plants growing in regular clusters. Fic. 3. Two plants growing on soil, one of which has been igaud and has grown out in an irregular way from the growing point. Fic. 4. New plants growing from apical cells of old thalli. Fic. 5. a, Decay of older part of thallus of the land form to give the floating form; b, plants collected in May. If these thalli should become stranded on the mud and growth should continue rosettes would be formed. Fic. 6. Plants decolorized in alcohol. The sporophytes appear as chains of dark bodies in the thallus. PLATE VI. Fic. 7. Longitudinal section of thallus parallel to the dorsal furrow, showing arrangement of sexual organs. Fic. 8. Cross-section of thallus showing archegonia. Fic. 9. Cross-section of thallus, showing the origin of archegonia in rows on floor of dorsal groove. Fic. 10.. Cross-section of thallus, antheridia. Fic. 11. Longitudinal section of thallus parallel to surface, showing the arrangement of antheridia. Archegonia have not begun to develop. 1Gs. 12-18. Stages in development of archegonium. 1G. 19. Archegonium in which egg-cell has not been fertilized and is shrunken Fic. 20. Cross-section of neck of archegonium. Fic. 21-25. Stages in development of sporophyte. Fic. 26. Spore mother-cells. : Fics. 27-33. Development of antheridium. Figs. 27-31, from materia] collected in October. TTR en Cpe secs BOTANICAL GAXETTE, XLI LEWIS on RICCIA BOTANICAL GAZETTE, XLI PEATE VI S zs, pele g =, U ve | SS UI gee O sal a LEWIS on RICCIA PEATE Vil ET bd BOTANICAL GAZETTE Seah 2, Site I o) c= man CS Y ne , Bh 6 on +] . Ss tx KS Petan NJ LEWIS on RICCIA y = rt Rog 4) oe ax f\ PLATE VIII 4 Bret y ieata BOR S ‘ LS SU OR LI PSI HS TOR A x REDE Py? sores 4, OR PAS EDO af AN 7 Ly aan Bese na Sa STORER TARE RES CSP RGAE AS uy Gh e, OK eee re ence CACY Car a is me aay Wy eats RSE Nf ¢, LOR ROR RS MG art a OB, ay oy uO a, J sae ry oe) es oe LEWIS on RICCIA NICAL GAZETTE, XLI BOTA BOTANICAL GAZETTE, XLI PLALE, 1X LEWIS on RICCIA 1906] LEWIS—DELOPMENT OF RICCIA 137 PLATE Vii. Fic. 34. Spore mother-cell in resting state. Chromatin on a fine linin net- Fic. 35. The chromatin is in the form of an irregular thread. Fic. 36. Chromosomes formed; weft of delicate fibres about the nucleus. Fic. 37. Nucleus elongated and showing a weft of fibres. Fic. 38. Spindle with chromosomes in plate. No centrosome. Fic. 39. Chromosomes moving to poles of spindle. Fic. 40. Chromosomes at the poles, thickening of spindle fibres to form cell Fic. 41. Daughter nuclei. G. 42. Cell plate. Daughter nuclei contain numerous spherical bodies of chromatin which stain bright red with safranin. Fic. 43. Daughter nuclei preparing for divi Fic. 44. Daughter nucleus with casa in plate. Neither centro- sphere nor centrosome. PLATE VIII. Fic. 45. Chromosomes moving to poles. 1G. 46. Daughter nuclei with chromosomes at the poles to form nuclei of spores. The cell plate formed in the first division persists. Fic. 47. Second division completed. Ss. 48-52. Stages in the division of a sporophyte cell. No centrosome. Fig. 48 shows slight radiation of cytoplasm from the poles of the elongated nucleus. Fic. 53. Cells of antheridium which show a single rather irregular body near the nucleus. Fic. 54. Cells of antheridium which shows the distinct centrosome-like bodies at the poles of the elongated nuclei. Compare jig. 74. Fic. 55. Spindle with centrosome-like bodies at the poles. Fic. 56. Centrosome-like bodies present when the chromosomes are almost at the poles. Fic. 57. Cell from young antheridium. Chromosomes at poles. No centrosome can be distinguis| Fic. 58. One cell preparing for last division, while the adjoining cel] has the spindle formed and arranged diagonally. Fic. 59. Diagonal arrangement of spindles in last division of cells in the antheridium Fic. Oa: Curved spindles. : Fic. 61. — nuclei formed after diagonal division; centrosome-like bodies presen Fic. 62. Cell of antheridium after last division. Fics. 63-67. Spermatids in mother-cells. Fics. 68-73. Stages in the development of the spermatozoids. 138 BOTANICAL GAZETTE [FEBRUARY PLATE IX. G. 74. Antheridium in which the nuclei are elongated and preparing for Fic. 75. Portion of section of an antheridium showing the spindles with the dense chromosomes in the plate and in some cases the centrosome-like bodies at € poles. Fic. 76. Two cells of the same section enlarged three times. _ Fic. 77. Chromosomes at the poles of the spindles. _ Fic. 78. Cross-section of thallus showing the hyaline hairs which extend up into the median grooves. BRIEFER R ARTICLES. NOTE ON THE RELATION BETWEEN GROWTH OF ROOTS AND OF TOPS IN WHEAT. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. LXXXI Srupts in the experimental morphology of plants have dealt mainly with the subaerial portions, comparatively little attention having been paid to variations in the growth of roots. Thus the literature of the subject is very meager. Several authors, especially. MOELLER,’ DETMER,? PERSEKE,3 Mer,‘ Gat, and FREIDENFELT,® have studied the relation of water to the growth of these organs, with the general result that in water itself they grow longer and thinner, with fewer branches and root hairs than are observed in moist soils. Curiously enough, in soils which are too dry for optimum growth, the response is very similar. The roots are long and slender and possess few branches excepting near their tips, which lie, of course, in moister soil. The only observation on the relation of root growth to that of tops, with which the writer is acquainted, is that of MoELLER, to the effect that in a series of nutrient solutions, of concentrations ranging from 1.0 to 0.05 parts per thousand, the actual weight of roots produced varies generally with the concentration, but that the ratio of weight of roots to that of tops is much larger in the most dilute solution than in any of the others. A number of experiments have suggested to the author that the accel- fertierns H., Beitrage zur Kenntniss der Verzwergung. Landw. Jahrb. 13: 167-173. 2DETMER, ie Ueber den Einfluss dusserer Verhiltnisse auf die Wurzelentwick- elung. “ Landw. Versuchst. I5: 107-113. 1872 3PERSEKE, K., Ueber die Formveranderungen der Wurzel in Erde und Wasser. Leipzig. 1877. 4MerR, E. De Vinfluence des milieux sur la structure des racines. Comptes Rend. 88: 1277-1280. 1879. Recherches expérimentales sur les conditions de développement des poils radicaux. Ibid. ep anee de structure et de forme qu’éprouvent les racines suivant les sie oti elles végétent. Assoc. Franc. pour l’avance. sc. Compt. rend. de la 9° session. Rheims. 1880. 5GAIN, E., Réle ieilaae de l’eau dans la végétation. Paris. 1895. ©FREIDENFELT, F., Studien iiber die Wurzeln kraiitiger Pflanzen. I. Ueber die Formbildung der Wurzel vom biologischen Gesichtspunkte. Flora 91: 115-208. Tg902. 139] {Botanical Gazette, vol. 4x 140 BOTANICAL GAZETTE [FEBRUARY erating or retarding effect of the soil upon plant growth may often be due primarily to a response of the roots themselves, and that the ordinarily observed effect upon the tops may be due to the nature of the roots rather than to that of the soil directly. This question deserves thorough study ; the results to be given here cover only a very small portion of the field. This work was carried on at the laboratories of the Bureau of Soils of the U. S. Department of Agriculture, Washington, D. C. The plant used was the Russian variety of wheat known as “Chul.” The plants were grown directly from the seed in paraffined wire baskets of the form described by Wuitney and Cameron.’ As these authors have already pointed out, such baskets possess the advantage over pots of producing a root system uniformly distributed throughout the soil mass, rather than the accumulation of roots on the inner surface of the vessel which occurs in the case of ordinary pots. The studies to be discussed in this paper were made upon the roots developed in the first six series described in the author’s previous publica- tion’ on the growth of tops, and in similar cultures. The medium used was a very poor soil from Takoma Park, Md., and the same soil with varying amounts of fermented stable manure added thereto.2 The cultures of any series were placed side by side in a greenhouse, the amount of water in all the baskets being kept practically uniform by weighing at intervals of one or two days and adding the amount of water which was found to have been lost by transpiration. It is thus seen that the different cultures were all subjected to the same conditions excepting those which depend upon the treatment of the soil. Series I of the paper on growth of topst° will serve as an example ; the results of all the series are in accurate agreement. The soils and culture numbers were as follows : Basket 8 Number - 2 3 4 5 6 7 Takoma | Do. Do.+ Do.+ Do.+ Do Do.+ | Do+ Soil Soil 5000 10,000 15,000 20,000 30,000 40,000 50,000 p.p.m.‘f | p.p.m. p-p.m. p.p.m. p.p.m. p.p.m. p-p.m. untreated | manure Manure | manure manure manure manure | manure Sram 7Wauirney, M. and Cameron, F. K., Investigations in soil fertility. U. 5. Dept. Agric., tiiean of Soils, Bull. 23. 1904 SLIvINGSTON, B. E., Relation of sentation to growth in wheat. Bor. GAZz- ETTE fhe: 78-195. figs. 21. 1905. a description of this soil and a discussion of its properties, see LIVINGSTON, _E, genes ON, J. C., and Ret, F. R., Studies on the properties of a sterile soil. Ss. De ept. Apric., siren of Soils, Bull. 28. 190 104 photograph of the tops and data for their — are given in that paper. — abbreviation p. p. m. is used to denote parts per million by weight of air soil. 1906] BRIEFER ARTICLES 141 At the end of the experiment, which lasted seventeen days, the soil masses were taken from the baskets and the roots removed from them with as little injury to the latter as possible. The fresh roots thus prepared are shown in jig. z. The numbers correspond to the culture numbers given above. It is at once evident that the root system increases in amount throughout the series. Closer observation shows that this is due mainly to differences in the relative number and length of secondary roots and succeeding branches; the primary roots are of the same number Fic. 1.—Roots from a series of wheat cultures grown in Takoma soil untreated and in te same with addition of 5000 p.p.m. to 50,000 p.p.m. of stable manure in culture 8 as in culture r. They are somewhat longer and more slender in culture 8. Photographs of single root systems from a similar series are shown in figs. 2-5, which bring out the last point more clearly than fig. z. In fig. 2, which represents the roots from natural Takoma soil, very few branches are to be seen, and these are exceedingly short; practically the whole root system consists of the primary roots, with a few adventitious roots developed at the extreme base of the stem shortly after germination. ig. 3 shows roots from a soil containing 5000 parts per million of manure. 142 BOTANICAL GAZETTE [FEBRUARY A very slight increase in number and length of branches is to be observed. Fig. 4, from a soil containing 10,000 parts per million of manure, shows numerous well-developed branches, while from a soil with 40,000 parts per million of manure, as shown in jig. 5, the branches have increased so markedly in number and extent as to make up by far the greater part of the system. From these facts it is seen that, for this series of soils the variation in growth of tops is correlated with the number and length of lateral roots. The water content of all the soils was the same, so that the variations in —Wheat roots grown in Ta- 3-—Roots apie in Takoma Fic. Fic. koma ot untreated. soil with poe p-p.m. manu growth cannot be related to this factor; therefore they must be connected with some unknown condition in the soil itself, a condition which is related _ to the amount of manure present. The comparative anatomy of these roots was investigated, both by hand and paraffin sections, with the result that in the poor soils the main roots have a strong tendency to swell by direct enlargement of the cortical cells, without increase in the number of these cells, while in the better soils this tendency is not nearly so marked. Very old wheat roots from autumm stubble in the field show this balloon-like enlargement of the cortical cells to a still greater degree. This is apparently a phenomenon of age, sug- gesting that roots in the poor soils age more rapidly than in the better ones. It was also found that the zone of root hairs, which normally has its lower limit s-1o™™ from the root apex, extends in the poor soils to within 1-3™™ of the tip. The outgrowth of root hairs from the piliferous layer may also be related to the age of the cells; as is well known, these Se 1906] BRIEFER ARTICLES 143 organs normally appear only after the cells from which they arise have passed through their period of most rapid growt It would seem that the poor soil, by inhibits branch growth and causing the enlargement of cortical cells, may render the root system unable to carry on an adequate amount of absorption for normal growth, and that this fact may be the main clue to an explanation of the stunted tops in such cases. That the inadequacy of the stunted roots is in regard to the water supply rather than to that of salts, is indicated by the fact that in distilled IG. 4.—Roots grown in Takoma soil Fic. 5.—Roots grown in Takoma soil with 10,090 p-p-m. manure. with 40,000 p.p.m. manure. water, for the first two or three weeks, a better growth of tops is obtained than in the natural Takoma soil. Determination of the relation of the dry weight of the root system to the nature of the soil was deemed advisable, but a number of tests yielded only negative results. The dry weight of the complete system was found to be practically the same from all of the soils. The variations were always irregular. This may, of course, be partially due to the fact that, while it is manifestly impossible to obtain anything like the entire root System of one of the better cultures (owing to the extreme fineness of the branches and their adhesion to the soil particles), yet from the poorer cultures, where branches are short and few, a much larger part of the System is obtainable—Burton Epwarp Livincston, The University of Chicago, March 5, 1905. CURRENT LITERATURE. BOOK REVIEWS. The Swiss moors. ~* Fring and Scurorer have published a remarkable work on the moors of Switzerland, and have thus placed all who are interested in bog studies under lasting obligations. More than half of the huge volume is given over to the discussion of general geological, chemical, physical and biological problems connected with peat formation and the ecology of bog plants. One of the most valuable features is the discussion and scurry of the mass of European and foreign literature touching these problem After defining the scope of the nveignton, the authors take up the peat- producing plant associations of Switzerland. These are described with the greatest detail, from the plankton to i seca! vegetation; and their relation to peat accumulation is explained. e moors are distinguished primarily as flat bogs, raised bogs, and bogs of the alpine regions. The first two types are of such general occurrence that their peculiarities may be briefly summarized here. The Flachmoor is characterized as occurring in connection with waters rich in mineral matter, especially lime, in both wet and dry climates. Usually they show centripetal growth, and are dominated by species of Cyperaceae, Gram- ineae, Juncaceae and Hypneae. Species of Alnus, Betula, and Frangula make up the woody growth. Sphagnum, Ericaceae, and Empetrum are entirely wanting. n the other hand, the Hochmoor type occurs under the influence of waters poor in mineral ee where the rainfall is abundant and the tem- perature mild or cold. The ace is convex. The oldest portion is toward the center; hence the BO: is ccokatiad The dominant plants are Sphag- um, Oxycoccus, Andromeda, Calluna, Vaccinium, Empetrum, Eriophorum vaginatum, Pinus montana uncinata, Betula pubescens, and B. nana. These plants do not occur on the flat bogs, and are driven out by irrigation with waters of si mineral content, especially lime. his summary suggests the essential difference between American bogs and the poner: of European authors. Grass marshes occur here, which are strictly comparable. But the term “flat bog” is usually applied to areas tFrtg, J. and Scuréter, C., Die Moore der Schweiz mit ii ir der gesamten Moorfrage. Beitrage zur Geologie des Schweiz, herausgegeben von = Geol. Komm. der Schweiz. naturf. Gesells. Geotechnische Sire III Lief. 4to, pp. xviii+751. Bern: A. Francke. 1904. M 40. 144 1906] CURRENT LITERATURE . 145 having a peat substratum, dominated by Sphagnum, Cassandra, Andromeda, Oxycoccus, Ledum, Vaccinium, etc.—-a vegetation more nearly related flor- istically to the Hochmaor. Further, these occur in areas whose soil water is rich in mineral salts, frequently overlying marl! The raised bogs of America are strictly comparable to the Hochmoor of Europe, but their occurrence appears to be localized by climate rather than by the character of the soil r For both the flat and raised-bog plants, the authors conclude that the substratum is physiologically dry because of the combined influence of three factors: (1) high water-content of the substratum, (2) consequent low tempera- ture and (3) the difficulties in the way of root respiration (due to wet soil and scarcity of oxygen), accompanied by a general impairing of all root functions. The plants of the Flachmoor are noted for their ready absorption of mineral salts, absence of mycor ni unusual development of the — nd parts, and the high percentage of as cause of the xerophytic characters of many of the plants is not clea They a re probably connected with the difficulties of absorption. These at are hes hydrophilous, never occur- ring in dry situations. The Huchmoor plants, however, absorb mineral salts with difficulty; mycorhiza bas occurs in all the Ericaceae, in Empetrum, Betula, and Pinus; carnivorous plants are common; the root systems are poorly developed; and the ash content is low. The xesophyie character of the plants is in part due to the difficulties of absorption, and in part to their evergreen habit. Many raised-bog plants also occur in dry situations. The third and fourth chapters discuss the conditions and processes involved in peat accumulation, the chemical and physical properties of the end products, the classification of peats, the bog minerals, and the relation of bogs to coal deposits The geographic distribution of the bogs of Switzerland, a geomorphic classi- fication of the moors of the world, the relation of settlement to moor develop- ment, the economics of the Swiss moors (with bibliography), the bog deposits as records of the postglacial history of northern ngniariag form the other principal chapter headings of the first portion of the wo The second division gives a detailed description . the individual Swiss bogs. In most instances these include not only plant associations of the present surface but also the succession of plant remains occurring in the peat. The location of the bogs is made clear by-an excellent topographic map, upon which the distribution of the several bog types is shown. The bibliography occupies seventeen pages and includes only the more important purely scientific . Papers. e importance of the work from the standpoint of future investigation is undoubted. To American students it furnishes not only a key to the present status of the subject, but also a model for the study and description of our own bogs, marshes, and swamps.—E. N. TRANSEAU. 146 BOTANICAL GAZETTE [FEBRUARY Reproduction of mildews. Harper has brought together the results of several years of study of nuclear activities in the mildews*in a lengthy and beautifully illustrated publication from the Carnegie Institution.2 It is impossible for us to consider more than the striking new features of his investigations. The paper contains a résumé of much of his earlier work and a broad discussion of many cytological principles which are of general interest and will richly repay the reader of this very creditable contribution to American botany. The author takes a strong stand for critical morphological analysis and classification of the stages in the life history of thallophytes, with a clear separation of phylo- anaes history from physiological functions. m mportant new features of HARpER’s research, chiefly in Phyllactinia, are @) the establishment of a “central body” within the nucleus, which constitutes a point of attachment for the chromatic elements and gives a clear polarity to the structure, and its continuous existence through the most important phases in the life history; (2) the evidence for the permanence of the chromosomes; and (3) the evidence that the triple mitoses preceded by synapsis in the ascus constitute a double reduction of the chromosomes which are quadrupled by the two nuclear fusions in the life history, the first fusion at the time of the sexual act and the second fusion within the young ascus. The central body is a permanent structure, always present in the resting nucleus, dividing with each mitosis, and the center for an arrangement of chromatic threads within the nucleus and for the attachment of spindle fibers during nuclear division. Its position determines a pole in the. nucleus around which are grouped the chromatic elements, which are thus always in connection with the central body, both in the resting nucleus and during mitosis. This e succession of mitoses in the life-history. HARPER has not been able to i the different sets of chromosomes after the nuclear fusions, for the chromatic elements and the central bodies unite very intimately. But the second fusion in the life history, that in the ascus, is followed at once by a period of synapsis and the triple mitoses out of which come the eight chromo- somes characteristic of the gametophytic phase of the form.—B. M. Davis. MINOR NOTICES. Observations in Spitzbergen——The flora of Spitzbergen is fairly known. Therefore, Dk. WuLFr, who accompanied the Swedish expedition for the measure- ment of an arc of the meridian, undertook to make ecological observations on the arctic plants,3 especially touching their transpiration, occurrence of mycorhiza 2 HARPER, R. A., Sexual ee oe and the organization of the nucleus i certain mildews imp. 8vo. pp. . pls. 7. Washington: er Institution of apAbae gos. ULFF, THORILD, Observations botaniques faites au Spitzberg. Missions scien- ious pour a mesure d’un arc de méridien au Spitzberg. Mission uédoise. ‘Tome Il, X¢ section, Botanique. Traduit de Allemand par H. Marcet Harpy & Dundee. 4to. pp. 63, pls. 4. Stockholm. 1903. 1906] CURRENT LITERATURE 147 and anthocyan, the vegetation of the “polygonal” soils, and to make miscel- laneous floristic notes at various stations. The transpiration he finds very feeble and almost without diurnal periodicity or plant control. This feeble transpira- tion he accuses of being a cause of feeble growth; instead, is not its feebleness due to the same cause as the feebleness of growth, the low supply of energy ? Mycorhizas, internal and external, are common. Anthocyan is found in fifty species, about half the known higher plants. It is always lacking in plants grow- ing on soil enriched by the droppings of wild birds, whereas the same species growing on poor soils show it abundantly. As to the rdle of anthocyan, he holds it for an absorber of energy, and without it no plant can become dominant in arctic regions. For other interesting observations one must consult the work itself —C. R. B. Polypodiacez and edible fungi—Not that there is any connection between them; but both are treated by CopELAND in a bulletin+ from the Government Laboratories at Manila. The section on Polypodiacee forms the bulk of the bulletin and is “an attempt to collect and publish descriptions of all the ferns known to have been found in these islands.”” The author adds: “I am not per- sonally acquainted with a large part of those ferns still known here only from earlier collections.” Which leads us to remark that he should then have abstained from describing a new genus and new species among them, as hedid in Dr. PERKIns’s last Fragmenta. In reprinting here these descriptions he has neglected to indicate that they have already been published elsewhere. He has sinned again in adding one more new name in this bulletin. The compilation of such descriptive floras is undoubtedly serviceable; but one who is not a taxonomist and who confesses the absence of indispensable books and specimens, should not take the chances of cumbering pteridology with new names which may or may not be justified. And the same may be said regarding the brief fungus part.—C. RB. Genera of Mexican plants.—The flora of Mexico is so closely related to our own that any work on it is of essential assistance to American taxonomists. So we welcome the assembling and description of the Mexican genera, and the list- ing of the species, undertaken by Professor ConzaTTI, director of the State Nor- mal School of Oaxaca, of which the first volume, on Polypetalae, has recently been published by the Ministry of Public Works. This volume 5 begins with an artificial key covering about 50 pages*including all genera, and contains descrip- tions of 667 genera of Polypetale, representing 71 families, and including close to 4,500 species. This is to be followed by another on Gamopetalae and a third 4CopELAND, E. B., I. The Polypodiacee of the Philippine Islands. II]. New species of edible Philippine fungi. Bureau of Government Labs. Bull. 28. 8vo. pp. 146. pls. 3. 1905. sConzatTI, C., Los géneros vegetales mexicanos. Imp. 8vo. pp. 449. Mexico: Oficina Tip. de la Secretaria de Fomento. 1905. $3 (Mexican). 148 BOTANICAL GAZETTE [vEBRUARY on Monochlamydeae, Monocotyledoneae, Gymnospermeae, and Pteridophyta, embracing in all about 1900 genera. The descriptions are very full, and though the diagnostic characters are not indicated, this is largely atoned for “ the tae system of synoptic characters under the tribes and subtribes.—C. R Germs of mind in plants.—A little book,® unknown to us in the original French, now translated into English by A. M. Sruons, well-known for his work in Chicago along social and philanthropic lines, shows that there exists in France the same sort of popularizers of science as in our country—writers who with a smattering of scientific knowledge lack the fuller knowledge that forms a back- ground and furnishes scientific perspective. The facts of plant ecology are herein so distorted in their relation as to become caricatures; the use of words is so fan- ciful as to convert sober ideas into grotesque fairy-tales. For this, doubtless, the author is chiefly responsible; but the translator slips occasionally through unfamiliarity with a technical use of some common word. The book is interesting; but it is as little “science” as a historical novel is history. It is difficult to see how such fiction can be “a contribution to the cause of socialism and science.” —C. R. B Hepaticae of France.—LAcouture has prepared a helpful series of descriptive analytical keys to facilitate the identification of French liverworts by amateurs.’? The keys are arranged in a convenient bracket fashion, which is easy to use but makes the form of the thin volume rather unhandy and pre- cludes its use as a field manual. The description of each species is accompanied by an excellent figure illustrating tl t essential features described. The keys, in the form of tables, are arranged in three series, of which the first, consisting of tables 1 and 1m gives the characters of the tribes; the second, tables 11I-Ix the characters of the genera; and the third, tables xm-xxxIx, the characters of the species and the illustrations. No attempt is made to exhibit the natural classification —C. J. CHAMBERLAIN. Index Filicum.— The fourth and fifth fascicles of CHRISTENSEN’s important work® were issued respectively in October and December last. They carry the references from Cyat/:ea lanuginosa to Gleichenia cryptocarpa. The huge genus Dryopteris alone takes fifty-two pages, which indicates something of the compre- hensiveness of the work. Let colleges and libraries hasten to support by their 6FrANcE, R. H., Germs of mind in plants. Trans. by A. M. Smmons. 12mo. pp. 151. Chicago: C. H. Kerr & Co., 1905. 50 cts Lacouturr, CH., Hépatiques de la France. Tableaux synoptiques de. caractéres sedllatite des tribus, des genres, et des espéces. 4to. pp- 78. /#gs- 200- Paris: Paul Klincksieck. 1905. /r. 10. 8CHRISTENSEN, C., Index Filicum, etc. Fasc. 4,5. Copenhagen: H. Hagerups Boghandel. 1905. Each 3sh. 6d. 1906] CURRENT LITERATURE 149 orders the stupendous and too thankless task which the author has undertaken. The employment of the American system of citation is notable-—C. R. B. Das Pflanzenreich.°—Of this work parts 22 and 23 have lately appeared, including respectively the Primulaceae by Pax and Knurs, and the Halorrha- gaceae by SCHINDLER. The rate at which these monographs are appearing is remarkable, and shows something of the energy of the editor and his sagacity in the selection of his collaborators. The publisher’s part, too, is admirably done—C., RB, Eucalyptus.— Marpen’s revision’® has now reached part 7, which includes EE. regnans, vitellina, vitrea, dives, Andrewsi, and diversijolia, and is illustrated by four plates.—C. R. B NOTES FOR. STUDENTS. Items of taxonomic interest —ZAHLBRUCKNER lists (Beihefte Bot. Cent. 19?: 75-84. 1905) the lichens collected by Professor D, H. MreveEr in the Ecuador highlands in 1903, describing six new species.—Carpor (idem 85-148. figs. 39) enumerates 125 species of the mosses of Formosa, collected by Abbé Faurre in 1903, bringing the total known species of this island to 130, of which 39 are new. Herpetineuron (C. Mill. as Anomodon §) is raised to generic rank.— ENGLER describes (Bot. Jahrb. Syst. 37: 95, 96. 1905) a new genus of Araceae, Ulearum, and in his tenth contribution to a knowledge of the Araceae, (idem) adds to the family nearly a hundred new species, chiefly from Central America, the subequatorial andine province, the Philippines, and East Indies——Drere1, in his sixth paper on Japanese Uredineae (idem 97-109) describes 16 new species, and in one on Japanese fungi (dem 156-160) ten others.—RADLKOFER (idem 144-155) describes 8 new species of Serjania and 8 of Paullinia (Sapindaceae) from Peru, Brazil, Bolivia, and Columbia.—StTEPHANI (Bull. Herb. Boiss. IT. 5: 885-900, 917-946. 1905) in his Species Hepaticarum concludes the treatment of the genus Plagiochila, describing 26 new species, a number of them from equa- torial America.—Domn (idem 947, 948) describes 2 new species of Koeleria from Asia, and BEAUVERD (idem 948) a new Burmannia from Brazil and (g90-991 a new Hesperantha from the Transvaal.—FERNALD characterizes (Ottawa Nat. 19: 156. 1905) a new variety of Antennaria nevdiica Green from E. Quebec.— SCHNEIDER, in a prodromus to a monograph of Berberis (Bull. Herb. Boiss. II. 5: 139 ff. 1905) recognizes 159 species, among them a number of new ones of his own creation, which he divides into 21 sections. The regions of their 9 ENGLER, A., Das Pflanzenreich. Heft 22, ere by F. Pax and R. KnurH. pp. 386 Ae. 75 (311), maps 2. M 19. 20.—Heft 23. Halorrhagaceae by ANTON K. SCHINDLER. pp. 133, figs. 36 ae M 6. 80. Leipzig: Wilhelm Engelmann. oer 10 MAIDEN, J. H., A critical revision of the genus Eucalyptus. 4to. pp. 183-205, , pls. 33-36. Sydney: Government N. S. Wales. 1905. 2sh. 6d. 150 BOTANICAL GAZETTE [FEBRUARY dominance are South America and E. Asia ——HeEtier describes (Muhlenb. 1: 124) a new Veratrum from Idaho, and (idem 125) a Linanthus or Gilia from California.—McALPrnE adds a new genus, Uromycladium, to the Uredineae (Ann. Mycol. 3: 303-323. pls. 6-9. 1905). It is based on 7 Australian species occurring on Acacias, and is intermediate between Uromyces and Ravenelia.—VUILLEMIN shows the identity of Hartigiella with Meria (idem 340-343).—SCHMIDLE found eases ; . proposes an extension of the moss family Pterobryaceae to include five other families, in whole or in part, and gathers from various genera some 25 species to swell his genus Pterobryopsis. He establishes a new monotypic genus Miillerobryum on an Australian moss already referred to 3 separate genera. Trachypodaceae is a new family, and Trachypodopsis its characteristic new genus, for both of which he has “gathered of every kind,” and Teil I is only begun!—Prck (Rept. N. Y. State Botanist 1904) describes new fungi; Boletus (3), Clavaria (2), Cortinarius, Lactarius (2), Pholiota—CARDOT finds 35 new species of mosses in SkoTTSBERG’s collections made on the Swedish antarctic expedition (Bull. Herb. Boiss. II. 5: 997-1011. 1905).— Hieronymus has studied (Bot. Jahrb. Syst. 36: 458-573. 1905) the Compositae collected by JretsK1 in Peru, among which he finds 58 new species—— DIELS idem Beiblatt 82: 1-138) makes hundreds of additions to his flora of central yee eee ng many new species and three new genera, Giraldzella Dammer ), Pleroxygonum Dammer and Diels (Polygonaceae), and Biondia per (Necletiadarena), —NELSON describes (Proc. Biol. Soc. Wash. 18: 171- 776. 1905) new species from Nevada in Cleomella (2), Sphaerostigma, Zauschneria, Rhamnus, Polemonium, Artemisia, and a new genus of Solanaceae, Bosleria— ENAULD and Carpor in their tenth paper on Musci Evxotici (Bull. Soc. Roy. Bot. Belgique 41: 7-122. 1905) describe, among many others, largely Mascarene and East Indian, 9 new species from Porto Rico, 3 from Costa Rico, one from Guadeloupe, 3 from Cocos Island (Pacific Cent. Am.), 1 from Mexico, and 1 from Hawaii. They also establish as a new genus of Hypnaceae Miiller’s section of Hypnum, Dimorphella. The same authors (idem 123 ff.) in their third article on Musci Costaricenses describe 22 new species—-HELLER has found some new species in his collections for 1905 in California and describes them (Muhlenb. 2: 1-6. 1905), under Eriogonum (3), Montia, Del- phinium, Ranunculus, Thysanocarpus (2), Lithophragma, Ribes, and Amelan- chier.—Howe adds several algae to our flora (Bull. Torr. Bot. Club 32: 563-586. pls. 23-29. 1905) from the Bahama-Florida region; Halimeda, Avrainvillea, Sarcomenia, Dudresnaya, and a new genus Cadoci pati (Codiaceae), besides changing several names.—UNDERWOOD (idem 587-596) maintains the genus Alcicornium Gaud. as valid, gives a synopsis of the species, and describes A. Veitchii as a new species—R¥DBERG, about to publish a Flora of Colorado, makes (idem 597-610) what he considers necessary changes in names, and describes new species of Deschampsia, Eatonia, Poa (9), Festuca (2), and 1906] CURRENT LITERATURE 151 Elymus (2).—OsrerHout proposes from Colorado (idem 611-61 3) new species of Allionia, Aster, Senecio, and Carduus (2), which are respectfully referred to Mr. RYDBERG.—SARGENT adds (Rhodora 7: 192-219. 1905) 24 new species of Crataegus, all from New England.—Rostnson describes (idem 219-222) a new Ranunculus from Gaspé and Labrador.—C. R. B. Fossil gymnosperms.—Two trunks of Cycadoidea have been found in the Portland beds of Boulogne, to which MM. Fricue and ZEILLER give the specific name C. pumila on account of their small size."* Another Cycadoidea is des- cribed without attribution of a specific name. An interesting and important dis- covery is a cone of Sequoia of the S. giguntea type, which is named 5S. portlandica. The oldest well authenticated cone of Sequoia previously known is Heer’s S. lusitanica from the Wealden beds of Portugal, which belongs to the type repre- sented by the living S. sempervirens. It thus is demonstrated that Sequoia existed in its two living types as far back as the Jurassic period and must thus be very much more ancient in its first appearance. Other important discoveries are pine-cones representing the two main series of the present day, viz., the sections Strobus and Pinaster. The cone of the Strobus type is very much flattened and does not yield any definite information as to its internal organization, so the authors include it under the provisional fossil genus Pinites, with the specific appellation P. strobijormis, which would appear to be too close to our western Pinus strobi- jormis to stand as a permanent name. The other cone is exceedingly well pre- served and resembles very closely, as the authors point out, small cones of the living P. Luricio. This cone is referred to Pinus as P. Sauvagei. ‘These obser- vations are of very special interest because they establish that Pinus too must be a very old genus, since examples of both the hard and soft pine series existed already in the Jurassic. GoTHAN calls attention to the somewhat Puts TEA condition of X ylopa- laeontolgie at the present time and by comprehensive study of fossil and living woods, including many type-specimens of the former, reaches a number of con- clusions of greater or less importance.t? The proposition of FELrIx to divide fossil woods presenting tracheary structure resembling that of living Araucarineae, into Cordaioxyla for the palaeozoic woods, which may be supposed to be those of Cordaites, and into Araucarioxyla for mesozoic and later woods, is rejected, since in the author’s opinion no distinction can be made histologically between the two. For these w ENDLICHER’s name Dadoxylon is retained. Cedroxylon. Kraus and Cupressinoxylon Goeppert are separated from each other, not on the basis of the presence of resiniferous parenchyma in the latter genus and ts absence in the former, but on the character of the medullary ray-cells, since many Cedroxyla and even Pityoxyla have resinous parenchyma. This distinction has LICHE, P., et ZEILLER, R., Note sur un florule portlandienne des environs de Roper Me Bull. Soc. Geol. de la France IV. 4: 787-812. 1904. 2 GoTHAN, W., Zur Anatomie lebender und fossiler a ABhasdl. ag pilcaed: geol. Landesanstalt, Neue Folge, Heft 44. 152 BOTANICAL GAZETTE [FEBRUARY already and previously been clearly made by PENHALLOW. The author also attempts to separate the woods of the Podocarpez from those of the Cupressineae in the larger sense, on the basis of the structure of the pits in the ray-cells. The success of this distinction may be judged from the fact that it results in putting Sciadopitys with the Podocarpéae. Pityoxylon of Kraus is broken up by this writer into two genera, Piceoxylon and Pinusoxylon. The latter genus repre- sents the wood of Pinus, and seems somewhat unfortunate, since it is doubtful if the mesozoic pines had the wood structure which is found as characteristic of that genus in Tertiary and modern times. There are also disquisitious on spiral _Striation in the wood of the nosperms and on the value of annual woody rings as diagnostic of geologic formations. The work closes with two tables for the determination respectively of living and fossil gymnospermous woods. There is likewise an index and an alphabetical list of the living woods investigated by the author.—E. C. JEFFREY Injury by smoke.—F requent controversies and law suits, arising from damage to agricultural crops by the smoke produced by manufacturing establishments in Germany, have made the recognition of this form of injury extremely impor- tant. In order to furnish a basis for distinguishing smoke-injury from injuries due to other factors, SorAvER'S has made a comparative anatomical study of various kinds of injury commonly occurring in the more important grains, oats, wheat, and barley. The paper contains detailed comparative descriptions of changes in the cell walls and cell contents which cannot be severally noted here. The general plan followed in each case is represented by the following heads: e behavior of the normal plant in its gradual, natural dissolution; abnormali- ties in smoke-free regions; the phenomena in plants injured by chlorin and by hydrochloric acid fumes; experimental tests of the influence of hydrochloric acid fumes; phenomena confused with smoke injuries. In natural death the cells lose a large part of their contents and finally (except the epidermal cells) collapse completely. This process first involves the tip and edges of the leaves. In cases of death resulting from other causes, as drought, the cells do not collapse so completely, since the contents are not fully resorbed. In injuries due to acid fumes from smoke, the contents of the mesophyll cells contract into an irregular greenish lump, while the cell walls partially collapse. The most striking feature about this form of injury is the collapse of the epidermal cells. The accompanying changes of the cell contents and cell walls in these and in many other forms of i injury are minutely described. The recogni- tion of smoke injury in general is based on the fact that the cells, dying rapidly, collapse partially without being emptied of their contents, the epidermal cells showing the same phenomena. The author continually emphasizes the fact, however, that no clearly defined symptoms for the absolute and certain recogni- tion of smoke injury can be given, but that in all cases a comparative study of . 13 SORAUER, P., Beitrag zur anatomischen Analyse rauchbeschadigter Pflanzen. Landw. Jahrb. 33:585-664. pl. 15-18. 1904. . * 1906] CURRENT LITERATURE 153 plants growing under the immediate influence of the acid fumes and others growing under similar conditions but not within the smoke zone, must be made. —H. HaAssE.princ. Viticulture—Recent publications from the Royal Hungarian Central Insti- tute of Viticulture are as follows: Volume III, part 2, consists of chemical analyses of the stems and shoots of American species used for stocks in Hungary.'4 The points determined were the moisture content, ether extractives (oils, fats, waxes, ms, and organic acids not further determined), alcoholic extractives (tannin, pashan, vanillin, and organic acids), nitrogen, starch, cellulose, and pro- teids. e paper contains a large number of analyses made at different seasons, but no general results have yet been reached, and it is difficult to see what may be expected. Part 3 of this volume is a small paper by IstvANFFI'S in which he describes a disease of the vine caused by Phyllosticta Bizzozeriana Massal. The disease is not of great importance, but has been mistaken for the black rot, one of the most dangerous vine diseases. In the part 4 IstvANFrFI’® gives the results of his investigations on the gray rot, caused by Botrytis cinerea. The first part of this paper is taken up with the effects of various kinds of poisons and other treatments as cold, drying, etc., on the spores of the fungus. One of the most striking results is the unusually high resistance which the spores are said to have to copper. Spores were kept twenty-four hours in different strengths of Bordeaux mixture ranging from 1 to 10 per cent., to which was then added must containing I per cent. of tartaric acid, so that the resulting solutions contained the equivalent of 0.3 per cent. CuSO,. Of the spores from the lowest strength mixture 38-40 per cent, germinated, of those in the highest 10-12 per cent. germinated. Spores sown on berries in 3 per cent. Bordeaux mixture germinated and penetrated the epidermis. Spores, kept one hour in a 2 per cent. solution of CuSO,, which was then diluted with ten times its volume of must, germinated. Many other similar experiments are given. The second part of the paper deals with the development and life history of Botrytis cinerea and methods of control. Ve little new is added to the life history of the fungus. For treatment, spraying with a 5 per cent. solution of calcium bisulfid is recommended.—H. HasseLBRING. Endotrophic mycorhiza.— The long and important paper of GALLAUD'? on this subject merits brief summary, as his conclusions are quite revolutionary. He has described for the first time the anatomical and cytological characters of ™ Gaspar, J., Analyses des sarments américains. Ann. Inst. Cent. Ampél. Roy. Hongrois 3; nen pls. 4-12. 1905. 5 IsrvANrFFI, Gy. de, D’une maladie de la vigne causée par le Ph-yllosticta Bizzo- zeriana. Idem, 167-182. ap 1}. 1906: ‘6 IsTvANFFI, ae de, Etudes scbitnistegiaiinn et mycologiques sur le rot gris de la vigne. Idem 183-360. pls. 14-21. 1995. 17 GALLAUD, L, Etudes sur les mycorhizes endotrophes. Rev. Gén. Bot. 17: pls. 4. 1905. 154 BOTANICAL GAZETTE [FEBRUARY a large number of endophytes, and his study enables him to distinguish four types: (1) type of Arum maculatum, hyphae intercellular after traversing the outer cells, their growth arrested by formation of simple terminal haustoria which penetrate the cortical cells; (2) type of Paris quadrifolia, hyphae intra- cellular, of indefinite growth, with complex lateral haustoria arising at definite points; (3) type of Hepaticae, hyphae intracellular, of indefinite growth, enter- ing via rhizoids and bearing haustoria transformed into sporangioles; (4) type of Orchideae, hyphae intracellular, of indefinite growth, forming tight pellets which are sometimes permanent and sometimes undergo more or less complete digestion. There is a remarkable uniformity in the constitution of the cell walls and in the cytological structure.’ Repeated attempts to isolate the fungi by direct extraction and by inoculation were unsuccessful. The first method failed, prob- ably because the fungus already in was already too much altered by the digestive action of the host, and the second leads the author to distrust utterly the identi- fications of previous authors. The endophytes, he holds, are saprophytes internes, which by their highly differentiated haustoria borrow some non-living nutritive material from the cells in which they live. These cells react very rapidly on the fungus, killing its haustoria, digesting and absorbing them in part; then they resume normal life, momentarily disturbed. It cannot be said that there is a harmonious symbiosis between the two plants, but rather a conflict between the caotine but little harmful, fungus and the cells which defend themselves by their digestive power—C. R. B Sexual reproduction of Stigeoclonium.—PascHeER in an account of the sexual reproduction of Stigeoclonium fasciculatum,*® touches briefly on the formation and behavior of the zoospores (macrospores), which in general agrees with that of other forms, but in a few cases the sporelings developed into filaments of a few cells only, which then formed in each cell a single four-ciliate zoospore (macro- spore) that developed like other zoospores. The microspores are four-ciliate and long motile; after losing their motility they become spherical and either form resting-cells, or (rarely) conjugate and form zygotes. The development of the latter was not followed, but from hasty observation he concludes that their germi- nation does not depart from that of the zoospores or the resting-cells. After an indefinite period the resting-cells germinate like the zoospores. Some, however, (akinetes or palmella stage), grow into a few-celled filament, each cell giving rise to four biciliate zoospores, resembling the microspores in size and activity, except that they will not conjugate but germinate at once like the zoospores Phylogenetically he claims for Stigeoclonium fasciculatum a position midway between Ulothrix and Draparnaldia, the three kinds of spores indicating that it is on the border-line of sexual reproduction. The same position was long ago claimed by Dopet-Porrt for Ulothrix zonata. But such generalizations will bear, 8 Pascuer, A., Zur Kenntnis der geschlechtlichen Fortpflanzung bei Stigeo- clonium. Flora 95: 95-107. figs. 2. 1905. RS Ae NO ENR 1906] CURRENT LITERATURE 155 revision, and investigations of the cytological phenomena involved are especially needed. PascHer’s observations were microscopic to be sure, but he has appar- ently attempted no cytological observations at all—R. THrESsEN. Sigillarian stems.—Owing to the rarity of sigillarian stems showing structure the description of new specimens is of particular interest to paleobotanists. IDsTON’® has given a well-illustrated and adequate description of Sigillaria elegans, which differs from the historic S. Menardi in that the primary wood of the former is continuous instead of broken up into bundles. The protoxylem is external to the metaxylem, and both are composed of scalariform tracheids. The secondary wood is about equal in thickness to the primary, and shows medul- lary rays which are mostly one cell thick and one to nine cells high. The outer margin of the primary wood is crenate, and from the furrows arise the leaf traces, of which there are about twenty-eight in a cross section; these do not seem to possess any secondary wood. As is usual in sigillarian stems the pith, phloem, and inner cortex have perished, and the outer cortex contains a broad zone of periderm. . elegans, with a continuous ring of primary xylem, S. spinulosa, with a mixture of continuous and discrete xylem, and S. Menardi, with separate bundles, form a good series, and judging from the scanty data available it seems that this series represents a sequence in time. The features of S. elegans support the view that the genus sprung from forms more like Lepidodendron.—M. A. CHRYSLER. Mycoplasmic propagation of grain rust—Erixsson has published another instalment of his studies on the demonstration of the propagation of grain rust by means of mycoplasm, this time dealing with Puccinia graminis.?° Four means are recognized by which the uredo stage of the rust may possibly arise in spring time in winter wheat: (1) from spores of the barberry aecidium, which in turn arose from the resting teleutospores that had remained dormant over winter; (2) direct infection of the wheat plant from the resting teleutospores (homoecism); (3) uredo infection from mycelium remaining alive in the wheat plant over winter; and (4) from endogenous germs of disease (mycoplasm) which pass the winter in a resting condition in the live wheat plant. He marshals a large array of data, drawn from his own observations and experiments and from a wide range of literature, to show that the first method, although it exists, is by no means uni- versal, that the second is highly probable, that the third never occurs in northern regions, if anywhere, and that the fourth is the most common method everywhere. Although the conclusions of the author will not be accepted by most investigators of this difficult problem, yet the array of data is interesting. Two clearly drawn gans of Brongniart’s 19 Kipston, RosBeERT, On Histoire des végétaux jossiles. Trans. mele Soc. as 4r: 533-550. pls. 1-3. 1905. a9 > ERIKSSON, JaKop, Ueber das vegetative Leben der Getreiderostpilze IV: nia graminis Pers. in der heranwachsenden Getreidepflanze. Kungl. Sy. Vet.- tay Handl. 395:1-41. pls. I, 2. 1905. 156 BOTANICAL GAZETTE [FEBRUARY colored plates are used to show the author’s interpretation of the transformation of the resting mycoplasm into the mycelium condition of the rust.—J. C. ARTHUR Light relations at high altitudes—Wirsner’s study of the Lichtgenuss of plants, already comprehensive for varying latitudes, has now been extended?! to include high altitudes. During a period of thirty days from Aug. 16, photo- metric observations were made in the Yellowstone territory at eight altitudes ranging from 515 to 2210™ above sea level. The investigation shows that the behavior of plants with advancing latitude does not agree with that manifested under increasing altitude. The relative amount of available light appropriated by arctic plants increases inversely with the distance from the pole. This relation holds with increasing altitude only to a certain limit, above which a smaller and smaller share of available light is appropriated. The cypress habit of growth is evidently intended to protect from increased intensity of light, whether this accompanies low latitudes or high altitudes. This seems all the more probable heat, which is manifested by other species that do not show it at lower levels.— Raymonp H. Ponp. Tomato rot.—Von Oven”? has recently described a disease of tomatoes caused by Fusarium rubescens Appel & Von Oven. This fungus causes a rotting of the tomato fruit, and evidently does not belong to the fungi in this group producing stem rot or wilt disease, although in cultures the pink and violet shades char- acteristic of the latter are also produced by this new species. As it is impossible to separate the species of Fusarium on morphological grounds, von OVEN has attempted to distinguish this species at least from several disease-producing fusariums by their physiological characteristics. It is thus distinguished from F. Solani, F. putrefaciens, and F. rhizogenum. In cultures on sterilized potato small sclerotia were formed, which produced conidia after being exposed during December and January. The author concludes that this is a hibernating stage of the fungus, although he does not mention finding them in nature—H. HassEL- BRING. Axillary scales of aquatic monocots.—As aquatic monocotyledons are by some held to be modern representatives of the more primitive angiosperms; 4S these forms may have been genetically related to some such type as Isoetes; and as he regards the ligule as an important phylogenetic organ, Grsson?3 has made a study of the vestigial structures of the following families: Potamogetonaceae, 21 WIESNER, J., Untersuchungen iiber den Lichtgenuss der Pflanzen im Yellow- — und in anderen Gegenden Nordamerikas. Sitzungsber. Kaiserl. Akad. Wiss. . Wien, Math.-Naturw. 2 lee II4':(pp. 74.) figs. 2. ; : 22 ne E. von, Ueber eine Fusariumerkrankung der Tomaten. Landw. Jahrb. 34:489-520. pls. 5, 6. fig. I. 1905. 23 Gipson, R. J. Harvey, The axillary scales of aquatic monocotyledons. Jour. Linn. Soc. Bot. 3'7:228-237. pis. 5, 6. 1905. 1906] CURRENT LITERATURE 157 Aponogetonaceae, Juncaginaceae, Alismaceae, Butomaceae, and Hydrochari- daceae. From an investigation of, adult structure and manner of development, he has concluded that the axillary scales found at the bases of the leaves in the plants of these genera are homologous with the more specialized and solitary stipules of Selaginella and Isoetes. It will be recalled that Grsson regards the ligule as a sort of specialized ramentum, protecting and keeping moist the young leaves and growing apex of Selaginella and Isoetes.—FLORENCE Lyon. Reserve food of trees.— NiKLEWSKI?4 confirms by macrochemical methods the observation of Russow and of FiscHeER, that in winter the fat-content of trees first increases and. then dimirishes. The process cannot be reversed by temperature changes. While a rise of temperature accelerates the formation of fat, no change affects its solution. The transformation of fat and of starch are not related. Low temperatures promote the enim: of sugar from starch. Complex phenomena result from a rise of temperature. So great is the loss of reserves by the sedis a that ee seems = ba sone a other than starch or fat share it Ca ‘ Conjugation of yeasts.—GuILLIERMOND’S has extended his studies on the conjugation of yeasts to several additional forms of the Schizosaccharomyces and Zygosaccharomyces. The union of the cells is followed by the fusion of the two nuclei, after which the fusion nucleus divides and the two cells separate or spores are formed in the fusion cell. In some forms conjugation takes place with the germination of the spores. GUILLITERMOND regards this cell and nuclear fusion as a sexual act, but of course chiefly on physiological grounds. Since we do not know the history of the yeasts, it is a matter of speculation whether or not these conjugating cells are phylogenetically gametes.—B. M. Davis. Amphispores in Uredineae.—ArTHuR has given an account of all species of rusts which have amphispores,”° 7. e., as defined by CARLETON, one-celled spores which resemble the teleutospores of Uromyces in appearance, but have two or more germ-pores, and in germination behave like uredospores, their function seeming to be to tide the fungus over unfavorable conditions. This account includes one species of Uromyces and eight of Puccinia, one of which, P. Garrettit, is new. All the forms are American, for thus far no cases of the occurrence of -amphispores have been reported from other parts of the world_—H. HassELBRING. Photosynthesis | extra vitam.—BERNarD has again examined carefully the 24 NIKLEWSKI, B., Untersuchungen iiber die Umwandlung einiger stickstoffreier Reservestoffe wahrend der Winterperiode der Baume. Beihefte Bot. Centralbl. gt: 68— UILLIERMOND, M. A., Recherches sur la germination des . et la con- siti chez les lévures. Rev. Gén. Bot. 1'7:337-376. pls. 6-9. figs. IT. 10905. 26 ArtHuR, J. C., Amphispores of the grass and sedge rusts. Bull. Torr. Bot. Club 32°35-42. figs. g. 1905. 158 BOTANICAL GAZETTE [FEBRUARY question of photosynthesis im vitro, and again with negative results.27_ He repeated Maccuiatr’s experiments (following his directions in litt.), and tried also those of Moutscu, which lent faint support to Maccurati’s conclusions. The gas disengaged seems due only to bacterial infection and when obtained at all does not conform in amount to that demanded by theory. This accumulation of negative results makes exceedingly doubtful the claims of FRIEDEL and MAc- carati.-~-C. K.-B. Measuring transpiration CaNNon describes? a method of studying the rate of transpiration upon plants in place, which he calls the polymeter method, because LaMBRecH?’s portable polymeter, a combined hygrometer and ther- mometer is used to ascertain the increase in humidity of the atmosphere around the experimental plant when enclosed in a bell jar. Certain defects in the method are noted, but the most important one, that it itself produces a variable decrease in transpiration, is not mentioned.— C. R. B. Diastase.— KLEEMANN, finding the known methods of determining the course of diastase formation not sufficiently accurate, proposes a new, and, as he claims, more satisfactory one.?? Using it he has determined that the amount of diastase formed depends, on the one hand, upon the water content of the barley, and on the other, upon how the water is supplied and taken up, and that the loss by respiration is greater the greater the water content.— C. R. B The sporophyte of mosses.—TRUE finds’° that the nodding of the capsusel of Mnium, and probably of Funaria also, is due to geotropic stimulation, while the direction of illumination determines the plane of the curve in the seta, the apex of the capsule sometimes curving toward and sometimes away from the incident light. The calyptra affords important protection to the growing sporo- phyte from mechanical injury and desiccation —C. R. B. Chloroform a stimulant— So Miss Latham: finds it in small quantities to Sterigmatocystis, especially at the time of germination, while larger quantities are inimical or fatal. Less acid formation and less sugar consumption under the stimulus indicate greater metabolic economy.—C. R. B Chromosome reduction.—A useful collective review of the recent literature on this subject is presented by K6rNIcKE in Bot. Zeit. 63?: 289-307. 1905.— 27 BERNARD, C., Sur l’assimilation chlorophyllienne. Beihefte Bot. Centralbl. IQ':59-67. 1905. 28 CANNON, W. A., A new method of pes a the transpiration of plants in place. Bull. Torr. Bot. Club 32: 515-529. 1905. 29 KLEEMANN, A., Untersuchungen iiber ie Landw. Versuchsstat. 63: 93-134- 1905. 3° TRUE, R. H., Notes on =f sanity of the sporophyte of Funaria and Mnium. Beihefte Bot. Centralbl. I9':3 3t LatHam, M. F.., ea Z ae by chloroform. Bull. Torr. Bot. Club 32: 337-357. 1905. NEWS. Dr. Enrico PANTANELLI has been appointed docent in botany at Rome. EMILE Boupier, the eminent mycologist, has been elected director of the Association internationale de géographie botanique for the year 1906. PRorEssor Dr. A. RicHTER has been appointed director of the botanic garden of the University of Kolosvar, the post recently vacated by the death of Professor V. BorsAs. A Portrait of Mr. Francts Darwin was lately presented to the botanical department of the prrsbrnc hs: of Cambridge, where he was for ‘many years.an active investigator and instruct PRoFEssor Huco De Vries ae sail for New York about’ April 1, to deliver an address at the bicentennial anniversary exercises in honor of BENJAMIN FRANK- LIN to be held in Philadelphia April 17-20, under the auspices of the American Philosophical Society. He expects to remain in this country two or three months. Dr. D. T. MacDovaat has resigned his position as assistant director of the New York Botanical Garden and has been appointed director of botanical research of the Carnegie Institute. Dr. B. E. Livincston has resigned his post as physiologist in the Bureau of Soils, U.S. Department of Agriculture, and Professor Francis E. Luoyp his chair in the Teachers College of Columbia University, to accept appointments as investigators on the staff of the Desert Botanical Laboratory, with Drs. CANNoNn and SPALDING. AFTER thirty years’ service Sir W. THIsELtoN-Dyer retired on December 15 from the directorship of the Royal Botanic Gardens, Kew, and was succeeded by Lieutentant-Colonel D. PRAIN, formerly director of the Botanical Survey of India, and superintendent of the Royal Botanic Gardens, Calcutta. Mr. Dyer will remain at Kew till March 31 next, and till that date will continue to act as botanical adviser to the secretary of state for the colonies and as technical adviser in botany to the Board of Agriculture and Fisheries, as well as to take charge of India Office work. From THE Journal of the New York Botanical Garden we learn that Mr. R. S. Wittrams has returned from two years’ explorations of the Luzon, Jolo, and Mindinao, three of the Philippine Islands, bringing large and important collections of herbarium and museum material, estimated at ten to twelve thousand specimens, in spite of the loss of about three months’ collections by fire. R. J. N. Rose with an assistant, Mr. PAINTER, spent the summer in the arid districts of central and southern Mexico, collecting cacti, of which they 159 160 BOTANICAL GAZETTE [FEBRUARY secured several hundred. Special arrangements are being made to study this family thoroughly, both in living and preserved material. me giant bamboos in the palm-house in the past season grew 65 feet (20™) in ninety-five days, an average of about 21°™ per day. ROM advance sheets of the seventeenth annual report of the Missouri Botani- cal Garden, we learn what extraordinary burdens the SHAw bequest has been carrying these sixteen years in the way of taxes, general and special, and real estate and street improvements. This has unhappily delayed the design of Director TRELEASE for development of the Garden as a research center, making impossible the prompt execution of the plan to maintain a staff of specialists and furnish them facilities for work. If the city and state were as just as Mr. SHAW was generous they would relieve the Garden of taxes at least, since it exists solely for the public good. Notwithstanding these unexampled inroads upon its income the institution has not stood still; the garden has not only been main- tained but greatly improved; a fine library and herbarium has been ‘accumulated, and notable researches have been published annually. The grounds now embrace 65 acres, the plant houses cover 30,000 square feet, the cultivated plants number 16,000 species, noteworthy groups being the cacti (678 sp.}, bromeliads (204 sp.), and orchids (942 sp.). The library isnow undoubtedly the best botanical library in the United States, and the herbarium contains over half a million specimens. We congratulate the Director and Trustees on the wise administration of their trust in the face of serious difficulties and discouragements. THE American Mycological Society held its third annual meeting in connec- tion with the American Association for the Advancement of Science at New Orleans, January 1, 1906. In the absence of the president, CHartes H. PEcK, the vice-president, F. S. EarLe, preside e new constitution recommended by the joint committee of the Botanical Society of America, the Society for Plant Morphology and Physiology, and the American Mycological Society, as a basis - for the union of the three societies, was adopted and the present officers . ART reasons for desiring a better classification of the Uredinales; S. M. TRACY, Uredineae of the Gulf States; W. G. FArtow, Some peculiar fungi new to America; F.S. Earte, North American gill fungi; Bruce Finx (by title), Lichens and recent conceptions of species; E. M. FREEMAN, The affinities of the fungus of Lolium temulentum; C. L. SaEar, Peridermium cerebrum Peck, and A Nps Quercuum (Berkeley); C. L. Sear, Romularia: An illustration of the present practice in mycological nomenclature; P H. Roirs, Notes om cultures of Collelotrichum ‘and Gloeosporium; P. SPAULDING, The occurrence of Fusoma parasiticum Tubeuf in this country; P. H. Rotrs, Notes on tes cocos; P. H. Roirs, Penicillium glaucum on pineapple fruit. —C. L. SHE NERVOUS DISORDERS The neryes seh one a es shabagt supply of il dualatas to keep them a and strong. A deficiency ~ the phosphates ve a lowering of nervous tone, indi- ated by exhaustion, eesthiaannan: tog eee or insomnia Hortorts Acid Phosphate (Non- Alcoholic.) furnishes Dae hcg po in a pure and abundan It supplies the phigh cells w ith h toe sivinglile force, repair waste, restores the strength a nd indu ieee restful Ses = reheat the use of i 18% ous drugs.” An Ideal Tonic in Nervou Diseases, If your druggist can’t supply you we will send a tials ye bottle, prepaid, on receipt of 25 cents. Rumford Chemical Works, Providence, R. 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SIyI ‘sRaIIg ge QUE, ‘ooo‘oSz‘1¢ ynoge soo 0} OSvoIYD jo AyisiaAIUp Oy} Jo saysuvapenb ay) ul Surpying Areiqiy & WITYy 0} [eIIoMaM & se yaI0 0} asodoid ‘19}0¥IvYO Poz]exXO SIyY PUL YIOM [Ny -asn sly jo Aroma ay} Yystsoyo 0} Surysim ‘1odsezy AouIey weI[IA JWOptIserg jo spuaisy ay2 ‘sRIIIg gy gun davagns* jeouayF LadaVGE MMIVVE wminyjpEge 34D ‘ODVOIHOD ‘ODV9 -IHOD 4O ALISHZAINN 3HL ‘S3aLSNYL JO GHYVOE AHL JO AYWLEAYHOAS AHL OL LIGNSS ONY 3D031d DNIMO1104 AHL NDIS GNV LNO T1114 OL GALSANOAY 3YuV IVINOWAW SIH OL 3ESINOSENS OL HSIM OHM Y3adYVWH LN3AGIS3Yd 40 SAGNFIY4 SECOND EDITION, ILLUSTRATED Methods in Plant Histology By eae” 98 J. CHAMBERLAIN, a Ph.D. tructor in Botany in the Unibersity of Chic. A Constant Help to Teachers and Students of Botany Contains Directions for Collecting and Preparing Plant Material forMicroscopic estigation T is based upon a course in botanical micro-technique, and is the first complete manual to be published on this subject. 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Cloth. Net $4.00. Postpaid $4,22. Ghe UNIVERSITY of CHICAGO Stace CHICAGO AND NEW YORK VOLUME XLI NUMBER 3 BOTANICAL GAZETTE MARCH, 1906 A MORPHOLOGICAL STUDY OF SARGASSUM FILI- PENDULA. CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. ETOILE B. SIMONS. (WITH PLATES X AND XI) THE family Fucaceae is less understood than its position and prom- inence in the Phaeophyceae warrant. Many important types have scarcely been considered at all, and, moreover, aside from the com- paratively recent cytological studies in the family, few investigations have been conducted with modern methods of technique. The prob- lems of morphology and cytology in the Fucaceae center chiefly around the sexual organs; the peculiar sunken structures in which they are borne, termed conceptacles; the likewise sunken but sterile structures called cryptostomata; and the sporelings. The present investigation of these structures in Sargassum filipen- dula Ag., a member of perhaps the most highly differentiated genus in the Fucaceae, was undertaken with the hope of filling some of the. obvious gaps in our knowledge of this family. It was conducted in the University of Chicago and at the Marine Biological Laboratory, Woods Hole, Massachusetts, under the direction of Professor BRADLEY Moore Davis, who suggested the research to me. It gives me pleasure to express here, both to him and to Professor JoHN MERLE COULTER, my appreciation of valuable suggestions and assistance given me in this work. My acknowledgments are also due the Carnegie Institu- tion for the use of a table at the Marine Biological Laboratory during the summer of 1904. References to anatomical and morphological work which concern 161 162 _ BOTANICAL GAZETTE [MARCH this subject will be given under the topics to which they belong. The early history of the genus with its taxonomic bearing is omitted, as having no place here, but the once credited distribution of Sargassum which was convincingly disproved by Kuntze (’81) is a matter of his- tory which.deserves at least brief mention. Kuntze relates that LINNAEUS believed that a vast area of sea was densely covered by Sargassum in active vegetative condition; Hum- BOLDT reported that the region surpassed Germany in extent six or seven times; Maury stated that it equaled the Mississippi valley; and HOECKEL estimated its area to be forty thousand square miles. That these views were generally accepted is well known. They led to instruction regarding a ‘‘Sargasso Sea,’ whose supposed limits were outlined upon maps of the world. Kuntze, by comparing his own observations and those of other travelers over routes which crossed in different places the outlined area, was able clearly to dis- prove the existence of such a “‘sea.”” Sometimes a voyage was made through the mapped region and little or no Sargassum was seen, and again it appeared somewhat abundantly, but without definite limits or fixed location. Storms which sweep tropical shores, near which attached Sargassum grows abundantly, were found to be in great part accountable for the appearance of the larger quantities of floating Sargassum. Kuntze obtained no evidence to substantiate the view that floating Sargassum vegetates. It had been believed that floating forms of Sargassum consisted of S. bacciferum only, but Kuntze found several species floating, and observed that the specimens in herbaria which had been collected in mid-ocean and labeled Sargassum bacct- ferum according to general belief, could be referred to various species. He therefore concludes that there is no characteristic floating species. The appearance in mid-ocean of floating masses now and then does not seem strange when the authentic distribution and abundance of attached Sargassum are recalled. According to KyELLMAN (’93) this genus, which includes one hundred fifty species, over half the number belonging to the entire family, is found attached along the coast of all warm seas, reaches north to Cape Cod in the Atlantic, to Japan in the Pacific, and in the south into Australian waters, where it is the most abundant. With the extent of this distribution in mind the presence of floating masses, especially after storms, is to be expected. a Te a ea ach al er ae a a DEO eds WoW rh tes 1906] SIMONS—SARGASSUM FILIPENDULA 163 MATERIAL AND METHODS. Material for this study was collected near the shores of Woods Hole, late in July and during August. Plants both in vegetative and in reproductive conditions were abundant. The weak solution of chromacetic acid of Flemming (1 per cent. chromic acid 25°°, 1 per cent. acetic acid 10°°, water 65°°) proved a satisfactory killing and fixing reagent. Microtome sections were cut from paraffin 5 » in thickness and stained either by iron-alum-haematoxylin after the method of Heidenhain or by safranin and gentian violet. The mucil- age on the surface of the plant and in young conceptacles and cryp- tostomata takes the anilin dyes readily, but is not especially trouble- some. GENERAL MORPHOLOGY AND HISTOLOGY. The habit of Sargassum filipendula is so like that of other species which have been described that it needs but slight attention. This species grows attached to rocks below low water mark, and therefore, unlike Fucus and Ascophyllum, is never exposed to the air. Vegeta- tive plants and reproductive plants bearing all stages of conceptacles are plentiful in summer. Sporelings are abundant also and easily collected, for the discharged eggs and their products, the sporelings, remain attached for some time by mucilage to the surface of repro- ductive branches near the parent conceptacles. The stem arises from a small disk-shaped holdfast and passes into long cylindrical branches which bear spirally arranged leaves, berry- like floats, which seem to be modified portions of teaves, as generally stated, and short reproductive branches. This form may attain a height of 60°™, but is commonly shorter. Cryptostomata develop upon stems, leaves, and occasionally also upon reproductive branches in Sargassum, which differs in this respect from Fucus, whose recep- tacles, according to Bower, contain no cryptostomata. KJELLMAN (’93) states that the conceptacles of Sargassum are her- maphrodite. In Sargassum filipendula both mature bisexual and unisexual conceptacles are formed. Some conceptacles contain only spermatocysts (antheridia); some, more rarely, contain many sper- matocysts and but one or two oocysts*(oogonia); and others bear only oocysts. The appearance of a conceptacle devoted to the forma- tion of oocysts differs decidedly from such a structure in Fucus. In 164 BOTANICAL GAZETTE [MARCH Sargassum the oocyst has no stalk cell. It is an embedded organ, being almost surrounded by wall cells of the conceptacle. As both the size and contents of a conceptacle are dependent upon the activity of wall cells (as described later), this conceptacle in Sargassum is smaller and has fewer sexual organs and paraphyses than the corresponding conceptacle in Fucus. The unisexual tendency in the conceptacle of Sargassum may be due in part to the unproductiveness of the many wall cells which abut upon the embedded oocyst. The anatomy of the thallus of Sargassum has been studied in four species. In 1876, REINKE reported its development in Sargassum Boryanum from a three-sided apical cell situated at the bottom of a pit in the apex of the stem. He stated that the holdfast is composed of rhizoids and that a few intercellular filaments occur in the old parts of the thallus. OxtManns (’89) in an anatomical investigation of Sargassum linifolium and S. varians, likewise described a three-sided apical cell, and in addition gave an account of the origin both of the apical cell of a leaf and of a branch. He believes that the branching in Sargassum holds no relation to dichotomy. He figures an enlarged epidermal cell near the apical cell of the stem, and states that it becomes a three-sided apical cell. This young cell develops an out- growth in which a second apical cell is soon differentiated, between the first and the stem. The first formed apical cell develops a leaf and the last a branch. OLTMANNs agrees with Kuntze (’81) that there are all gradations between leaves and floats, and that floats are modified portions of leaves. In 1892, HANSTEEN published the results of an anatomical and physiological investigation of Sargassum bacciferum. He also reported a three-sided apical cell, but did not trace its origin in any structure. He described three kinds of tissues, naming them the assimilating system, the storage system, and the conducting system. The assimi- lating system, according to HANSTEEN, includes only the outer layer of cells, or epidermis. Its cells are twice as long as broad, have undu- lating walls, like the epidermal cells in higher plants, and contain “‘phaeoplasts.” The cells of this system add to their own number by radial, and to the cells below by tangential, divisions. The storage system occupies a zone several cells wide between the assimilating system and the innermost tissue which constitutes the conducting 1906] SIMONS—SARGASSUM FILIPENDULA 165 system. Most of the cells in the storage system are large. HANSTEEN found them empty in alcoholic material of Sargassum, but he did not doubt their function to be that of storage, because he had found much reserve material in similar cells of living Fucus. The conducting system consists of an axial cylinder of long cells with small diameter and oblique end walls. These cells are believed by HANSTEEN and others to function as sieve tubes. The cells of the three aes communicate by pores. HANSTEEN observed in the storage cells of Fucus serratus a several other types, spherical grains of different sizes, which he named fucosan. He believes that the same structures have been variously considered as fat, proteid, and starch by other observers. The grains do not stain blue with iodin, and are soluble in water. HANSTEEN, who made a chemical analysis to determine their composition, con- siders them as a carbohydrate with the formula (C;H,,O,),. CRaTo (°92) described in Chaeto pteris plumosa spherical or elliptical bladder- like structures which he named physodes. He reported (’93) that they contain phloroglucin as a constant ingredient, function in direct- ing the chemical exchange and transportation of food material within the cell, have motion, and are independent cell organs like the nucleus and chromatophore. Crato stated further that HANSTEEN had confused various cell contents, and that fucosan grains and physodes" are the same. KocH (’96) denied the presence of phloroglucin in these bodies. Ina later paper HANSTEEN (:00) again discusses fuco- san grains. He maintains that CRATO’s physodes are fucosan grains, and that they are not independent cell organs but products of the phaeoplast. HANSTEEN has made no further chemical analyses to determine the nature of the bodies, but holds that they surely repre- sent a product of photosynthesis. HANSEN (’95) after an investiga- tion of several forms (Dictyota dichotoma, Taonia alomaria, Haly- seris polypodioides, Asperococcus, Hydroclathrus, and Cystoseira), states that the Phaeophyceae contain oil and no starch, and OLTMANNS (:04) expresses the same view. It is seen therefore, that the character of the reserve material in the cells of the Phaeophyceae is still some- what problematical. Every stem and leaf structure in Sargassum filipendula, as in other species studied, develops through the activities of a three-sided apical 166 BOTANICAL GAZETTE [MARCH cell. The tissue systems described by HANSTEEN are present and each seems to have the function ascribed to it, although without _rigidity. . Each system, too, has its origin in the group of segments surrounding the apical cell and can be traced very near it. The cells of every system are meristematic in the apical region, but the epider- mal cells are apparently the only ones which retain this activity. The cells of any one of the three systems correspond well in general appearance with the similarly placed cells described by HANSTEEN, but an interesting modification was observed in the cells of the con- ducting system. All are long and of small diameter, but in respect to thickness of walls the tissue is differentiated into two regions. The inner cells have thin walls, while the outer ones have thick walls. The thick-walled cells may be both supporting and conducting in func- tion. The conducting system of a leaf blade consists only of thin- walled tissue. No intercellular filaments, as reported by REINKE, have beenfound. Sometimes, however, a filamentous alga creeps into the mucilaginous walls of cells near the surface of a leaf or old stem, and gives the appearance of intercellular filaments. As the little alga contains true starch, its cells when stained with iodin present a sharp contrast to the unstained cells of Sargassum. HANSTEEN _ (792) figures pores in thin areas consisting of the middle lamella in Sargassum baccijerum, and REINKE (’76) represents similar areas but without pores in cell walls of Fucus vesiculosus. Such thin areas are common between cells in the tissues of Sargassum filipendula, but pores, though probably present, are rarely seen. The character of the reserve material in Sargassum proved of great interest. Sections from plants which have been preserved in forma- lin contain much more stored material than tissues which have been kept in alcohol. Preparations, however, which have passed through alcohol, xylol, paraffin, the heat of the bath, etc., still contain within the cells of the epidermis and outer cortex, many bodies which in all probability represent reserve food material. These’ bodies, which stain readily, vary in size and structure, but are evidently related, for transitional stages can be found between the most extreme forms. Judging by the appearance of the structures,some are intact and others modified. Those which seem intact are spherical, with a diameter which equals or exceeds the length of a chromatophore. Each con- 1906] SIMONS—SARGASSUM FILIPENDULA 167 sists of a more or less homogeneous ground substance and one or more refractive areas which are somewhat centrally placed. The modified structures vary from spheres, whose ground substance has been changed only at the periphery, to swollen masses which have an entirely modified ground substance with an irregular outline. Both the intact and modified bodies may occur within the same cell; but the former and the least modified are more common in epidermal cells, whereas the most modified are in cortical cells. The occurrence of such bodies within epidermal cells where photosynthesis is the most active, suggests that they represent a manufactured food. The varied modifications in the structures indicate the solvent action of the killing fluid, or an intercellular enzyme. As the inner cells contain bodies presenting greater modifications than the epidermal cells, the agent producing the change is apparently applied from within the tissue. If then within, it is probably an enzyme, for a solvent used in the process of killing would attack the contents of epidermal cells, doubtless before any others. The intact bodies may represent a newly formed product, perhaps a carbohydrate, and the modified structures, the product in process of digestion. ‘The bodies do not stand with iodin in any con- dition. If they are carbohydrate they probably differ as much or more from the starch of higher plants as does inulin. The presence of many small spheres in formalin material and their absence from tissues preserved in alcohol indicates that oil globules are present in the cell, in addition to the structures described above. Future investigations on living material will probably disclose the presence of both oi and a carbohydrate in the Phaeophyceae. THE ORIGIN AND DEVELOPMENT OF THE CONCEPTACLE,. The conceptacle in the Fucaceae had been but little studied when Bower (’80) gave an account of its development in four genera and six species (Fucus serratus, F. platycarpus, F. vesiculosus, Ozon- thallia nodosa, Halidrys siliquosa, and Himanthalia lorea). Accord- ing to him the development of the conceptacle in every species con- forms to one scheme with minor variations. The “‘initial cell’? of the conceptacle, as stated by Bower, is the terminal cell of a linear series which is produced by a modification of the regular divisions in the segments of the apical cell of a receptacle. 168 BOTANICAL GAZETTE [MARCH This initial cell, strangely, contributes nothing essential to the concep- tacle. It either degenerates directly without having divided at all, or it produces a short filament whose terminal portion degenerates. A cortical cell below the initial is termed by BowER a “‘basal cell.”” This cell and others which adjoin the initial cell laterally, divide and form the walls of the conceptacle from which the sexual organs and paraph- yses arise. The initial cell, therefore, according to BoWER, takes no part in the development of the conceptacle, whereas the cells adjacent to the initial produce all that is important, the walls and their products. The prominent features of this scheme for the development of the con- ‘ceptacle are, it is seen, degeneration of an unimportant initial cell or a part of its filamentous product, and the activity of cells adjacent to the initial in producing the entire conceptacle. Nearly all contributions in this field since 1880 have been in the main confirmatory of the work of BowER. VALIANTE (’83) states that the development of the conceptacle in Cystoseira is due to the growth of neighboring tissue, about one or two cells. OLTMANNS (’89) describes the walls of the conceptacle of Halidrys siliquosa, Himanthalia lorea, and Ascophyllum nodosum, as also formed by neighboring cells, with the one exception that in Ascophyllum the initial cell develops a mass of tissue in the base of the conceptacle. This tissue, he reports, shares with the rest of the inner surface formed from neighboring tis- sue, in developing the sexual organs. As no degeneration of tissue was observed in Ascophyllum, and as its initial cell does contribute some important tissue the development of the conceptacle, this genus presents an exception to a part of the scheme which Bower reports. Although Sphlachnidium should no longer be included in the Fuca- ceae, as shown by the Misses Mirc#ELL and WHITTING (’92), it is of interest to note that these investigators report its conceptacle as devel- oping by the radial division of cells adjacent to a persistent but incon- sequential element, which they believe to be homologous with the initial cell of Bower. GRUBER (’96) states that the conceptacle of Sezro- coccus axillaria is more like that of Halidrys than Ascophyllum, which means, again, that it has an initial cell which contributes nothing of consequence to the conceptacle, whose walls are formed by cells which are adjacent to the initial. - Hotz (:03) reports that in the development of the conceptacle of 1906] SIMONS—SARGASSUM FILIPENDULA 169 Pelvetia jastigiata several epidermal cells cut off basal segments which divide transversely until six or more tiers are formed. Over these tiers, one or more epidermal cells break down and a cavity results, which is gradually enlarged by further disintegration of epidermal and meristematic cells. After a time this process ceases, and a “healthy surface”’ is formed from the deeper meristematic cells. This surface, which comprises the walls of the conceptacle with the exception of the upper part that is formed by “cortical rows”’ of cells, produces sexual organs and paraphyses. The prominent features which distinguish the conceptacle of Pelvetia from others, as thus described, are the presence of several epidermal or initial cells, the more extended disin- tegration of tissue, and a difference in the behavior of the basal cells. The development of the conceptacle in Sargassum filipendula is at variance with all the prominent characteristics in the development of the conceptacle as described by Bower. The initial cell of Sar- gassum does not break down. It is an active cell which produces the entire conceptacle. As the whole conceptacle is the product of this one cell, adjacent cortical tissue takes no part whatever in the devel- opment of the structure. The first indication of the conceptacle is a clearly differentiated epidermal cell which lies near the apical cell of a reproductive branch (jig. 7) and constitutes the initial cell of the con- ceptacle. The upper portion is surrounded laterally by epidermal tissue, whereas its central and basal regions are bounded by cortical. The initial is much larger than any of the cells with which it is in con- tact and differs much from them in shape. Though it may vary some- what in length it is always flask-shaped. Its oval bowl, sometimes slightly narrowed at the base, tapers above into an elongated neck whose outer end is flush with the surface. The initial cell is circular in cross section at its apex (fig. ra) and elliptical at its base (fig. 1b). The initial cell never breaks down. On the contrary the develop- ment of the conceptacle is initiated by its activity. Its large nucleus divides. Then a curved wall is formed with concave surface above, separating two very unlike cells (fig. 2), which form the two-celled stage of the conceptacle. The upper cell, which may be designated the tongue cell, is a long somewhat cylindrical structure; whereas the lower is somewhat conical or wedge-shaped. The initial cell and the two-celled stage of the conceptacle have similar outlines both in longi- 170 " BOTANICAL GAZETTE [MARCH tudinal (figs. r and 2) and in transverse sections (figs. 1a, m, b, and 2a, m, b).. That the lower portion of the tongue cell is surrounded by the upper part of the cell below it is well shown in both longitudinal and transverse sections of the two-celled stage of the conceptacle. The lower cell of this two-celled structure divides longitudinally into two similar daughter elements, thus producing the three-celled stage of the conceptacle (jig. 3). The longitudinal wall reaches to the lower portion of the tongue cell, whose basal portion is surrounded now by two cells instead of by one. _ The relative position of the three cells is made clearer by an examination of their transverse sections. A cross section near the base of the three-celled structure shows two similar cells (fig. 3b). A cross section about midway between the apex and base shows three cells (figs. 3m and 3bm), the tongue cell and the two lower cells which surround its base. A section of the apex is circular in outline and consists of the tongue cell alone (fig. 3a). The three-celled stage of the conceptacle is apparently formed occasionally in another way. The two longitudinal sections of an initial cell are shown in figs. 4, 5, containing three nuclei but no walls. Two nuclei appear in one section and one in the other. It seems that the nucleus of the initial cell in this instance divided first with its spindle perpen- dicular to the axis of the cell, and that one of the daughter nuclei divided with its spindle parallel to the axis. After the three-celled stage, the development of the conceptacle is readily followed. The two lower of the three cells divide longitudi- nally in various planes. A condition thus results which exhibits five cells in longitudinal median section (fig. 6). Four of the five cells are young cells of the recent divisions, and one is the centrally placed tongue cell. Longitudinal divisions continue as before, and a struc- ture showing six or seven cells in longitudinal section is formed (fig. 7)- The tongue cell is still conspicuous in this and in several succeeding stages. Longitudinal divisions continue as illustrated in figs. 8, 9, 11; until the walls of the entire conceptacle are formed. Some of the wall cells begin to develop sexual organs when the conceptacle is very small (figs. 9, 11). - This activity of the cells, however, does not prevent them from contributing to the growth of the conceptacle. The mouth of the conceptacle is surrounded by a marginal ring of epidermal tissue about one or two cells deep (figs. 8, 11). As these ~aihinnbilpleiehin ae > einen ean bp ees Sis bay eS ee 1906] SIMONS—SARGASSUM FILIPENDULA 171 cells are not aggressive they may be omitted from further consider- ation. Excluding this minor detail every portion of the conceptacle is the product of one initial cell. Cortical tissue adjacent to the ini- tial takes no part in its development. The behavior of the tongue cell is similar to that of the “initial cel]”’ in other forms as reported by Bower. It may show signs of degenera- tion (fig. 8), may remain inactive for some time (fig. rr), or may even divide to form a filament of two or three cells (figs. 9, 10). In-no case does it contribute to the walls of the conceptacle, but on the contrary after its divisions resembles a paraphysis. The tongue cell is very : conspicuous until sexual organs begin to develop, but shortly after their appearance it cannot be distinguished. The upper and lower cells which result from the transverse division of the initial cell (fig. 2) cor- respond in appearance and behavior with the “‘initial cell” and “basal cell” as described by Bower and others. It seems probable that Bower saw both the initial cell and the two-celled stage of the concep- tacle, but failing to observe the division in the initial cell, considered the initial and the upper cell of the two-celled stage identical. With this construction, degeneration of the upper or tongue cell was be- lieved to be degeneration of the initial cell itself, and the division of the lower cell of the two-celled stage, a product of the initial cell, was regarded merely as the division of an unrelated cortical cell. A conceptacle developed from cells which happen to be adjacent to a degenerating and unimportant cell would be a very different structure from a conceptacle developed from one active initial cell. | THE ORIGIN AND DEVELOPMENT OF THE CRYPTOSTOMA, The references embodied in the preceding treatment of the concep- tacle constitute the chief source of information bearing upon the cryp- tostoma. The structure which produces the sexual organs has com- monly and naturally been given first attention, but investigators who have studied both, generally agree that the conceptacle and cryp- tostoma are homologous. Different theories regarding the signifi- cance of the cryptostoma have been offered, but no safe generalization can be made until a more extended investigation of both structures has been made in a variety of forms. Miss Barton (’91) gave an account of the cryptostoma in Turbi- 172 BOTANICAL GAZETTE [MARCH naria, stating that an initial cell divides longitudinally, thus forming two daughter elements which produce paraphyses. In demonstration of this two paraphyses are figured arising from the base of a many-celled structure. Miss Barton does not report the origin of the walls of the cryptostoma, but as the initial cell is believed to develop directly into paraphyses, we may assume that she believed the walls to arise from neighboring tissue in accordance with the views of the earlier writers. The development of the cryptostoma in Sargassum follows step by step the history of the conceptacle. The initial cell arises near the apical cell of a leaf or vegetative branch. Longitudinal and cross sections of this cell (figs. 12, 12a, m, b) show the same form and struc- ture as the longitudinal and cross sections of the initial cell of a conceptacle (jigs. 1, 1a, m, b). The activities of the two initials are also identical. The initial cell of the cryptostoma divides transversely, forming a two-celled structure (fig. 13) which is comparable in every way to the two-celled stage of the conceptacle (fig. 2), consisting as it does of a tongue cell and a larger lower cell. The lower cell divides longitudinally. A group of three cells then results (figs. 14, 15) which is entirely similar to the three-celled stage of a conceptacle (fig.3). The two lower cells of this three-celled stage then divide longitudi- nally in one or more planes, forming a structure which shows four or five cells in longitudinal section (figs. 6, 17). The center of this structure and of several which follow is occupied by the conspicuous tongue cell (figs. 16, 17, 18, 19). Thus by the continued longitudinal divisions of the products of the lower cell of the two-celled stage, the walls of the entire structure are gradually developed. Paraphyses begin to appear in the cryptostoma (jigs. 18, 20) as early as do the sexual organs in the conceptacle (fig. g). The activity of the wall cells in producing paraphyses, however, does not interfere with their functioning further in developing the cryptostoma (fig. 21). Enpider- mal cells at the mouth of the cryptostoma form here, as in the concep- tacle, a marginal ring one or two cells deep (compare figs. 8 and 19). The origin of the true walls of the structure, however, may be traced as in the conceptacle to the lower cell resulting from the transverse division of the initial. The behavior of the tongue cell in the cryptostoma is similar to SC 1906] SIMONS—SARGASSUM FILIPENDULA 173 that of the corresponding element in the conceptacle. Occasionally the tongue cell of the-cryptostoma may develop a prominent filament (fig. 20), which is clearly identical in structure with a typical paraph- ysis (fig. 21). The young conceptacle and cryptostoma are so alike that they can only be distinguished by their respective positions on fruiting branches or on young vegetative structures, until the appear- ance of sexual organs in the one and paraphyses in the other defines their mature characters. The development of the paraphysis is interesting for its regularity. A wall cell enlarges, pushing into the cavity of the cryptostoma, and then divides transversely (figs. 18, 20). The upper cell produces the paraphysis, whereas the lower functions in the development of the wall. The growth of the paraphysis results from the transverse divi- sions of the cell next the wall (figs. 21, 22), a method of growth termed trichothallic. ‘The development of a paraphysis in the cryptostoma of Sargassum is, therefore, characteristically basipetal, as BARTON (’91) found in Turbinaria. ; A somewhat advanced paraphysis is composed of three regions. That which adjoins the wall of the cryptostoma consists of the large turgescent meristematically active basal cell (fig. 22). The middle region is occupied by six or eight short cells which have so recently been formed that they have not had time to lengthen much. The upper region contains several greatly elongated cells. This region in a mature paraphysis is partly within the cryptostoma and partly without, for fully developed paraphyses extend far beyond the sur- face of the plant. A peculiar condition found in many cryptostomata deserves special attention. Structures frequently appear between the paraphyses which seem to bear no relation to them. These are papillae and stalked cells, the former like the papillae which precede male organs in a conceptacle and the latter like the male organs themselves. The stalked cells, although slender and probably always sterile, appear to be spermatocysts no longer functional. This surprising condition is of great interest and importance in relation to the homology and sig- nificance of the cryptostoma, a structure formerly believed to contain only paraphyses, but which appears also to have sexual organs or their degenerate representatives. That the cryptostoma and conceptacle 174 BOTANICAL GAZETTE [MARCH are homologous cannot be doubted, since: their origin and early development are identical in all details. The occasional appearance of sterile representatives of sexual organs within the cryptostoma further confirms this view of their relationship and strongly supports the theory of Bower (’80) that the cryptostoma in the Fucaceae is derived from the conceptacle. The occurrence of conceptacles in special reproductive branches only, the appearance of cryptostomata in both vegetative and repro- ductive branches, and the development of representatives of sexual organs within the cryptostomata, suggest a line of evolution from plants bearing conceptacles scattered over leaf and branch struc- tures indiscriminately, to the type now under consideration with localization of the conceptacle upon special branches. Certain branches were set apart to bear conceptacles as the conceptacles in all other parts of the plant body were rendered sterile and thus changed into cryptomostata. The presence of sexual organs or their degenerate representatives within a cryptostoma indicates, according to these views, that the process is not carried to its farthest point in Sargassum. The production of conceptacles upon small special branches only, instead of upon the entire plant, naturally results in fewer concep- tacles upon one plant. The conceptacles, however, are much more closely placed than the cryptostomata. On account of their com- paratively small size the initials and young conceptacles occupy very little space in the apex of a branch, but farther down on the receptacle the bulging bowls of the developing flask-shaped conceptacles require more and more space, until the mature structures nearly fill the interior of the receptacle and there is only enough intervening tissue to hold the conceptacles together. The cryptostomata, on the other hand, are well scattered upon vegetative branches and mature leaves. The contrast in the placement of cryptostomata with that of concep- tacles is, therefore, very marked. THE SPERMATOCYST. The male sexual organs (antheridia), which will be called sper- matocysts in this paper, according to the terminology of DAvis (:04), develop from wall cells of the conceptacle in Sargassum as in other forms of the Fucaceae. A wall cell puts forth a papilla (fig. 23) which I a a i ti I 2 1906] SIMONS—SARGASSUM FILIPENDULA cy gs is cut off by a transverse wall (jig. 24). The lower cell becomes a part of the wall occupying the place of the cell from which it arose. The upper cell enlarges for a time and then divides, forming the sperm mother-cell or spermatocyst and its stalk (jig. 24, at the right.) A stalk cell may have no other relation than that which it bears to the spermatocyst which it supports, or it may function in other ways. It may produce several spermatocysts directly, without individual stalks; it may put forth a papilla which gives rise to a spermatocyst and stalk (figs. 25, 26); or it may develop a hair (fig. 27). Hairs, however, are comparatively rare within a conceptacle of Sargassum. Owing to the variety of activities which belong to a stalk cell, the growths within a conceptacle lack uniformity. Some structures reach but a little distance above the wall of the conceptacle, whereas others form conspicuous branch systems. Although these systems are prom- inent in this conceptacle, they are considerably smaller and less dense than the branch systems in a conceptacle of Fucus, and there is far more unoccupied space within the cavity of a conceptacle of Sargassum than of Fucus. The young spermatocyst contains dense cytoplasm, a centrally placed nucleus and deeply staining granules, the nucleus remaining in a resting condition for a long period. The divisions of the nucleus were not studied in detail. Sixty-four sperms are apparently formed (figs. 27 and 28), agreeing, therefore, with the count announced by BEHRENS (’86) for Fucus vesiculosus. The sperms within the spermatocyst have an elliptical outline, a cytoplasmic ground mass, and a somewhat spirally arranged band, which is probably the nucleus. The discharge of sperms was not seen, but a rent, partly terminal and partly lateral in empty spermatocysts, indicates their mode of escape. THE OOCYST. The female sexual organ (oogonium) or oocyst, according to the _ terminology of Davis (:04), is peculiar among the Fucaceae, as far as is known, in that it is not borne upon a stalk but is a partially embed- _ded organ (fig. 31). The sister cell of the oocyst, instead of develop- ing into a pedicel cell as is usual in this family, functions as one of the wall cells of the conceptacle. The oocyst enlarges greatly, but becomes nearly surrounded by adjacent wall cells. 176 BOTANICAL GAZETTE [MARCH Its development is simple. A somewhat enlarged wall cell of a young conceptacle divides transversely, forming two cells much alike in size and contents (jig. 29). The inner cell, which is the homologue of the stalk cell of the female organ in Fucus, cannot be distinguished from neighboring wall cells shortly after its formation. The outer cell, which has a free surface toward the interior of the conceptacle, increases greatly in size and soon becomes the spherical oocyst. Fig. 30 represents a young oocyst and its sister cell, already unequal in size. There now follows a long period of growth, during which the oocyst attains a remarkable size, finally containing a great quantity of reserve material, many chromatophores, much cytoplasm, and a large nucleus. The mature organ, drawn under a lower magnification than fig. 30, is represented in fig. 31. No trace of its sister cell could be found. The oocyst of Sargassum develops but one egg. The mitosis within the wall cell whose division produces the oocyst is normally the only mitosis in the process of oogenesis. Particular attention was given to this point. The one nucleus of the oocyst remains in a resting condition throughout the entire period of the growth of the cell, and therefore becomes the nucleus of the egg. In the other genera of the Fucaceae, as is well known, there are three mitoses within the oocyst, resulting in eight nuclei. Each of the eight nuclei may become a center for the development of an egg as in Fucus, or some nuclei may degenerate and a less number of eggs be formed, as in Ascophyllum and Pelvetia. It might be supposed from these conditions in the Fucaceae that the oocyst of Sargassum would show similar nuclear divisions and degeneration, but this is not the case. The mitoses characteristic of oogenesis in Fucus are normally suppressed in Sar- gassum. The tendency in the Fucaceae to reduce the number of eggs produced by an oocyst reaches its culmination, therefore, in Sargassum. It is interesting to note that Sargassum still gives proof that it be- longs to the reduction series which has its beginning in Fucus and allied forms that produce eight eggs in an oocyst. Out of the great number of conceptacles examined, one oocyst was formed which con- tained two eggs, and two oocysts which contained eight. The oocyst with two eggs was formed in an immature conceptacle that held five normal oocysts. The two eggs appeared fairly vigorous. One of the 1906] © SIMONS—SARGASSUM FILIPENDULA 177 two oocysts which contained eight eggs was an old conceptacle, from which other sexual elements had apparently long been discharged. The eight together were smaller than one normal mature egg. The other oocyst which contained eight eggs shared a conceptacle with two normal oocysts. It was attached in the side of a conceptacle near the surface of the plant, which for a slight distance was modified in struc- ture as if in response to an injury. — It is possible in this case that the wound incited the reversion.. The appearance of an oocyst contain- ing more than one egg in Sargassum must be regarded as a rare rever- sion to the Fucus type. The resting nucleus of the oocyst is always large, but varies in structure. Sometimes it has few granules and no conspicuous reticu- lum, whereas at other times it contains many. granules and a dense network. The nucleolus is also large in size and variable in structure. At the present time no suggestion can be made to account for the changes in nuclear structure, excepting that they are the concomitants of growth and varying nutritive conditions. The method of discharge of the egg from the conceptacle of Sar- gassum is somewhat unlike that reported in Fucus and other genera. In Fucus the outer membrane of the oocyst remains attached to the conceptacle, as explained by THuRET, and the eggs escape in a group surrounded by a very delicate inner membrane. In Sargassum the entire oocyst becomes freed from the conceptacle and escapes. In Fucus the inner membrane dissolves or breaks, thereby freeing the naked eggs which it has enclosed. In Sargassum the wall of the oocyst ‘swells, stretches, and sometimes ruptures, but it may persist for a long time, even enveloping later a many-celled sporeling formed within it. The inner membrane enclosing the eggs of Pelvetia is separated from the outer as in Fucus. In Pelvetia, however, as figured by THURET, this membrane persists about the eggs, apparently offering no great resistance to the entrance of sperms. Whether the sperm enters the egg of Sargassum through a break in the oocyst membrane, whether it passes through the membrane, or whether the eggs develop par- thenogenetically, isnot known. A study of fertilization in Sargassum is surrounded by serious technical difficulties because both eggs and sperms develop upon the same plant, thus making it difficult to isolate the sexual cell. 178 BOTANICAL GAZETTE [MARCH THE SPORELING. Many if not all of the eggs of Sargassum on leaving the conceptacle become fastened immediately by the mucilaginous wall of the oocyst, which still surrounds it, to the surface of the reproductive branch. In this position the eggs segment. ‘The first division of the egg in Sar- gassum does not differentiate a rhizoidal region, as in Fucus and Ascophyllum. Instead, a many-celled ellipsoidal structure is formed, the divisions occurring with mathematical precision. Rhizoids then develop at one end with no apparent relation to a substratum or to gravity, so far as could be observed in fixed material. Sporelings sometimes occur at opposite sides of a branch with rhizoids directed toward the stem, thus showing no relation in the development of rhi- zoids to gravity; and again, sporelings occur with rhizoids directed away from the branch in various directions, indicating that the parent plant exerts no special influence. It is possible that the attachment of a sporeling upon a plant is so insecure that the direction of its axis may be shifted in the manipulation of material. Otherwise it is difficult to account for the conditions which were observed. When the many-celled sporeling has reached the condition for rhi- zoid formation the cells at one pole elongate, thereby giving rise to a tuft of rhizoids of approximately equal length. This mass of rhizoidal filaments finally produces the characteristic disk-shaped holdfast of the mature plant. Fig. 32 shows a sporeling in about the oldest con- dition in which it remains attached to the parent plant. No apical cells were found in these sporelings and therefore its differentiation must occur after the sporeling has separated from the parent plant. The germination of the oospore deserves careful cytological investi- gation. Many preparations have been made and studied, but further attention will be given the subject before the observations are pub- lished. A few conditions may be noted, however. There are numer- ous radiations at the poles of the early spindles. The asters contain granular inclusions which suggest centrosomes, although their origin and relation to the processes of mitosis have not been traced. Walls following the mitoses are developed somewhat slowly, being formed in part at least by the membranes of contiguous vacuoles. 1906] SIMONS—SARGASSUM FILIPENDULA 179 SUMMARY. Each stem, branch, and leaf structure develops through sic activi- ties of a three-sided apical cell. The thallus consists of three compact tissues, called for conven- ience the epidermal, cortical, and conducting tissues. The latter con- sists of only thin-walled cells in the leaves, but in mature stems con- tains both thick and thin-walled elements. A ring of thick-walled cells, which may have both a mechanical and conducting function, surrounds the thin-walled conducting cells in the center of the axis. The tissues normally contain much reserve material, a part of which is oil, and a part, whose nature is undetermined, appears to be a carbohydrate. Both the conceptacles and cryptostomata originate in a single flask- shaped initial cell which develops the entire structure. The first division of the initial cell results in two unlike segments: a large lower cell which develops the walls of the conceptacle and cryp- tostoma; and an upper cell, the tongue cell, which either remains inactive, divides to form a short filament, or degenerates. The “‘initial cell” of Bower is apparently the tongue cell, a product of the true initial cell. The conceptacle and cryptostoma are undoubtedly homologous structures. Every stage of development in both structures is the same, from the appearance of the similar initial cells to the develop- ment of paraphyses in the cryptostomata and sexual organs in the conceptacle. The paraphyses are developed sag i by the division of the lowermost cell in each structure. Spermatocysts or their degenerate representatives occur in some cryptostomata. Such conditions indicate that the cryptostomata have been derived from conceptacles whose sexual organs have become sterile. The spermatocysts develop as in other Fucaceae, each finally pro- ducing sixty-four sperms which are discharged from a partly terminal and partly lateral rent. The sister cell of an oocyst does not become a stalk and conse- quently the oocyst is an embedded structure. 180 BOTANICAL GAZETTE [MARCH The oocyst normally gives rise to but one egg. The nucleus of the oocyst accordingly becomes the nucleus of the egg. | The oocysts were found containing eight eggs each. These must be considered a rare reversion to the Fucus type. The entire oocyst of Sargassum, unlike other genera of the Fuca- ceae which have been studied, is discharged with its enclosed egg. The oocyst wall may break, partially freeing the egg, or it may persist even enveloping a many-celled sporeling. Segmentation of the egg takes place while it is attached to the sur- face of the plant by the mucilaginous wall which surrounds it. This segmentation results first in a many-celled undifferentiated ellipsoidal sporeling. Rhizoids develop late at one end of the multicellular spore- ling, with no apparent relation to gravity or other stimulus. Asters, containing granular inclusions suggesting centrosomes, appear at the poles of the spindles in the early mitoses of the segmenta- tion of the egg. THE UNIVERSITY OF CHICAGO. LITERATURE CITED. Barton, E. S., ’91, A systematic and structural account of the genus Turbinaria Lamx. ‘Piers: Linn. Soc. Bot. 3: 215-226. pls. 54-55. BEHRENS, J., ’86, Beitrag zur Kenntniss der eg ia Rarer bei Fucus tices: lige Deutsch. Bot. Gesells. 4:92-10 Bower, F. O., ’80, On the development of the sciceplecks in the Fucaceae. Quart. Jour. Micr. Sci. 20: 36-49. pl. 5. Crato, E., ’92, Die Physode, ein Organ des Zellenliebes. Ber. Deutsch. Bot. Gesells. 10:295-302. pl. 18. Davis, B. M., :04, The relationships of sexual organs in plants. Bot. GAZETTE GruBer, E., 96, idee Aufbau und Entwickelung einiger Fucaceen. Bibli- otheca Bot. 38:3 HANSEN, A., ’95, Usher Stoffbildung bei den Meeresalgen. Mittheil. Zool. Sta. Neapel rz: 255-305. pl. 12. HANSTEEN, B., ’92, Studien zur Anatomie und Physiologie der Fucoideen. Jahrb. Wiss. Bot. 24: 317-360. pls. 7-10. ,:00, Ueber das Fucosan als erstes scheinbares Product der Kohlensaure- assimilation bei den Fucoideen. Jahrb. Wiss. Bot. 35:611-625 pl. 14. Hotz, F. L., :03, Observations on Pelvetia. Minn. Bot. Studies 3:23-45- pls. 7-12. KJELLMAN, F., ’93, Engler and Prantl, Pfl. fam. I. 2: 268. PLATE X ~ ~ be Sa Ny hy Ry ~ S = 1S) ~ q & 2) Q 1 2 Etoile B. Simons, del. * sIMONS # SARGASSUM 1906] SIMONS—SARGASSUM FILIPENDULA r81 Kocu, L., ’96, Untersuchungen iiber die bisher fiir Oel oder Phloroglucin gehaltenen Inhaltskérper der Fucaceen. Inaug. Diss. Rostock. Kuntze, O., ’81, Revision von ae und das sogenannte Sargasso-Meer. Engler’s Bot. Jahrb. 1: 191-239. pls..1-2. MitcHett, M. O., and Wuarrtrnc, F. G., ne On Splachnidium rugosum Grev., the type of a new order of algae. Phycological Memoirs Part I. pp. 1-10. Learn F., ’89, Beitriige zur Kenntniss der Fucaceen. Cassel. Meaghleige und Biologie der Algen. Jena a4 L 76, Beitrage zur Kenntniss der Tange. Jahrb. Wiss. Bot. 10:317- 382. pls. 25-27. VALIANTE, R., ’83, Le Cystoseirae del Golfo di Napoli. Fauna und Flora Golfes Neapel '7: 1-30. pls. 15. EXPLANATION OF PLATES X AND XI. All figures were sketched with a camera and reduced one third in reproduc- tion. Figs. r-30 were drawn with Zeiss apochromatic objective 1.5™™ and com- pensating ocular number 4, magnification 1140. Figs. 31 and 32 were drawn with dry objective, magnification 570. PLATE X. Figures 1-11. Development of the conceptactle. Fic. 1. Initial cells, longitudinal section; 1a, cross section of apex; 1m, cross section of median portion; 1b, cross section of basal portion. Fic. 2. Two-celled stage, longitudinal section showing the slender upper tongue cell, and a larger lower cell; 2a, cross section of the apex showing the tongue cell only; 2m, cross section of the median portion with the centrally placed basal region of the tongue cell surrounded by the upper part of the lower cell; 26, cross section of the basal portion showing the lower cell only. : Fic. 3. Three-celled stage, longitudinal section; 3a, cross section of the apex showing tongue cell only; 3m, cross section of median portion with the cen- trally placed lower part of the tongue cell, surrounded by the upper part of its two companion cells; 3b, cross section a little below 3m, showing the same cells; 3, cross section of the basal portion showing the two lower cells only. Fics. 4 and 5. Longitudinal sections of a peculiar trinucleate stage of one conceptacle. The first division of the nucleus of the initial cell must have been with the axis of the spindle perpendicular to that of the cell. Fig. 5 contains one of the nuclei of the first mitosis and fig. 4 the products of a division, now in late telophase, of the other nucleus of the first mitosis. Pb o = Oo ag MW MW walls Celi ~~ the tongue cell. - tudinal section of a later stage with six similar wall cells and the Fic. 7. Longitu centrally placed tongue cell. 182 BOTANICAL GAZETTE [MARCH Fic. 8. Longitudinal section of a more advanced stage illustrating the forma- tion of the cavity of the conceptacle. IG. 9. Pesca aes section of a young conceptacle some of whose wall cells are ie papi he tongue cell contains two nuclei. Fic. 10. A aaa of three cells formed from the tongue cell. Fic. 11. Young conceptacle showing simultaneous development of wall cells and papillae. Figures 12-22. Development of the cryptostoma. Fic. 12. Initials, longitudinal section; 12a, cross section of the apex; 12m, cross section of median portion; 126, cross section of basal portion. Fic. 13. Longitudinal section of the two-celled stage. Fic. 14. Longitudinal section of the lateral surface of the three-celled stage. Fic. 15. Longitudinal section of the interior of the same group of cells repre- sented in fig. 14. Fic. 16. Longitudinal section showing four wall cells and the tongue cell. Fic. 17. Longitudinal section slightly more advanced. Fic. 18. Longitudinal section of a young cryptostoma beginning to form paraphyses very early. Fic. 19. Longitudinal section of an older stage which has not yet begun to develop og aa Fic i Loaditadinal section showing five paraphyses developing from wall cells sia one from the ton e Fic. 21. More ‘vaiee illustrating the simultaneous development of paraphyses and wall ce Fic. 22. Still more iluaspach PLATE XI. Fig. 23. The development of papillae which will later give rise to spermato- sts. Fic. 24. At the left a cell which results from the separation of a papilla from a wall cell. At the right a spermatocyst and stalk which have been formed by the division of a cell similar to the one shown at the left. Fic, 25. A stalk cell has given rise to a papilla, now separated by a wall. Fic. 26. A branch system formed through the activity of stalk cells. Ah Fic. 27. A spermatocyst containing sperms. The stalk cell has developed a air. Fic. 28. A mature spermatocyst, the stalk cell pushing out at one side. Fic. 29. Very young oocyst with its sister cell, which is the homologue of the stalk cell in Fucus FIG. 30. Slightly older oocyst and its sister cell already unequal in size Fic. 31. A mature embedded oocyst containing many hnshatophines and much reserve material. Fic. 32. A sporeling still attached to the surface of the parent plant. At one pole rhizoids have begun to develop. The old wall of the oocyst surrounds the sporeling. CHROMOSOME REDUCTION IN THE MICROSPORO- CYTES OF LILIUM TIGRINUM.? Joun H. SCHAFFNER. (WITH PLATES XII AND XIII) THE progress recently made in our knowledge of hybrids has given a new impetus to the study of chromosome reduction. Unfortunately, there is still much disagreement in the accounts of various observers. In order to continue my own investigations on a very favorable object, the microsporocytes of Lilium tigrinum were selected, since material is easily obtained in large quantities. CHAMBERLAIN has studied the pollen grain of this plant and has also given figures of the microsporo- cyte in the spirem stage. The chromatin granules are exceptionally distinct, and this facilitates the correct interpretation of the complex figures to be seen in the reduction karyokinesis. Recently papers on the reduction division have been published by FARMER and Moore, STRASBURGER, MONTGOMERY, WALLACE, and others, which are in essential agreement with the interpretations of Drxon on Lilium longiflorum and my own observations on Lilium Philadelphicum and Erythronium. On the other hand BErcus, REGOIRE, and ALLEN have come to somewhat different conclusions. The observations of RosENBERG on Drosera have opened up an important field of investigation on the individuality of the chromo- some. These papers have been reviewed so recently by various writers that it is needless to discuss the results here. It is sufficient to say that it must appear to an impartial judge that the cytologist is at present able to see in his preparations almost anything which may be conceived of as taking place in the structures investigated. This, however, should not hinder work in such an important field, for the proper interpretation can be attained only by continued observation. Little can be regarded as certain until there is a more general agreement among competent investigators. So far as the present research is concerned, the extent and variety of the preparations ‘Contributions from the Botanical Laboratory of Ohio State University. XXIV. 183] [Botanical Gazette, vol. 41 184 BOTANICAL GAZETTE [MARCH studied seem to preclude the possibility of mistake. Such doubts as were expressed in my former papers have been practically removed. So far as the writer is concerned, the interpretation given below is conclusive. Another investigator might perhaps come to different conclusions by using other methods. MATERIALS AND METHODS. Stamens of various ages were collected in Clay County, Kansas, during July 1904; killed in weak chrom-acetic acid (chromic acid 0.3%”, glacial acetic acid 0.7°°, water 99°‘), passed gradually through the grades and preserved in 70 per cent. alcohol, imbedded in paraffin, cut 10-18 » thick, and stained on the slide. Some old slides, the mate- rial of which had been killed in the ordinary chrom-acetic acid solu- _ tion, were also at hand. Various stains and combinations were used, but for bringing out the chromatin network and chromatin granules of the early stages, Delafield’s haematoxylin, when properly developed, gave by far the best results, being superior in this respect to either Heidenhain’s iron-alum-haematoxylin or safranin and gentian violet. Nucleoli, both in the nuclear cavity and in the cytoplasm, are stained very distinctly by the safranin-gentian-violet combination, but are only slightly affected by Delafield’s haematoxylin. _ I am indebted to my wife, MABEL ScHAFFNER, for the prepara- tion of most of the two hundred serial slides on which the present paper is based. INVESTIGATION. Before the microsporocytes are beginning to separate the promi- nent chromatin network is being transformed into slender delicate threads. These threads appear to be discontinuous in some places. However, the appearance may be due to injury during the process of cutting. The threads are small in diameter as compared with the single chain of spherical chromatin granules (figs. 1-3). After the spirem becomes fully developed it shows no free ends and is much wound, looped, and twisted. In this stage it appears to be entirely free in the nuclear cavity and is usually in the so-called synapsis stage. Sometimes the contraction is to one side of the nuclear cavity, some- times near the center, but often little or no contraction is evident. Whether this is an artifact or a real stage in the process of karyokin- = Y? — 1906} SCHAFFNER—LILIUM TIGRINUM 185 esis appears still to be an open question. Such contractions are so easily produced by the ordinary killing reagents used, and have been described for such a variety of the early stages of nuclear division, that it seems to me no importance is to be attached to observations which have been made on killed material. So far as any opinion is to be expressed upon the appearances in Lilium tigrinum, I am still convinced that the extraordinary distortions commonly figured owe their origin to the action of the fluids used before imbedding in par- affin. : The spirem remains simple during the entire synapsis. The linin thread becomes thickened and the chromatin granules are usually more or less elongated (figs. 4, 5). At this stage the spirem has already a strong tendency to be thrown into loops and coils (fig. 4). After the microsporocytes have become partly separated and more spherical, they were rarely observed to be in synapsis. The nuclear cavity enlarges and the chromatin ribbon becomes thicker, with the granules still more prominent (jigs. 6-8). This is an important point to consider, for we have here a clear case of sporocytes, long past the supposed synapsis stage, showing with remarkable clearness a simple continuous spirem, with a single chain of chromatin granules. Synapsis, therefore, can have nothing to do in this case with a sup- posed longitudinal conjugation of two spirems or two networks of chromatin before the spirem is formed. After the nuclei have passed on to the stage represented in fig. 6, the stages are so easily followed and the threads so prominent, that a longitudinal conjugation, if one occurred, could not escape notice. Shortly after the stage shown in jig. 6 the spirem begins to show double rows of elongated chromatin granules, but the relative quantity of chromatin ribbon present in the nuclear cavity is not much diminished (figs. 9, 10). If the amount of spirem were diminished one-half by a longitudinal conjugation the fact would certainly be noticeable. Often a part of the spirem appears still single, or it will appear with a double row of granules and gradu- ally change to a row apparently single (figs. 11-17). The appearance would be the same whether the granules were dividing or conjugating. The uniformity of the pairs of granules on the linin thread is remark- able, and the pairs themselves suggest a division. If a conjugation of the chromatin granules were established, it would certainly show 186 BOTANICAL GAZETTE : [MARCH an almost inconceivable regularity. The granules would have to be definite and fixed, of the same number in both the egg and sperm, and not be subject to increase or diminution (jigs. 14-16). In the early stages of the spirem there are, of course, numerous instances of threads lying parallel, side by side, but these appearances are equally common after the chromatin granules appear double (jigs. 15, 17). The ribbon now begins to show an arrangement into definite loops (fig. 10). It becomes much shorter and thicker (jigs. 18, 19) and finally shows a definite twisting together into twelve loops, which have their heads toward the nuclear wall (jig. 20). At this stage the chromatin - granules can still be distinguished lying side by side (fig. 21). But at this time the whole ribbon begins to undergo a change, so that it stains of a uniform, dense color throughout, and before the loops separate all evidence of chromatin granules is lost (figs. 22, 23). That the loops shown in figs. 18-23 are the incipient chromosomes is self-evident. By no manner of interpretation can such a conclusion be explained away. By the time the twelve loops have separated, the nucleoli have entirely disappeared from the nuclear cavity (fig. 24). The nucle- oli break up into micronucleoli and are thrown out into the cytoplasm (figs. 38, 51). The figures in which they do not appear were taken from material stained in such a way that the nucleoli were not evident or not very distinct. The chromosomes are exceedingly interesting on account of the | many fantastic shapes produced by the coiling of the ribbon. A series of distinct shapes is given in jigs. 25-37. Occasionally the loop shows evidence of its double nature (jig. 29), but usually it appears homo- geneous throughout. Much time was spent in a study of these chro- mosomes, and the variety of shape and coil could be extended indefi- nitely. In all cases the chromosome is continuous, the outer end of the loop always being closed. Occasionally the coiling takes place in such a way as to form a double loop (figs. 26, 27, 35). The chro- mosome is situated on the spindle with its head or closed end outward (figs. 28-30). Sometimes it is very difficult to unravel the nature of the coil, as in fig. 35. In other chromosomes it is an easy matter to follow out the details of the loop, as in fig. 28. The spindle threads are evidently attached some distance back of the free limbs of the loop (figs. 25-28). The limbs are gradually = Pee Le | Oa . 1906] SCHAFFNER—LILIUM TIGRINUM 187 uncoiled and pulled apart. They are separated at the closed outer end. A transverse splitting of the chromosome is thus accomplished. There is no evidence that the two limbs of the chromosome are a male and female chromosome joined end to end and twisted together until they make a longitudinal pair. But theoretically such a proposition is easily possible or even probable. Because the spindle threads are attached some distance from the ends of the limbs, the daughter chro- mosomes are developed as V- or U-shaped loops (jigs. 40, 43, 51).- In favorably stained sections some evidence of chromatin granules may be observed. These still show a distinct pairing in some cases (jig. 44), but in others the arrangement is considerably disturbed (jig. 45). The daughter chromosomes, as they appear in the daughter star, are of various forms and are sometimes twisted (figs. 44-50). The micronucleoli are gradually collected below the two daughter skeins, and are finally all inclosed in the nuclear cavities of the daugh- ter nuclei (figs. 52, 53). The daughter chromosomes do not appear to form a very definite resting network, but are transformed into the mother skein of the second division rather rapidly. Whether a con- tinuous ribbon is formed was not ascertained. The loops are already separate at an early stage, and it is possible that the daughter chro- mosomes of the first division, after forming an imperfect reticulum, break up directly into the twelve chromosomes of the second division (fg. 54). This point, however, is doubtful. But the absence of a defi- nite resting stage in sections having microsporocytes with loose daugh- ter skeins in close proximity to loose mother skeins of the second division, gives support to the above supposition. The micronucleoli are again distributed in the cytoplasm before the mother-star stage of the second division (figs. 54, 55). The karyokinetic figures of the second division are easily distinguishable from those of the first. This is especially true of the mother star (figs. 38, 55). The chro- mosomes in the second division have their free ends directed out- wards, as in an ordinary vegetative division. Commonly they are more or less tangential to the spindle. Sometimes, however, they Stand at right angles, as represented in jigs. 56, 57. One of the most difficult points to determine was the nature of the splitting in the second division. However, it was definitely ascer- tained that the splitting is longitudinal. Dividing chromosomes are 188 BOTANICAL GAZETTE [MARCH represented in jigs. 58-61. The daughter loops are completely sepa- rated very early in their migration towards the poles (jig. 62). They are also of various shapes. Frequently straight chromosomes are present in the daughter star and very commonly the J shape pre- dominates (figs. 63,64). Uand V shapes are also present. After the chromosomes have passed into the daughter skein stage they show an irregular outline with a row of irregular chromatin granules (jigs. 65, 66). The chromosomes develop into irregular networks, show- ing the remains of the original loops, and thus pass into the spore tetrad stage (jig. 67). During the early germination stages of the microspores the figures are again remarkable for the large nucleoli in the cytoplasm (jigs. 68-70). From the appearance of the nuclei during the several divisions, it is evident that the nucleoli do not contribute directly to the formation of the chromosomes, but that they are uniformly thrown _ out into the cytoplasm, in a fragmented condition, during the earlier stages of karyokinesis. No study of the achromatic structures was attempted, and though some interesting points were observed from time to time, there was nothing which calls for special mention. SUMMARY. 1. The first division of the microsporocyte is a true reduction divi- sion. 2. A continuous spirem is formed with a single row of chromatin granules. 3. The spirem passes through and comes out of synapsis without a conjugation or division of chromatin granules. 4. The chromatin granules divide but the linin thread does not show a distinct separation. 5- The continuous spirem shortens and thickens and twists up into twelve loops, which are the incepts of the twelve separate chro- mosomes. 6. The chromosomes are arranged in the mother star with the loop or head end turned outwards and the spindle threads are attached near the ends of the free limbs or about half way between the free ends and the head. pe | —— ieee Se cenilaiiiaiiiea is 1906] SCHAFFNER—LILIUM TIGRINUM 189 7. During the metakinesis stage the chromosomes uncoil and separate by a transverse division at the middle. 8. The chromosomes of the second division appear to represent the daughter chromosomes of the first division. g. The division of the chromosomes in the second nuclear divi- sion is longitudinal. 10. The nucleoli fragment and pass out into the cytoplasm during the first and second divisions and also during the germination of the microspore. OHIO STATE UNIVERSITY, Columbus. LITERATURE. ALLEN, C. E., Nuclear division in the pollen mother-cells of Liliwm canadense. Annals of Botany 19: 189-258. 1905 Bercus, J., La formation des chromosomes hétérotypiques dans la sporogénése végétale. I. La Cellule 21:171-189. 1904. II. Idem 21: 381-397. 1904. III. Idem 22: 41-53. 1904. IV. Idem 22: 139-160. 1905. CHAMBERLAIN, C. J., The pollen grain. Bor. GAZETTE 23: 423-430. 1897. Dixon, H. H., The nuclei of Lilium longiflorum. Annals of Botany 9:663- 5- 1695. FARMER, J. B. and Moore, J. E. S., New investigations into the reduction phe- nomena of animals and plants. Proc. Roy. Soc. London 72: 104-108. 1903. Grécorre, V., La réduction numérique des chromosomes et les cinéses de matu- ration. La Cellule 21: 295-314. 1904. Montcomery, T. H., Some observations and Oke upon maturation plichomena of ooh cells. Biol. Bull. 6:137-158. 1 ROSENBERG, O., Ueber = Individualitat der {Rx minicess im Pflanzenreich. Flora 93: aks 259. , Ueber die ‘ecoheica aioe im Drosera. Med. Stockholms Hégs. Bot. Tnst: (Reprint, 1904.) SCHAFFNER, J. H., The division of the macrospore nucleus. Bot. GAZETTE 23: ay an iby. , Acontribution to the life history and cytglogy of Erythronium. Bor. Gazerre 31: 369-387. 1 STRASBURGER, E., Ueber Remi ciecline! Sitzb. Kénig. Preuss. Akad. Wiss. 18: 587-614. 1904. Wattace, L. B., The spermatogenesis of the spider. Biol. Bull. 8: 169-184. 905. “> * 190 BOTANICAL GAZETTE [MARCH EXPLANATION OF PLATES XIT AND XIII. The plates are reduced five-eighths in reproduction. The figures represent- ing entire cells and nuclei were studied with a Leitz no. 4 ocular and +g oil immer- sion objective; the others with a Zeiss no. 12 ocular and a Leitz +g oil immersion objective. PLATE XII. Fic. 1. Microsporocyte with chromatin network beginning to form the spirem. Fic. 2. Microsporocyte with delicate threads showing a single row of promi- nent chromatin granules. ' Fic. 3. Single thread of the same. Fic. 4. Sporocytes beginning to separate. Nucleus with distinct arch contracted spirem in the so-called synapsis stage. Fic. 5. Single thread of the same showing a single row of chromatin granules. 1G. 6. Sporocyte some time after the synapsis stage, showing prominent spirem with single row of chromatin granules. The free ends are cut. Ics. 7, 8. Single threads of the same. Fic. 9. Sporocyte with chromatin granules mostly double and flattened in appearance. Fic. 10. Somewhat later stage, showing the beginning of looping of spirem, and division of chromatin granules. One nucleolus beyond the nuclear cavity. Fics. 11-17. Single threads from the same stage, showing chromatin gran- ules still single, some with granules partly double, and others with typical double rows of granules. Fic. 18. Microsporocyte with loops nearly developed. Fic. 19. A single loop of the same stage. Fic. 20. Chromatin loops some time before separation; somewhat cut. Fic. 21. A single loop from the same stage. Fic. 22. Section of microsporocyte, showing chromatin loops near the time of separation. Fic. 23. A single loop from the same nucleus. Fic. 24. Microsporocyte, showing the twelve chromosomes. Fics. 25-37. Individual chromosomes, showing various types of loops and coils and their position on the spindle threads. PLATE XImI. Fic. 38. Mother star, showing position of chromosomes. Micronucleoli in the cytoplasm. Fics. 39-42. Chromosomes from mother-star stage, showing mode of sepa- ration of limbs of loops. Fic. 43. Microsporocyte with daughter stars. Fics. 44-50. Daughter chromosomes from the daughter star, showing char- acter of of the loops. Fic. 51. Daughter-star stage, with micronucleoli in the cytoplasm. BOTANICAL GAZETTE, XLI SCHAFFNER on LILIUM PLATE Xil BOTANICAL GAZETTE, XLI SCHAFFNER on LILIUM PLATE XUil — 1906] SCHAFFNER—LILIUM TIGRINUM IQI Fic. 52. Loose daughter-skein stage with micronucleoli collected below the open ends of the chromosomes. Fic. 53. End of loose daughter-skein stage, with mnjcgemuriank collected among the chromosomes. 1G. 54. Beginning of second division, with chromatin loops in the nucleus and aaa ania in the cytoplasm. . 55. Mother star of the second division, showing characteristic appear- ance the chromosomes, with micronucleoli in the cytoplasm. Fics. 56, 57. Single chromosomes from the mother-star stage, showing posi- tion on the eres? threads. IGS. . Chromosomes from the metakinesis stage, showing the nature of the aiid splitting. 1G. 62. End of metakinesis stage. Fic. 63. Daughter-star stage of second division. Fic. 64. A pair of daughter chromosomes from the daughter-star stage of the second division. Fics. 65, 66. Pieces of chromosomes at the end of the second division, show- ing a single row of irregular chromatin granules. 1G. 67. Tetrad at end of second division. Fic. 68. Microspore at beginning of germination, with two large nucleoli in the cytoplasm. Fic. 69. Same stage as the preceding, showing three nucleoli in the cytoplasm. Fic. 70. Microspore in germination stage, showing a number of nucleoli in the cytoplasm. CYTOLOGICAL STUDIES ON THE ENTOMOPHTHOREAE. I. THE MORPHOLOGY AND DEVELOPMENT OF EMPUSA.'" EDGAR W. OLIVE. (WITH PLATES XIV AND XV) THE general development of the Entomophthoreae and the exter- nal morphology of its various members have been studied by a number of investigators. For a detailed review of the literature pertaining to the group, the reader is referred particularly to the papers of BREFELD (’71,’81,’84) and of THAXTER (’88). CoHNn’s (’55) results in his classical paper on the developmental history of Empusa muscae have been in certain respects considerably modified by later investigation. According to him, the fly first be- came diseased and the fungus followed as a consequence. The first indications of the disease which CoxN could find in the blood of the fly were numberless minute globular or irregularly shaped bodies, whose presence he could not explain otherwise than by the assump- tion of spontaneous generation (p. 334). These bodies, according to him, grow larger, become globular or ellipsoidal, and finally grow into the filament, which, by the formation of partitions, becomes the three-celled hypha characteristic of the mature fungus and consisting of spore, stalk-cell, and root-cell. This three-celled character of the hyphae of Empusa muscae was disproved, however, the next year by LeBert (’56) and later by BREFELD and others. Every investigator who has attempted to infect insects artificially has testified to the difficulties which he has encountered. COHN (’55, P- 342), in speaking of his own lack of success, wisely empha- sizes the caution which should characterize such experiments, noting that one should be certain that the insects experimented with are not already stricken with the disease, a more difficult task than would at first appear. I am under obligation to the Carnegie Institution of Washington for grants, which have rendered possible this investigation. Botanical Gazette, vol. 41] [192 ~ - = 1906] OLIVE—DEVELOPMENT OF EMPUSA 193 BREFELD (’70, "71, 77), however, was successful in transmitting the parasite through external inoculation of spores, and he found that in the case of Empusa muscae infection took place only through the thinner whitish parts of the skin on the under side of the fly’s body; whereas in another species, E. sphaerosperma, the infected hyphae gained an entrance at any part of the skin of the cabbage larva. This author has contributed more on this point than any other, in that he was able to germinate the conidia in artificial media as well as on the surface of the insect body, and to find with the microscope the germinating hyphae actually boring through, the skin of the host. In his series of articles, BREFELD has described the complete course of development of two species of Empusa, E. muscae Cohn and E. sphaerosperma Fres., which furnish quite differ- ent types of vegetative growth. According to him, in E. muscae there are formed from those germ-tubes which have penetrated into the body-cavity of the insect numerous detached non-nucleate cells, which reproduce by repeated yeast-like sprouting, and which grow within the fat-bodies of the host. At a certain advanced stage of the development of the fungus, the reproduction of the cells by budding ceases, and each grows at one or both ends into a long unbranched tube, which grows through the body-wall and produces at its external end a single conidium. In the other species, E. sphaerosperma, BREFELD found that the germ-tube produces, on the other hand, a copiously branching mycelium with many cross-partitions, which finally fills the body-cavity of the host. At the end of the vegetative period, this mycelium sends out hyphae which grow to the surface, branch digitately, and finally produce acrogenously at each ultimate end a single conidium. Em pusa sphaeros perma further differs from E. muscae in producing resting spores. COHN (’55, p. 343) had already suggested, since he could not make the conidiospores of E. muscae germinate, that per- haps the conidia themselves required a year of rest. But BREFELD (°70, ’71), proved conclusively that the spores of this form were short- lived, living only for a few days; hence his first suggestion in regard to the puzzling question as to the wintering of such a species was that this form was probably heteroecious, and that resting spores were pro- duced in some other host. Later, however (’84, p. 68), he seems 194 BOTANICAL GAZETTE [MARCH inclined to believe that the disease is continued over winter on flies in warmer regions, and that it migrates northward with the insects on the return of summer; the fallacy of which theory THAXTER has pointed out. THAXTER (’88) in his account of the Entomophthoreae of the United States, gives the results of morphological studies based on a considerable number of new as well as old forms. This author dis- agrees with BREFELD in regard to certain important points. In the first place, he maintains that the vegetative growth in E. sphaero- sperma is not filamentous in all cases, as is stated by BREFELD, and he appears to be inclined to think that both the filamentous mycelium and the broken-up, budding segments may occur in the same form under different conditions. He asserts that the usual multiplication of the hyphae is not by branching and continuous growth but by the formation of ‘‘ hyphal bodies,” which “ consist of short thick fragments, of very varied size and shape, that are continually reproduced by budding or division, until the insect is more or less completely filled with them.’’ But he continues further: ‘‘In cases where a direct mycelial growth follows the entrance of the hypha of germination, if indeed such instances occur, this mycelium must fall to pieces into hyphal bodies, before the commencement of growth the direct object of which is reproduction, in a fashion resembling that above described at a similar stage for Conidiobolus’’ (p. 140). This con- ception of reproduction by means of ‘‘hyphal bodies, ’’ however, for reasons that are stated later in this paper, must be abandoned, at least as a generalization. The segments of the vegetative hyphae, or “ hyphal bodies” as THAXTER terms them, under unfavorable conditions may each form a thick-walled resting chlamydospore; or, when the conditions are favorable, they may at once proceed to develop into the fructifying state. In the latter case, according to this author, each hyphal seg- ment sends out one or two (in some species more) hyphae which develop into conidiophores. In the simplest case, a simple conidio- _ phore grows directly to the outer air and produces a single conid- ium (Empusa muscae, e. g.). Or, the conidiophore may become compound and produce a set of conidia. Or, under very favorable conditions, ‘‘a single primary hypha may branch indefinitely, each . silane er 1906] OLIVE—DEVELOPMENT OF EMPUSA 195 ultimate branch becoming a conidiophore similar to those of the more simple case just mentioned” (p. 142). A singular method of germination of the “hyphal bodies” occurs in E. aphidis and E. virescens, according to THAXTER’s observations. Spherical bodies, evidently regarded as “hyphal bodies” with highly refractive con- tents, germinate and send out a mass of hyphae in all directions. In this condition they are said to resemble a head of Aspergillus, although the author does not show in either of his two drawings of the phenomenon any trace or remnant of the central cells or “hyphal bodies” from which the radiating hyphae are said to arise. The conidium is regarded by THAXTER as having a double wall, and thus is to be interpreted more properly as a simple single- spored sporangium. MATERIALS AND METHODS. In March 1904, the writer found in horse-dung cultures in the laboratory a small species of fly belonging to the genus Sciara, which was infected with an Empusa. This small fly with its attendant disease has been propagated in horse-dung cultures since that time, and many successive generations of the insect, the larval condition as well as the adult, during the year and more of its cultivation, have fur- nished a wealth of material for an almost complete cytological and developmental study of this species of Empusa. A number of other forms of the Entomophthoreae, most of them in the fructifying stage of their existence, have been used for comparison, but no others have as yet been traced through their entire life history. Enough has been learned, however, to show the existence of a most interesting series of distinctive variations. My material has been killed with a variety of fixing agents, mostly with varying strengths of Flemming’s chromic-acetic-osmic acid mix- ture. The insect body was generally cut in two or pricked to allow direct contact of the fixing fluid and the fungus hyphae in the body cavity. The material was sectioned usually 3-6 » thick and stained with Flemming’s safranin gentian-violet orange-G, or with Heiden- hain’s iron haematoxylin. Six species of Empusa altogether have been thus studied. These species, determined Zaccording to the descriptions in THAXTER’S 196 BOTANICAL GAZETTE [MARCH account of the group, are as follows: Empusa muscae Cohn, on the common house fly; E. culicis, A. Braun, on a small green species of Chironomus; Empusa sp., on a large fly; EZ. aphidis Hoffman, on vari- ous aphides; E. americana Thaxter, on a blue-bottle fly; and one other species, which is the principal one studied in this paper, on the small fly, Sciara sp. The ovoid conidia of this last form coincide closely with the description of the conidia of EF. montana Thaxter. They show decided differences, however, from THAXTER’s drawings of this species; and from £. ovispora Nowakowski and E. echinospora Thax- ter, with which the measurements of the conidia also almost coincide, the form is distinguished by the characteristic zygospores of the latter species. While it is possible that, under certain conditions unknown to me, this species on Sciara may produce zygospores, yet the fact remains that after over a year of continuous observation, I have failed to find any resting spores. It is therefore thought advisable to give a new name provisionally to this species, which will hereafter be referred to as Empusa sciarae. Empusa sciarae n. sp.—Vegetative hyphae forming a branching, septate mycelium, which in advanced stages is cut up into few- (gener- ally 3-5-) nucleate cells. Radial hyphae branched; conidiophores 3-5, bearing at each ultimate end a single ovoid uninucleate conidium with a rounded basal papilla, 12-16 X 18-25 w. Zygospores unknown. VEGETATIVE STAGE. My own observations of purely vegetative stages are concerned with two forms only, Empusa aphidis and E. sciarae, both of which agree with the type described by BREFELD for E. sphaerosperma. In all the other species studied, the vegetative condition had ceased and conidiophores had grown out from the vegetative hyphae. As observed by BREFELD, THAXTER, and others, the insect dies at the end of the nutritive period of the fungus; but they do not seem to have emphasized the fact that living insects alone must furnish data for the study of the vegetative hyphae. After the initiation of the repro- ductive period and the consequent death of the insect, the radial growth of the conidiophores produces a mass of hyphae which might readily be taken, in certain instances, for a vegetative filamentous mycelial growth. The probability has suggested itself to the writer 1906] OLIVE—DEVELOPMENT OF EMPUSA 197 that the fat cells, detached in the search for the fungus, must have been mistaken by some for stages in the development of Empusa; since these bodies may frequently resemble closely short hyphal segments in their fatty, granular contents, as well as in their assumption of globular or irregular shapes, which give the suggestion of budding cells. These detached fat-bodies, which prove in sections to be small aggre- gates of insect cells, are particularly abundant in the adult just pre- ceding ovipositing. An easy and certain method of distinguishing the vegetative cells of Empusa from fat-bodies which oceur with them in the body cavity, is to stain with a dilute methyl-green solution, acidulated with a few drops of acetic acid, when the fungus cells stand out conspicuously, distinguished by their relatively large and characteristic nuclei. In some preliminary observations on the manner of infection of Empusa sciarae, I have not been able to make certain of this point, but I wish to record here some notes of interest pertaining to it. The small fly, Sciara sp., was sometimes accompanied, in the vessels in which the dung cultures were kept, by three other species of small flies, which hatched generally in less abundance, Psychoda sp., a so- called moth fly, and two other undetermined forms. None, however, other than Sciara were infected, although many times I have noted conidia stuck on the surface of the bodies of the larvae as well as of the adults of the other species. Successive generations of infected Sciara larvae as well as adults have appeared with great regularity every month or six weeks. The fly lays its eggs on the surface of the dung or on the sides of the vessels, and the young larvae, soon after hatch- ing, crawl below the surface of the substratum. It would appear reasonable to assume that in this case infection would occur with ease in this very young condition, when the larval skin is thin and deli- cate, and before they had crawled below the surface cf the dung, where they would obviously not be reached by shooting conidia. My preliminary unsuccessful experiments at infecting healthy adult larvae, confined for a week in bottles with diseased ones, at least suggest the possibility of infection occurring in young insects only. If the cultures in which young larvae were being nourished were kept quite moist, perhaps the majority of individuals in this stage were killed by the disease. When, however, the conditions were 198 BOTANICAL GAZETTE {MARCH drier, the insect developed frequently into the adult fly, which itself, after ovipositing, generally died of the disease. In order to determine whether the disease could be carried from larval to adult condition, I have examined many pupae, but only in one or two instances have hyphae been found. In one case in which a young insect was struggling to get out of its pupal shell, a few Empusa hyphae were noted when the body was cut open, together with many small eggs and globular fat-bodies. When cultures were kept at some distance from infected ones, in an adjoining room, Sciara fre- quently developed to maturity by hundreds and died finally a natural death, evidently without infection. Resting spores in this species, as in E. muscae, have not been observed ; hence the puzzling question confronts us here also as to the method of wintering of the fungus. An interesting suggestion is made in this connection by the spontaneous appearance of Empusa sciarae in laboratory cultures as early in the spring as March, in 1904. This fact apparently still further renders useless BREFELD’s hypothesis as to the migration of the fungus north from warmer countries. The host, in this case, must have been breeding in the dung of the warm stables all through the winter, and it is quite reasonable to suppose that the short-lived fungus must have been continued at the same time on successive generations of insects. The successful cultivation of the form for more than a year in the laboratory gives additional reason for this belief. It is possible that Empusa muscae and such similar forms may have lost their sexual stage, because of their success in propaga- ting the disease by means of conidia alone. It is well known that a few house-flies survive the winter by hibernation or otherwise, and it is probable that some of the individuals during the winter may continue breeding in stables or in other favorable places, and in this way carry over the disease, even in cold climates. Living larvae of Sciara furnish beautiful material for a study of the vegetative conditions of this type of Empusa. When placed on a slide in water, a glance with the naked eye is sufficient to determine whether the larva is infected or not, since the diseased individuals are whitish in appearance, due to the presence of the more or less copious mycelium, while uninfected ones are transparent. Under the micro- scope, the diseased Jarvae show clearly the long, branched, mycelial | | ? ag eee eee ee Spee le eee > MA Naas 2. er a es Oe ee FA Tt WR a PRS FM RE aoa On eee Se RS a a oo =, 1906] OLIVE—DEVELOPMENT OF EMPUSA 199 filaments, which in a young condition may have but few septa, while in a later vegetative stage they develop numerous septa. As the larvae crawl about over the surface of the slide, the hyphae can be seen sometimes extending the full length of the body-cavity, some- times copiously developed in the posterior portion only, again more abundant in the anterior region. As the body parts move, or the blood flows in the cavity, the mycelium shows. corresponding pul- sations and movements. Fig. z showsa ‘portion of a branched vegetative hypha, taken from such a larva which was crawling just below the surface of the dung culture. Evidently the vegetative activ- ities have here almost ceased, and the 2-, 3-, or 4-nucleate cells are about ready to send out the radial conidiophores. Fig. 2 shows a section of a younger hypha, in which the cells contain a varying number of nuclei; while fig. 3 represents a still younger stage, the - earliest condition within the body of the host which I have succeeded in obtaining, growing in a larva in which the mycelium was composed of but a few scattered filaments. The section of the hypha, of which jig. 3 represents only a portion, shows in the preparation twenty-two nuclei, but in the entire length, as far as traceable, not a single cross partition. While in my own investigations of the vegetative stages of Empusa sciarae there remains as yet a small gap, from the penetration of the infecting hypha to the production of such multinucleate mycelial fila- ments as are shown in fig. 3, I feel reasonably certain as to the method of procedure. Germinating conidia, growing in a sterilized decoc- tion of cooked larvae, are shown in figs. 4-9. Figs. 4-7 illustrate the germination of the uninucleate conidia into germ-tubes which have grown out of the liquid decoction, thus resulting in the formation at once of secondary conidia; jig. 7 shows the beginning of the forma- tion of a tertiary conidium. In figs. 8 and g the germ-tube has grown into a hypha in which all of the protoplasm appears to be in the end cells; in the latter, sixteen apparently empty cells separate the old conidial wall from the terminal protoplasm, which still remains uninucleate. ‘Both BREFELD (’71, figs. 5, 29, 31) and THAXTER (’88, fig. 240) give figures showing a somewhat further. advance over what I have obtained in the cultivation of conidia, in that in their forms branches 200 BOTANICAL GAZETTE [MARCH are beginning to appear. Whether an increase of the amount of protoplasm and of the number of nuclei has accompanied this branching is doubtful. It is highly probable, therefore, that in Em pusa sciarae the hypha which has penetrated into the body-cavity of the fly from the germi- nating conidium grows rapidly at the expense of the nutrient fluids in which it floats. After the protoplasm has increased in volume, the nuclei increase in number by division, and from the uninucleate condition, in the case of Empusa sctarae, the hypha finally becomes a multinucleate branching filament, such as is shown in fig. 3. Partition-walls in this form at first occur but sparingly; later, how- ever, at the culmination of vegetative activity, septa are abundant and branching becomes more frequent. Finally the body-cavity of the insect becomes completely filled with the mycelial filaments, vegeta- tive activity ceases, and the fructifying state begins. Empusa aphidis furnishes a somewhat modified vegetative devel- opment. The advanced condition, which is my only source of infor- mation in this case, shows in sections branched coenocytic hyphae, which appear to be but rarely divided by septa, unlike the corre- sponding stage in Empusa sciarae. Even after the rhizoids have grown out from the under side of the body of the insect (figs. 10, 11), the vegetative activities appear to continue, as evidenced by the fact that the nuclei in this instance are still undergoing division. Also in Empusa sp. there occurs a similar prolongation of vegetative activity, since at an advanced stage even fewer cross-partitions can be found in the coenocytic mycelium (jigs. 23, 25). REPRODUCTIVE STAGE. At the culmination of vegetative growth, the body-cavity of the larva appears to be completely filled with a mass of long hyphae. Toward the close of this stage, the larva crawls to the surface of the substratum or high up on the side of-the culture dish; or, in the case of the adult Sciara, the fly seeks a conspicuous position, as is common with such infected insects, and death ensues with the beginning of the fructifying condition. The initiation of the fructifying condition is marked in Empusa sctarae by the sudden formation of radial branches from the short ial Fens, | an he oemeeneeee chien et eC ee ee ee ee ee A nes 1906) OLIVE-—DEVELOPMENT OF EMPUSA 201 cells which make up the hyphal filaments. These branches appear to be put forth more or less simultaneously, in this species but one from each cell. The cell swells up and becomes rounded off somewhat at its ends (fig. 13). One or more vacuoles appear in the protoplasm and a protuberance is pushed out from one end of the cell; this grows into the radial hypha destined to become the branched conidiophore (figs. 12, 14,15). The two forces, the swelling and consequent round- ing off of the cells at the ends and the pushing out of the branch, com- bine to split the partition-walls between the cells, thus causing the hyphae to become easily broken up into one-celled segments, the “hyphal bodies” of THAXTER (figs. 1, 12). I have seen similar hyphal segments, forming in these instances also the origin of the radial conidiophores, as well in my preparations of E. culicis and FE. americana; but other species promise interesting variations from this common type of pre-reproductive development. In Empusa sp. and E. aphidis, for example, the vegetative hyphae remain, up to the very initiation of the reproductive stage, either unicellular, or at least with cross-partitions at only rare intervals. In such instances, there- fore, no breaking up into “hyphal bodies” occurs; but the vegetative hyphae appear to grow out directly into the conidiophores. As BREFELD and THAXTER have pointed out, the first hyphae to appear in the external growth of the fungus form the rhizoids, by means of which the host is fastened firmly to the substratum. In the common house-fly, the host is attached by means of its proboscis. . In E. sphaeros perma, according to BREFELD, bundles of rhizoids break out more or less irregularly from the under side of the body of the insect and attach it to the substratum. According to THAXTER, these rhizoidal hyphae may branch, and may terminate in a kind of expanded sucker, which apparently secretes a viscous substance. . In Empusa sciarae, rhizoids are developed more or less abund- antly from the under side of the abdomen of the fly, or, in the case of the larva, from almost any point on the under side of the body. In certain forms of E. aphidis, groups of rhizoids break out from the under surface of the insect and form large sucker-like hold-fasts. In several instances, I have counted three of these hold-fasts from aphides parasitic on Solidago. Fig. 10 shows one of these sucker-like bundles of rhizoids in section; and fig. rz a highly magnified hypha from near 202 BOTANICAL GAZETTE [MARCH the inner region of the hold-fast. The latter presents clearly the char- acter of the typical rhizoid; thick, frequently yellowish walls, contain- ing but a thin layer of protoplasm, which bounds externally the large vacuolar cavity that almost fills the hypha. The walls of the ordinary vegetative hyphae, on the other hand, are thin; those of the rhizoids in this case appear to have undergone a gelatinous or slimy modification, and the contents seems to be undergoing degeneration. The growth of the conidiophores, in the case of E. sciarae, proceeds more slowly than that of the rhizoidal hyphae, a phenomenon which is probably due, in part at least, to the slower absorption of water by those hyphae destined to bear conidia. At any rate, the vacuoles which are formed at this time in the cells increase slowly in size, and a conidiophore arises from near the end of each cell and grows out radi- ally, boring its way through the tissues of the host (figs. 12, 14, 15)- As THAXTER and others have noted, the conidiophores of certain species remain simple and unbranched, as in the case of E. muscae (fig. 40); or, in other species, they may become normally septate and branched, as shown in E. sciarae (fig. 16), and Empusa sp. (fig. 23)-° From this fact arise most interesting cytological variations in the vari- ous conidia of these species. THAXTER has brought out clearly the variation in size, shape, etc., of the conidia of many species, but CAVARA (’9Q9) was the first to contrast the multinucleate condition of the conidia of Empusa muscae with that of the uninucleate conidia of Entomophthora Delpiniana. Of the six species studied by me, four have uninucleate conidia (E. sciarae, fig. 27; E. americana, figs. 36, 37; E. aphidis, figs. 42, 43; and Empusa sp., figs. 22, 26): The conidia of E. culicis are normally two-, rarely three-nucleate (figs. 31, 32); while those of E. muscae have a more or less indefinite number, frequently about 15-18 (figs. 38, 39).2. The more common uninucleate conidia arise primarily from the septation of the conidiophores into uninucleate segments; whereas, on the other hand, the simple conidiophore of E. muscae does not usually become septate except at the conidium. In the last case, therefore, all of the many nuclei of the last-formed vegetative cell, which forms the origin of the 2 While convinced of the value of CAVARA’s suggestion as to the use of nuclear characters in the classification of the Entomophthoreae I do not think that this one character alone would justify the separation of Entomophthora from Empusa. ee rr we 1906] ' OLIVE—DEVELOPMENT OF EMPUSA 203 conidiophore, flow out into the single, large, bell-shaped conidium (fg. 39). In E. culicis, which has binucleate conidia, septation of the conidi- ophore occurs, by which the protoplasm cuts itself off from behind; but, unlike the case of E. sciarae, I have found no branching with it, so that as a consequence, all of the binucleate protoplasm of the conidiophoric hypha flows into the one terminal conidium (figs. 30-35). The simple conidiophore of E. culicis resembles, therefore, that of EF. muscae in being unbranched, but the origin of the conidiophores may differ in the two cases. One may find, in fact, in E. culicis, two sorts of “hyphal bodies;”’ either small cells which, like the corresponding ones of E. muscae, give rise to but one conidiophore; or, on the other hand, larger cells, which may give rise to several conidiophores by budding from several points simultaneously, in a similar manner to Conidiobolus. In both instances, each conidiophore remains simple and ultimately bears, terminally, the binucleate conidiospore. In the species with uninucleate conidia, E. sciarae, E. americana, E. aphidis, and Empusa sp., the coenocytic conidiophore, as was indicated above, is cut up by septa, in a manner to be described later, into uninucleate segments (figs. 16-21). In all these cases, this results in a branching growth and successive abjunction of the acrogenous spores. Below the terminal cell the penultimate cell pushes out to one side, and thence bores its way to the surface of the host, where it abjoints a single uninucleate conidium (figs. 16, 23). From this habit in certain forms, arises a profuse system of branching, frequently of a digitate type (e. g., Empusa sp.) or corymbose (E. sciarae and others), to enable the uninucleate segments to reach the surface and to discharge their protoplasm by means of the abjointed conidia. But the last-formed vegetative cells of E. sciarae contain only 2-4 nuclei (figs. 1, 12, 15), so that, in this instance, only a correspondingly small number of branches are formed. _ The process of abjection of the conidia of Empusa is apparently similar in a general way to that described for the sporangia of Pilob- olus, except that in these conidia there is no gelatinous collar visible. The formation of the partition at the base of the conidium in Empusa also is quite a different process from the formation of the columella in the case of Pilobolus. In Empusa, the vacuole which appears in the basal portion of the 204 BOTANICAL GAZETTE (MARCH ‘cell becomes larger and larger (fig. 27), and a small protuberance, which has a diameter equal to about half that of the conidiophore, is pushed out from the end (figs. 34, 41). There is now formed at the apex of this narrowed sterigma a swelling (jigs. 27, 35, 39, 42), which, after continued enlargement, finally receives the greater part of the protoplasm and all of the nuclear content from the basal portion. The process of cell-division, by means of which this apical conidium is cut off by a wall from the penultimate cell, will be described in some detail later, in connection with the description of cell-division in Empusa. Certain points should be noted here, however, and among them, that the term basidium, as applied to the penultimate cell, although in common use in this connection, should, in my opinion, be confined to the Basidiomycetes, where, morphologically, the true basidium is a very different spore-bearing structure from the penultimate cell bear- ing the conidium in Empusa. The penultimate cell forms the explosive mechanism by which the conidium is shot off. As the ring-formed wall which cuts through the base of the conidium travels progressively inward, the protoplasm passes through the narrowing opening leading from below, until at the close of abjunction, the basal cell retains only a thin parietal layer of protoplasm, but no nucleus. Continued swelling, due to the absorption of water, finally results in the bursting of the basal vesicle, thereby breaking the wall of the vesicle where it joins that of the conidium. In some forms, a ring-shaped scar is noticeable near the base of the conidium, marking the circle where the summit of the swollen basal vesicle was formerly attached. The septum which separates the conidium from the subterminal cell is at first usually pushed upward, thus resembling a columella (figs. 28, 30, 31, 36), but when the spore is shot off, this partition-wall reverses its former position, and in the conidium it appears as a prominent papilla (jigs. 29, 37, 43): When the basal vesicle bursts, its contents are thrown out of the open ruptured end and frequently persists as a slimy covering about the spore, serving in this case, perhaps, the double purpose of protection against excessive evaporation and of sticking the spore to the sub- stratum which it strikes. I have noted that the explosion, in the case of Empusa sciarae, sometimes throws the spore a distance of 67"; while BREFELD has recorded an even greater distance in the case of E. muscae, in which the spores are said to be sent as far as 2-3°™. . a —T 1906] ' OLIVE—DEVELOPMENT OF EMPUSA 205 I have not yet completely solved satisfactorily to myself the pecul- iar method of abjection of the spores of my undetermined species of Empusa. I am inclined to think, however, that this method, in certain respects, is unlike that described above. In this form the conidio- phores come to the surface, become cut up by septa into uninucleate segments, and proceed to branch profusely (jigs. 23, 25). The cell terminating each branch pushes out in a peculiar manner. Instead oi forming a large basal vesicle as an explosive mechanism in the usual manner, as described above, here the protoplasm appears to cut itself off from behind by means of one or more successively formed walls (figs. 22, 24, 26). A minimum of protoplasm seems to be lost in the process, and this cut-off protoplasm soon assumes a peculiar granular appearance. In this condition it is probably dead, for these cut-off cells appear soon to lose their turgescence. _ It is difficult to con- ceive of a forcible discharge of the spore in this instance, especially if it be true that the protoplasm of the basal cells is dead and thus inca- pable, through loss of turgidity, of functioning as an explosive mechan- ism. The process here rather seems to be that, by means of these successive abjunctions, the uninucleate spores are pushed off with but little force, and that they are probably followed out of the thick, gelatin- ous wall of the mother hypha by other cells pushed up from below. It may be, however, that further studies on fresh material of this species will change this impression of subterminal proliferation. Figs. 45 and 46 represent the terminal portion of large hyphae of E. culicis which are destined to form resting spores; and figs. 47 and 48, two fully formed, thick-walled resting spores. Such hyphae as are shown in the first two figures are distinguished from conidiophores by being much larger, and, further, they contain four or five nuclei, instead of two. I have traced these hyphae in sections back to large “hyphal bodies,” but I was unable to follow their complete history. Whether the thick-walled resting spores of this species are therefore true zygospores, or azygospores, as they are termed in THAX- TER’s monograph, I am not prepared to say. I have not been able, however, to confirm VUILLEMIN’s assertions (:00) as to the nuclear fusions in the azygospores of Entomophthora gloeos pora. UNIVERSITY OF WISCONSIN, Ison. 206 BOTANICAL GAZETTE [MARCH LITERATURE CITED. BREFELD, O., "70, Entwicklungsgeschichte der Empusa muscae und E. radicans. Bot. Zeit. 28; 161-166, 177-186 , 71, Untersuchungen iiber die Entwicklung der Empusa muscae und E. radicans. Abhandl. naturf. Gesells: Halle 12: rf. pls. 4. » > pas die Entomophthoreen und ihre Verwandten. Bot. Zeit. 35: 345-355, 368- —— a En hes radicans. Untersuch. tiber Schimmelpilze 4:97- Tit. -7- oe Conidiobolus ae und minor. Untersuch. tiber Schimmel- niin 6:35-72. pls. 3 Cavara, F., ’99, Osservazioni a sulle certo thchone: Nuovo Giorn. Bot. Ital. N. S. 6: 411-466. pls. Coun, F., ’55, Empusa muscae in ae Krankheit der eset Nova Acta head: Caes. Leop. Carol. Germ. Nat. Cur. 25:3 Lesert, S., ’56, Die Pilzkrankheit der Fivesen. Abhandl. Naturf. Gesells. Zuri THAXTER, R., ’88, The Entomophthoreae of the United States. Mem. Boston . Nat. Hist. 4:133-201. pls. 14-21. VurtLemtn, P., :00, Développement des azygospores d’Entomophthora. Compt. Rend. Acad. Sci. Paris 130: 522-524. EXPLANATION OF PLATES XIV AND XV. The drawings were made with the aid of an Abbé camera-lucida, and for the most part with various compensating oculars combined with Zeiss 2™™ apochro- matic obj. N. A. 1.30. PLATE XIV. 1Gs. 1-9, Empusa sciarae. Fic. 1. A freshly killed pie fixed and stained with acetic methyl green. % 275. Fic. 2. Section of a younger filament. Fic. 3. Section of a still younger vegetative hypha. 275. Fic. 4. A germinating conidium, cultivated in a decoction of cooked larve, oe a Ain conidium. Killed in acetic methyl green. X575. IGS. 5, 6. Conidia with still younger germ-tubes. Killed with acetic methyl ok *575- Fic. 7. A germ-tube has formed a secondary conidium, which in turn has started to form a tertiary conidium. Killed ditto. 575. Fics. 8, 9. The conidia have produced long germ-tubes instead of secondary conidia. From the same culture as those above. Killed ditto Fics. to-11, Empusa aphidis Fic. to. A cross-section of one of the sucker-like rhizoidal disks. Fic. 11. A section of a single rhizoid. x 1080. Ne BOTANICAL GAZETTE, XLI ae ; PLATE XIV pene ae ten ; me é E. W. Olive, del. PLATE XV OLIVE on EMPUSA . we del. E. W. Olive, ni qo ne a sees BOTANICAL GAZETTE, XLI 1906] OLIVE—DEVELOPMENT OF EMPUSA 207 Fics. 12-21, Empusa sciarae. Fic. 12. A section showing two cells of the old vegetative mycelium, or “hyphal bodies,” from the upper one of which has grown out radially a conidiophore.. Fic. 13. A hyphal segment freshly killed with acetic methyl green. X 480. Fic. 14. The same, showing the beginning of a conidiophore. X 480. Fic. 15. The same, showing a still longer conidiophore. 4 Fic. 16. Section of a radial conidiophore which is just ees > deca the epi- dermis of the host, showing the subterminal branching. x1 Fic. 17. Section of a conidiophore showing a palsy ERS between the two cells. 1080. IG. 18. A half-completed stage of cell-division in a conidiophore. X 1080. Fic. 19. Cell-division in the vegetative mycelium. 00. IG. 20. An almost-completed stage of cell-division in a conidiophore; the iron- haematoxylin stain has been almost completely washed out of the preparation, except at the innermost margin of the cleft. x 1500. Fic. 21. A poorly differentiated construction in a vegetative hypha. Fics. 22-24, Empusa sp. ? Fic. 22. Section of a conidiophore, showing the peculiar method of the cutting off of eee cells, which apparently soon die. x1 - 23. Section of a conidiophore showing a subterminal branch, and also be hae appear to have assumed amoeboid shapes. X 1 Fic. 24. A similar section to that shown in fig. 22, but with aly one basal cell cut off. * 1080. Fics. 25, 26, Empusa sp. Fic. 25. End of a conidiophore, from which has been cut off two uninucleate segments. X 1080 Fic. 26. A re which is apparently being pushed out of the thick, slimy wall of the mother hypha. 575. Fics. 27-29, Empusa sciarae. Fic. 27. A section showing two protruded terminal cells of the conidiophore, from which young conidia are in process of formation. x1 Fic. 28. A section of a conidium in process of shetrintion, 7 in which the iron- haematoxylin stain has not differentiated the nucleus aay has brought out some of the metachromatic bodies. X 1500. Fic. 29. A mature conidium. Fics. 30-32, Empusa culicis. 1500. G. 30. The upper portion of the terminal cell of a conidiophore, showing at sigs side the cleft marking the ring-formed cleavage-furrow. Fic. 31. A section showing a somewhat older stage, in which the cleft has almost abstricted the binucleate conidium Fic. 32. A rare occurrence, showing a ‘pontiac conidium. 208 : BOTANICAL GAZETTE [MARCH PLATE XV. Fics. 33-35. Empusa culicis. X1500. Fic. 33. A thin, lightly stained section of the end of a conidiophore, showing the usual binucleate condition. Fic. 34. An older stage, in which the sterigma is ‘ poditins ou Fic. 35. From a still older preparation, in which the end : the sterigma has become swollen to form the binucleate conidium. Fics. 36, 37, Empusa americana. 1500. Fic. 36. A preparation in which the nucleus is poorly differentiated, but which ina especially clearly the vacuolar cytoplasm and the columella-like wall which finally abstricts the conidium. Fic. 37. A mature conidiospore, showing the vacuolated cytoplasm (probably filled vation living with oil-globules), the single nucleus, and the basal papilla which has been formed by the reversal of the columella. Fics. 38-40, Empusa muscae. X1080. Fic. 38. The partition-wall which cuts off the multinucleate conidium is here Sihirteint: The slimy enucleate protoplasm of the basal cell appears to be but ittle shrunken. IG. 39.. The end of a young conidiophore, just extruded from the body of the fly, in which the conidium is in process of formation, showing the multinucleate character of the protoplasm. : Fic. 40. A portion of a conidiophore still within the body of the fly, which is growing toward an opening in one of the abdominal joints. Fics. 41-44, Empusa aphidis. X1500. Fic. 41. A section showing at the tip of the conidiophore a young conidium in process of formation. | Fic. 42. An older stage of conidium-formation, Fic. 43. A mature conidium, showing the uninucleate character. Fic. 44. A nucleus from a conidiophore, showing two nucleoles, and portions of the chromatic thread. ; Fics. 45-48, Empusa culicis. Fic. 45. The end of a young azygosporic hypha. Fic. 46. An older condition, which shows the beginning‘of the formation of a resting spore. X 1080, Fic. 47. A mature ‘“azygospore.”’ X 1080. Fic. 48. A thinner section of a mature eeu ”” showing but one nucleus. Other nuclei lie below and above in the section. X 1500 a DRIEPER A REGLE s NEW NORMAL APPLIANCES FOR USE IN PLANT PHYSI- OLOGY III (WITH TWO FIGURES) In the two preceding articles I described several pieces of apparatus newly devised for educational work in plant physiology and explained the objects I have in view in their development. In brief I aim to provide for each of the principal physiological processes such apparatus as will be accurate in results, convenient in manipulation, and obtainable by pur- chase from a supply company. The company to which the manufacture has been delegated is the Bausch & Lomb Optical Co., of Rochester, N. Y. In the earlier articles I named the appliances ‘‘precision-appliances,” which some, though not all of them are; they are, however, more properly normal appliances, which I shall henceforth call them. VI. PHOTOSYNTHOMETER. No fact in all the physiology of plants is more important, and hence more imperatively demands complete demonstration in botanical educa- tion, than the absorption of carbon dioxid by green plants in light, with the equivolumetric release of oxygen. There are simple ways of demonstrating the process in part, and somewhat complicated ways of demonstrating it completely; but hitherto there has been no simple method of demonstrating the entire process in one operation. This is effected, however, by the new photosynthometer described below, and illustrated in the accompanying fig-1. tis called by this name for the reason that it permits photosynthesis (the quantity of the photosynthate being a function of the quantity of the gases absorbed and released) to be measured as well as demonstrated. The instrument consists essentially of a pear-shaped plant-chamber set in a firm iron base, a graduated measuring tube with a small stop-cock at the upper end, and a connecting stopper furnished with a stop-cock of con- siderable bore. The total capacity of the apparatus when closed is exactly 102°, of which the 2° is for a shoot and 100° for the gases concerned. The proper amount of shoot is provided by selecting a small-leaved plant and pushing a branch down into a measuring-glass until it displaces exactly 2°¢ * Continued from Bot. GAZETTE 39: 152. February 1905. 209 | [Botanical Gazette, vol. 41 210 BOTANICAL GAZETTE [MaRcH of water; the water-level is then noted on the stem, which is cut at this point under water, the shoot being later, when dried, placed upright in the chamber. (It hardly PIGS: shows in the figure because of irregular reflections from the surface of the chamber.) Taking advantage of the fact that the shoot will carry on photosynthe- .sis for a time in an atmosphere con- taining carbon dioxid in demon- strably large amount, even up to 10 per cent. or more, we add some selected percentage of that gas to the apparatus in this way. The measuring tube, with stop - cock closed, is inverted and filled with water of room-temperature, up to a figure of the graduation expressing the selected percentage, for the tube is graduated in cubic centimeters, which are, of course, percentages of the total gas capacity of the appara- tus.. The stopper is then placed on the tube, and its stop-cock closed; its hollow is filled with water and inverted in a pneumatic trough (or equivalent dish of water), which has been standing in the laboratory long enough to take the temperature of the air. The lower stop-cock is then opened and carbon dioxid from a generator is allowed to enter the tube, either from below or, as is most convenient, through the top of the tube. The admission of the gas may be perfectly controlled by cautious manipulation of the upper stop-cock, which is closed at the moment when the water has been wholly driven out to the bottom of the bore of the lower stop-cock, which point is held exactly at the water-level. The lower stop-cock is now closed, and the combination, which now contains exactly the desired percentage 1906] BRIEFER ARTICLES 211 of carbon dioxid, is lifted from the water, shaken free from all adhering drops, and placed in position on the chamber. To prevent compression, and therefore the presence of too great a quantity, of air in the chamber when the stopper is pushed into place, tiny holes (visible in the figure), matching in stopper and chamber-neck, allow free release of such pressure, and the chamber is perfectly sealed by twisting the stopper a little. The lower stop-cock is then opened, permitting the carbon dioxid of the tube to diffuse into the chamber, a process hastened by its gravitational flow. The apparatus now contains obviously 2° of plant and 100° of gas, of which a known percentage (say 5, 8, or 10) is carbon dioxid, and the remainder is air. The instrument is now placed in a bright light (not direct sunlight) for three or four hours; then the lower stop-cock is closed (as shown in the figure), shutting into the tube a sample of the gas of the chamber at the close of the experiment. The analysis of this gas can be made at leisure, and is accomplished thus. The stopper and tube are removed and placed upright in the pneumatic trough, deeply enough to allow the stopper to be taken off without admission of air to the tube. The zero mark of the tube is then brought exactly to the water surface; the upper stop-cock is cautiously opened, permitting the water to rise slowly to the zero mark, when the stop-cock is again closed, shutting into the tube exactly ro®° of the gas to be analyzed. First the quantity of carbon dioxid in the tube is determined, which is accomplished by aid of a reagent tube, of the form shown at the bottom of the figure. This tube, of glass, is provided with an extension of soft rubber tubing closed by a screw clamp, and it is filled completely to the clamp with a strong solution of caustic potash. It is slipped under the water of the pneumatic trough, with the clamp closed; the air, if any, is squeezed from the upper part; and the rubber is slipped over the lower end of the measuring tube which it grips firmly. The whole is then lifted from the water, the clamp is opened, the combination is inverted and the liquid is allowed to flow back and forth several times from one tube to the other, when it will completely absorb any carbon dioxid present. The clamp is then closed, the combination is slipped again under water, and the rubber tube is pulled off, when the atmos- pheric pressure will instantly force up the water to the exact extent of the carbon dioxid absorbed, permitting the amount, and hence the percentage, to be read off directly. Next a determination of the percentage of oxygen present is made. This is effected by a precisely similar method, using an- other reagent tube containing pyrogallate of potash, freshly made up in the usual manner. A few inversions of the combination will result in absorp- tion of all the oxygen, and the water-level in the tube when the rubber is 212 BOTANICAL GAZETTE [MARCH removed will give directly the percentage originally present. Some slender vessel may then be slipped under the tube which is removed and supported as shown by the figure. Thus may the gas exchange in photosynthesis be demonstrated accu- rately, completely, logically, and conveniently. n studying the process with beginners, the demonstration is the more striking and conclusive to them if a second instrument is set up like (and beside) the first, but covered completely from light; while even a third, like these two except that it has no plant, may advantageously be added. The comparative analyses of the gases at the close of the experiment give results leaving nothing to be desired in logical demonstration. For all ordinary purposes, water may be used in the pneumatic trough, its slight absorption of carbon dioxid being negligible; but for great accu- racy mercury may be employed. Similarly the corrections for capillarity, vapor tension, etc., may for elementary demonstration be ignored, though in precise work they would be taken into account. Temperature and baromet- ric pressure are compensated, obviously, by the method of use of the instru- ment. A much larger instrument, capa- ble of taking large leaves, branches, or even entire potted plants, but operated upon substantially the same principle, is now in advanced prep- aration and will be described later. VII. ALUMINUM SHELLS FOR TRANS- PIRATION EXPERIMENTS, ETC. In transpiration studies with potted plants it is of course neces- sary to eliminate evaporation from pot and soil. There are many ways of effecting this, of which the best, perhaps, is the use of a tin cup or glass jar to cover the pot, and a roof of rubber sheeting. The advantage of this method over those in which the plant is wrapped in rubber, sealed in melted wax, etc., is this,— that the rubber roof may be readily detached from the can or jar and lifted, permitting a complete change of air to the roots when the plant is * watered, thus contributing greatly to the health of the underground parts. Pies Ss. eT ee ee ae ee 1906] BRIEFER ARTICLES 213 The aluminum shells here figured (fig. 2) are designed to provide light, neat, clean, and easily applied covers for pots on this principle. Flower pots are now made so nearly in standard sizes that it is possible to make the shells to fit them closely, and shells will be made for the present in 3-inch, 34-inch, 4-inch and 5-inch sizes. To hold the rubber roof tightly to the shell, a tightly-fitting band or strap of aluminum, resting in a groove just below the strengthened top of the shell, may be drawn to any desired tightness by a convenient screw-nut, shown (though dimly) on the right in the figure. The rubber roof may be attached to the plant in any of the ways ordinarily used, but I find upon the whole the best method to be the following. In the middle of a proper-sized piece of medium-thick rubber-sheeting, a hole a little smaller than the stem of the plant is made with a cork-borer, and a cut is made with scissors from this to the margin of the piece. It is then placed around the stem, the cut edges of the central hole are stretched to overlap a little, sealed together with rubber cement and held clasped until this sets. Then a line of the cement is run to the margin, sealing one edge over the other. When fully set, the margin of the rubber is clasped to the shell, all surplus material is cut away, and a very neat and perfectly tight roof temains.—W. F. GANonG, Smith College, Northampton, Mass. CURKENT EITERATURE, BOOK REVIEWS. Bacterial diseases. Tuts quarto volume on bacterial plant diseases, by Dr. ERwin F. SMITH,’ is a manual for plant pathologists, which, with the author’s characteristic care in clearly stating details, covers the subject, from the angle at which a tube should be held during inoculation to plans for the construction of a water still and sugges- tions as to the proper developer for photographic plates. One of the fundamenta reasons for the abundance of half-finished work on bacterial plant diseases which is annually imposed upon the public has been the lack in all literature of just such a sae? " ah areca =nitee which the author has scattered through the work uncertainty which now pervades the subject in the minds of many would be oan dissipated. The Carnegie Institution is surely treading on dangerous ground in publishing such a manual, but if such action can be justified at all, it can be in this case. Here the need is great and the subject is so specialized as to make such a publica- tion impossible except at a financial loss. So great was the need that a proper presentation of the material promised for the second volume could hardly be made without a considerable portion of the matter here given. The volume is divided into three parts; 186 pages being devoted to the text, 16 to useful laboratory formule, and 63 to bibliography. The latter covers the gen- eral field of bacteriology, with the exception of plant diseases, this division being reserved for the second volume. The publication is well illustrated, many of the cuts showing the effect of bacteria upon plant tissue. The index covers the entire volume and will be especially useful in connection with the bibliography, which is arranged by subjects in the body of the volume. g the many original things in this suggestive work, the discussion of “keeping of records” and “nomenclature and classification” are of special interest. Any one who has attempted to keep an accurate history of the behavior of a plant parasite in the laboratory and in its host when the work has extended over a number ‘of years, has felt the need of an improved system. The subject of bacterial classi- fication has been fairly quiescent for some years, and we have been busy trying to fit the forms as they are found into MicuLA’s somewhat artificial framework. The proposition of the author to replace MicuLa’s Pseudomonas by Bacterium and send the group represented by B. anthracis masquerading under the name of A planobacter will come to some as a discouragement 1SMiTH, E. F., Bacteria in relation to plant diseases. Vol. I. 4to. pp- xii 285. Washington: Carnegie Institution, 1905. 214 ap eee ne 1906] CURRENT LITERATURE 215 In spite of some minor things which seem inseparable from originality, this work is of the first quality and should be in the hands of every plant pathologist. —H. A. Harpine. MINOR NOTICES. Grasses of Iowa.— As a supplementary report for 1803 the Iowa Geological Survey issues part u of the Grasses of Iowa,? prepared by PAmmet, BALL, ScCRIBNER, and others. This is a descriptive and geographical study of the grasses of the state, their general and economic aspects having been treated in part 1. Under each genus there is the generic description, with synonymy, a key to the species, a description of the species, often a figure, a list of localities and a map showing the distribution of each form in the state, and a statement of distribution in North America and elsewhere. There isa chapter on physiography and geology, with a map, a section onecology, and a partial bibliography of works on grasses. The work seems very complete and should be especially serviceable to Iowa botanists. It is a pity state printers are so seldom skilfuy ook-makers.—C. R. B Connecticut fungi.— The recently established natural history survey of Con- necticut has begun to show results, in the publication of two bulletins listing the Hymeniales and Ustilagineae of the state. The formers lists 375 species in 65 genera, gives analytic keys to the genera, and illustrates the commoner species by admirable half-tones, most of which are original. The species of smuts+ are described with lists of hosts and distribution, and notes on economic features.— oe NOTES FOR STUDENTS. Photosynthesis and temperature.—The interesting results of Miss iia on temperature as a limiting factor for photosynthesis’ have now been extended by her work in cooperation with BLACKMAN. ey have endeavored to interpret the quantitative variations of photosynthesis, under approximately natural conditions, in terms of the three limiting factors thereto, viz. (1) intensity of illumination, (2) temperature of leaf, (3) pressure of CO,. When a leaf is - 2PamMeL, L. H., Batt, C. R., and ScriBNer, F. L. The grasses of Iowa. Part 0, Iowa Geological Survey, co report. 1903. 8vo. pp. xiv+ 436, figs. 270. Des Moines, Iowa. 1904 3 WHITE, E. A., A pr re report on the Hymeniales of Connecticut. State Geol. and N. H. Survey Bulletin 3. 8vo. pp. 81. pls. 4O. 1905. 4CLinTon, G. P., The Ustilagineae or smuts of Connecticut. Idem, Bull. 5. 8vo, Pp. 45. figs. 55. 190 5 See Bot. GAZETTE 38 : 476. 1904. BLackan, F. F., and Matraaet, G. L. C., Experimental researches in vege- table assimilation and reanirations. IV. A quantitative study of carbon dioxide assim- ilation and leaf temperature in natural illumination. Proc. Roy. Soc. London B. 76: 402-460. 1905. 216 BOTANICAL GAZETTE [MARCH exposed to diffuse daylight alone the amount of photosynthesis is a measure of the light, and it varies with varying light only when the amount of carbon dioxid in the atmosphere is artificially increased and the temperature is kept high. Ifnot, photosynthesis is limited i and is constant though the light vary. Isolated leaves may rise more than 10° C. above a bright mercury thermometer in the sun, a result quite at variance with Brown and EscomBr’s results,7 which, how- ever, were calculated, not observed. Further study of this point is needed. At normal temperature leaves are not able to utilize the full amount of energy absorbed; helianthus could reach its maximum at 29° C. with about 68 per cent. full sunlight and cherry laurel with about 36 per cent. When light is the limit- ing factor equal intensities produce equal photosynthesis with leaves of most various structure and type. At low temperatures leaves as different as helianthus and cherry laurel have similar photosynthetic maxima, but at high temperatures these diverge. Thus at 29.5°C. the former can fix twice as much CO, as the latter, requiring twice as much energy to do it, of course. The essential differ- ence in the photosynthetic activity in different leaves lies, then, in that they have different coefficients of acceleration of this function with increasing temperature. So in nature it appears that the low pressure of CO, (entailing slow diffusion after solution at the surfaces of the leaf cells) and the low temperatures are the serious impediments to food making.— Root tubercle cultures——Much interest has been excited during very recent years by work done in the Department of Agriculture concerning soil inoculation with various root tubercle bacteria. Widespread and rather unfortunate notori- t knowledge concerning the root tubercle is to be attributed to the recent investi- gations conducted i in the Department. This popular impression is of course erroneous. The two distinctive contributions to this subject claimed by the workers in the Department of Agriculture were that the nitrogen-gathering ability of the bacteria was heightened by new cultural methods, and that a method of transportation in dried condition, upon cotton, had been devised, whereby pure cultures could be distributed readily to farmers. Much skepticism has existed concerning the possibility of practically height- ening the nitrogen-gathering power of the bacteria, and in a recent bulletin ® Harptnc and Prucwa claim to have demonstrated by an examination of eighteen of these cotton cultures that such packages are worthless for practical — since the organisms are unable to survive upon the cotton or survive uch small numbers as to be practically valueless. ‘Substantially identical sl upon six of these packages were obtained in five separate laboratories,” and the reviewer may add that similar results were obtained in his own 7 See Bor. GAZETTE 40 : 473. 1905. 8 Harpine, H. A., and Prucua, N. J., The apa of commercial cultures for legumes. N. Y. Agr. Exp. Sta. Bull. 270: 345-385. I 1906] CURRENT LITERATURE 217 laboratory. The inability of the cultures to live is attributed to the method of preparation and not to any knavery upon the part “of the commercial producers. A test conducted by the authors of this bulletin demonstrated the inability of the organism to survive to a satisfactory degree upon the cotton. Any intention of opposing the idea of treating the seed of legumes with living bacteria is distinctly disavowed. It is exceedingly unfortunate that this method should have been given such wide publicity and launched as a commercial enterprise until the question as to its efficiency had been thoroughly tested.—F. L. Stevens. Streaming of protoplasm in mucors.—This go-naten although very striking and_easily observed, has been little studied. The ment was noticed by WorRoNIN in 1866 in Ascophanus pulcherrimus. It was pears with consid- erable detail in a number of species belonging to different genera by SCHROTER, writer of the latest account,? in 1897, and the conclusion was drawn that the Movement was dependent upon osmotic conditions. A careful study was also ment was affected only very slightly by variation in the intensity of light. The action of ether, extremes of temperature, pressure, wounds, variation in amount of carbon dioxid, was similar to that of the same agents when applied to the higher plants. The streaming is found to be due to osmotic action and trans- piration and therefore does not occur in a homogeneous substratum, as for instance when the fungus is wholly submerged, or in a saturated atmosphere. The stream- ing is not a rotation or circulation, as in the hairs of roots and stamens and in the cells of Chara, Nitella, Vallisneria, etc., but a backward and forward-movement, in which the protoplasm, vacuoles, and nuclei participate. Occasionally the acropetal movement is somewhat balanced by a thin peripheral layer of proto- plasm without vacuoles setting up a basipetal movement. Usually the movement -is toward one end of the hyphae for a longer or shorter time, then stops and starts again in the opposite direction —J. C. ARTHUR. Germination and radium emanations.—KO6ORNIcKE’® has continued his study of the effect of radium emanations on the germination of ungerminated seeds which have been exposed in both the dry and wet condition. His earlier tests were made with radium bromid contained in glass tubes. In his later study he has used a much more powerful mixture which was contained in tubes having one side of . 9 SCHROTER, ALFRED, Ueber Protoplasmastrémung bei Mucorineen.* Flora 95: I-30. 1905. satis : 10 K6RNICKE, M., Weitere Untersuchungen iiber die Wirkung von Réntgen- und Radiumstrahlen auf die Pflanzen. Ber. Deutsch. Bot. Gesells. 23: 324-332. 1905. 218 BOTANICAL GAZETTE [MARCH aluminium, through which the emanations pass more readily. In all the trials he finds that although the germination is not prevented there is a period of retarded growth in the seedling. The elongation of the root or stem may be temporary or rmanent according to the duration of the exposure. In the latter case the injured organ persists indefinitely without disorganization, but further growth of the seedling occurs in the form of secondary members. In the case of Vicia Faba such a condition will follow an exposure of only one hour; yet an exposure of four- teen days does not prevent germination. Since the retardation of growth occurs sooner in the root than in the stem of a given seedling, the author favors the expla- nation offered by other investigators, who have worked on entirely different mate- rial, that organs engaged in photosynthesis are more resistant to the emanations. The author’s experiments offer no conclusive evidence on this point. Organs of seedlings from seeds exposed to emanations retain geotropic sensibility as long as they are capable of growth, the two capacities being concurrent. The same is true of heliotropic sensibility. His earlier view that radium emits enough luminosity to induce heliotropism, which was questioned by Mo tiscu, is maintained. Im- portant as these results are, it seems to the reviewer that their value would be much greater if obtained under standardized conditions—RAyMoND H. Ponp. Anatomy of Matonia.—TansLry and Luria describe the development and mature anatomical structure of a number of specimens of Matonia pectinata gathered by one of them on Mount Ophir in the Malay Peninsula.": The coty- ledons in this species are bilobed as in the polypodiaceous ferns. Below the first leaf the central cylinder of the young stem consists of a rod of xylem, surrounded by parenchyma alone; later phloem appears on the outside of the stele and in the center as well. Subsequently the endodermis and “oround tissue”’ likewise appear within the stele, which becomes typically siphonostelic. By a process of “local dilatation of the m rgin of the leaf gaps” an internal mass of fibrovascular tissue appears, which ultimately becomes tubular and lies within the original fibrovascular tube. This inner tubular fibrovascular bundle subsequently gives off an internal tracheary strand, which may also become tubular, so that there may be in Matonia as many as three tubular bundles lying one within the other. These join each other only in the region of the nodes. The authors consider the internal fibrovascular system asa storage tissue only, since it has no direct con- nection with the roots, which are attached to the external cylinder, as in other ferns of this type. The views as to the morphological nature of the complex fibrovascular system of the stem in this species may be regarded as “ orthodox,” since the conclusion is reached that it constitutes a single stele. The hypothesis that the pith is intruded cortex is accordingly rejected, since the authors are of the opinion that the only trustworthy criterion as to the morphological value of tissues is to be derived from a study of their relation to the aera meristems of the growing point.—E. C. JEFFREY. _TANSLEY, A. G., and LuLHaAM, Miss R. B. J., Astudy of the vascular system of Matonia pectinata. Annals of Botany 19:476-519. pls. 31-33. 1905- | | 1906] CURRENT LITERATURE 219 Chloroplasts of sun and shade plants. —LuBimENKO” refuses to accept as a general law the statement formulated by Turriazerr that the maximum photo- synthesis occurs under an intensity of psiaenian ome to renee one half that of direct insolation. By measuring the rate of phot hilous plants (Tilia and Abies) and of ombrophobous plants (Betula and Pinus) under both artificial and natural light he finds that plants differ as to the minimum insolation necessary to initiate photosynthesis. In this result the author finds basis for the conception of a specific quality of the chloroplasts. Further investigation coh- vinces him that the curve of photochemical work is determined primarily by the specific quality of the chloroplasts and by the anatomical structure of the leaf. The influence of the latter factor is particularly evident during periods of moderate sunshine, but the potency of the former is manifest under insolation of high or low intensity. The chloroplasts of the ombrophilous plants have greater dimensions and a sensibility almost five ne Lereaiet than that of ie ca ameaamne of ese ena plants. Other test te that the pig roplasts of the former. : While auxiliary data support the author’s main conclusion, the chance of error through imperfect technique and ignorance of all the factors is so great that final conclusions are better withheld—RayMonD H. Ponp. Microsporangia of Sphenopteris.—It has been suggested that the microspo- rangia of Lyginodendron Oldhamium =Sphenopteris (Crossotheca) Héninghausii might be found on Telangium Scotti. K1pston"s concludes from its structure that . Telangium Scotti cannot be the microsporangia of Sphenopteris Héninghausii. In no instance was organic connection between the two demonstrated. He found a few microsporangiate pinnae referable to Crossotheca Zeiller in organic connection with barren pinnae of Sphenopteris Héninghausit. The fertile pinnule is oval, entire, and attached to the rachis by a short pedicel, which is thick- ened very slightly upwards before merging into the pinnule, to which it seems to be united fora short distance. The pinnules appear to be rather thick, and the vascular bundle which enters the pinnule divides into two branches, which separate slightly from each other. Each fertile lobe bears six to eight broadly lanceo- late, sharply pointed, bilocular microsporangia, which in early stages bend inward, forming a small hemispherical bunch, with their apices meeting in the center, Later, the microsporangia spread outward and appear as a fringe hanging from the margin of the pinnule. The microspores are either slightly oval or circular and Measure 50 to 75 “ in diameter. The walls are roughened, being covered with minute blunt points. The tri-radial ridge, marking the line of cleavage of the tetrads, is sometimes apparent.—W. J. G. Lanp. 12LUBIMENKO, M. W., Sur la sensibilité de 1’ ig chlorophyllien des plantes ombrophiles et ombrophobes. «Rev. Gén. Bot. 17: 381-415. pls. 10, 11. 1905 ™3KipsTON, R., Preliminary note on the occurrence 3 microsporangia in organic connection with the foliage of Lyginodendron. Proc. Roy. Soc. B76: 358-360, pl. 6. 1905. 220 BOTANICAL GAZETTE [MARCH Photic sense organs.—GuTTENBERG' has demonstrated that two of the om- brophilous species of his local flora have a photo-sensitive epithelium, whose response consists in maintaining the leaf in the transverse heliotropic position. The mechanism is essentially the same as was found by HABERLANDT in the so- called velvet leaves, so abundant among the ombrophilous species of the tropical hydrophytic forests. The épidermal cells function as converging lenses, so that the protoplasmic membrane which covers the floor of the cell is not uniformly illuminated. In HABERLAND?I’s studies the bright spot was centrally located, but GUTTENBERG finds that for his species that it is excentric, because the papillosity is not centrally located. The result is the same in both cases, for the leaf is attuned to the distribution of interior illumination which exists when the leaf is in the transverse position. Actual tests showed that the petiole is not a factor in securing this position. Curiously enough the leaf assumes the horizontal position in diffuse light, such as occurs under the open sky on a cloudy day. In this cee, however, the internal distribution of light is the reverse of that which exists under parallel rays, the central area of the floor wall being dark with a margin of increasing brightness. The stimulus apparently consists in an unequal illumination of the cell lumen.—RAyMonpD H. Ponp. Nature of chromatophores.— MErEsCHKOWSKY'S holds that these bodies are not organs of the plant cell and never have been, but are foreign organisms which penetrated into the colorless plasma of the cells and live there as symbionts. In support of this notion he adduces the facts that the chromato- phores multiply continuously by division and do not arise de novo; that they are in high degree independent of the nucleus; that they are completely analogous with zoochlorellae and zooxanthellae which inhabit hydras, infusoria, etc.; that there are organisms, (e. g., the lower Cyanophyceae, such as Aphanocapsis and Microcystis) which can be considered as free-living chromatophores; that certain Cyanophyceae actually live as symbionts in the cell plasma. This theory he thinks, is the only possible explanation of the polyphyletic origin of primeval plants, which were merely amoebae and flagellates into which Cyanophyceae migrated; that the green, red, and brown Cyanophyceae account for the algae of these colors; that the plant cell-wall is due to the formation, by the symbiotic chloroplasts, of carbohydrates easily polymerized into cellulose ; which w. makes impossible the further taking of solid food and entails the quiescent nature and simple organization of plants, minus nerve, muscle, and psychic life. Here is another pyramid of theory resting on its apex.—C. R. B ‘4GUTTENBERG, H. R., von, Die Lichtsinnesorgane der Laubblatter von Adoxa Moschatellina und Cynocrambe prostrata. Ber. Deutsch. Bot. Gesells. 23:265-273- pls. 10, II. 1905. 5 MERESCHKOWSKy, C., Ueber Natur und Ursprung der Chromatophoren im OT en a Biol. Centralbl. 25: 593-604. 1905. * 1906] CURRENT LITERATURE 221 Idioblasts of Cruciferae. —ScHWEIDLER"® has decided to assign a systematic value to the peculiar idioblasts of the Cruciferae. The author at present reserves judgment as to their generic value, though this is expected to be established by further work. He has no doubt, however, that suborders and tribes can be accu- rately defined. On this basis he divides the family into three suborders. The first is characterized by the presence of idioblasts which contain chlorophyll and which are located exclusively in the mesophyll. The idioblasts of the second sub- order occur in the vascular tissue and differ from those of the first group in not containing chlorophyll. The third suborder is composed.of members which have both kinds of idioblasts. Just what would happen to the systematic standing of an individual so unfortunate as to have had the development of its idioblasts inhibited is certainly not for the reviewer to say, but in view of the urgent necessity of estab- lishing systematic work upon an experimental basis rather than morphological, it is difficult to escape the conviction that more or less futility is involved in all those efforts of which this paper is an example.—RayMonD H. Ponp. Araucarineae.—A preliminary note by THOMSON?’ states that in Agathis there are many supernumerary nuclei in the pollen tube and that in Araucaria as many as thirty were counted. The pollen tube grows along the surface of the ligule for 22“ or more before entering the micropyle. The anatomy of the ovule and development of the archegonia, as well as of the pollen tubes and megaspore membranes indicate that the Araucarineae occupy a very isolated position among © the Coniferales SEWARD and Forp in an abstract of a paper'® read before the Royal Society Dec. 14, 1905, indicate the scope of an extensive investigation of the Araucarieae. The section headings are: Introduction, distribution, diagnosis and synonymy, seedlings, root bis stem ATONE neers 7 presi rate reproductive shoots, fossils, and phylogen lusion The ions ‘Annee conclusion is that the group, unlike the Cycadales, has been derived from lycopodiaceous ancestors. The Araucarieae differ so greatly from the other Coniferales that the — suggest the substitution of the term, Araucariales for Araucarieae—CHARLES J. CHAMBE Inhibitory action —ERrerA’ suggests that the non-development of lateral branches or their growth in a particular position (e. g., of certain conifers) is determined by inhibitory stimuli (de nature catalysatrice si l’on veut) traversing either bark (Araucaria) or all living cells (Picea). We may conceive, he says, the apex of the stem or root asa sort of tyrant who forbids the subjacent 16SCHWEIDLER, J. H., Die systematische Bedeutung der Eiweiss- oder Myro- sinzellen der Cruciferen nebst Beitragen zu ihrer anatomisch-physiologischen Kennt- niss. Ber. Deutsch. Bot. Gesells. 23:274-285. pl. 1905. 17THOMSON, R. B., Preliminary note on the Araucarineae. Science 22:88. 1905. 3SEWARD, A. C., and Forp, Sibille, O., The Araucarieae, recent and extinct. TOERRERA, L., Conflicts de préséance et excitations inhibitoires chez les végétaux. Mém. Soc. Roy. Bot. Belgique 42 : 27-43- 3- Aug. 1905 222 BOTANICAL GAZETTE [MARCH branches to erect themselves or in other cases to develop, though they have the same tendency to do so as he; their geotropism or their power of growth is held in check by his own. Suppress the apex, let it die or become enfeebled, and the subjugated branches lift their heads. Several could erect themselves and take the lead, and that is sometimes observed. But ordinarily a new conflict for precedence occurs among the branches; the one nearest the apex or the most vigorous near one early asserts its supremacy and in its turn keepsits rivals at its feet. Cj. the independent and almost simultaneous amas of the like idea y McCattium, Bot. GAzETTE 40 : 262. Oct. 1905.—-C. R Ecological survey.—P 1 PRAEGER?° have a8 another vegeta- tion map and ecological description to the list of vegetation surveys of the British Isles. The area discussed lies south and west of Dublin. After a historical introduction the geology, physiography, floristics, and survey methods are briefly explained. The vegetation is divided primarily into littoral, agrarian, hill- pasture, and moorland zones, and the woodlands. The zones are further sub- divided into associations. These are described in detail and as far as possible related to the factors determining their occurrence. The text is accompanied by a map and five excellent plates of vegetation types. The paper will prove of especial interest to those who have followed the work of R. SmitH, W. SMITH, and Lewis in Scotland and England.—E. N. TRANSEAU. Alternation of generations in animals.—In criticism of CHAMBERLAIN’S paper on this subject?? Lyon?? holds that the phylogeny of animal gametes gives no evi- dence of their being reduced or vestigial generations, comparable with the gameto- phytic generation in plants; similarity of cytological processes does not prove identity of morphological value in the two cases. He refers to the alternation in Hydrozoa, and calls attention to the earlier proposal by BEARD and MuRRAY of a theory similar to CHAMBERLAIN’s. In reply CHAMBERLAIN maintains’$ that his -critic fails to distinguish between a gametophytic generation and a gametophytic plant. He holds that the generations in Hydrozoa do not alternate in the botanical sense, and points out that although reduction of the gamete-bearing generation has not been proved for animals, there is strong evidence for its aving occurred in plants—M. A. CHRYSLER. Mechanics of secretion.— PANTANELLI*4 has attempted to ascertain whether or not true secretion of enzymes occurs. He defines secretion as “‘the emission 2°PETHYBRIDGE, G. H. and PRAEGER, R. L., The vegetation of the district lying south of Dublin. Proc. Roy. Irish Acad. B. 25:124-180. 1905. 2tBoT. GAZETTE 39: 137-144. 1905. 22Lyon, H. L., Alternation of generations in animals. Science N. S. 21: 666-667. 05. ?3CHAMBERLAIN, C. J. Alternation of generations in animals. Science N. S. 22! 208-211. 1905. ?4PANTANELLI, E., Meccanismo di secrezione degli enzimi. Annali di Bot. 3: i13-142. 190 line 1906] CURRENT LITERATURE 223 of substances by living protoplasm, a thing possible through a self-regulated change in the condition of permeability of the plasmatic membranes such that the organism is able at pleasure to reverse it.” He finds that the ferment of Roman bread and Chianti wine truly secretes invertase, by the augmentation of the permeability of the protoplasm during the period of fermentative activity. is increased permeability is general, various salts escaping more freely at the same time. In Mucor stolonijer, however, the emission of invertase seems to have the character of a free exit of materials from dying parts, coincident with spore formation. Whether it has a true but weak secreting power remains for further study.—C. R. B Respiration.— PALLADIN distinguishess three sources of the respiratory CO. of plants: (1) nucleo-CO, produced by the action of enzymes, which, partly soluble, partly insoluble in expressed sap, are intimately bound up with the pro- toplasm; (2) stimulation-CO,, formed by the protoplasm itself (apparently directly) under the action of stimuli; (3) oxydase-CO., produced various oxidases. The process which characterizes animal and plant life consists in the excretion of nucleo-CO, which is formed by decomposition without the partici- pation of atmospheric oxygen. Intramolecular respiration is a primary phe- nomenon, whose CO, is principally nucleo-CO, and in some cases also stimulation-CO,. But alcoholic fermentation is no simple phenomenon, and, as KostyrscHew has shown, must be distinguished from intramolecular respira- tion.— ub an and pear rot.— LonGyEAR?® has published the results of his study of a rot of apples and pears due to an undescribed species of Alternaria. The same disease has so far been found in California, Colorado, and Michigan, and in Colorado is one of the most widely distributed and common diseases of apples. PappocxK?7 was the first to cali attention to it. In the case of the apple the disease attacks the fruit only, but it attacks the fruit, leaves, and young sprouts of the pear. In the apple it appears frequently first at the blossom end of the fruit and, in the case of sorts having a deep calyx-tube, a core-rot may occur. The disease may be controlled by spraying with Bordeaux mixture and plowing under or removing the diseased fruits in which the fungus is able to pass the winter.— E. Mrap WILcox. Reduction division.—The earliest enh were scat non- a seron, In phylogeny, according to SCHAFFNER,”® the conjugat 8 25 PALLADIN, W., Ueber den verschiedenen Ursprung der wahrend der Atmung der Pflanzen ausgeschiedenen Kohlensaure. Vorlaufige Mitteilung. Ber. Deutsch. Bot. Gesells. 20: 240-247. 1905. . 26 LoncyEar, B. O., A new apple rot. Bull. Col. Agric. Exp. Stat. ros: 1-12 pls. I-4. 1905. 27 PApDock, W., A new apple disease. Rept. Col. Exp. Stat. 17: 99. 1904 28 SCHAFFNER, JOHN H., The nature of the reduction division and related phe- nomena. Ohio Naturalist 5:331-340- 1905. 224 BOTANICAL GAZETTE [MARCH a disturbance into the life cycle and a reduction division of some kind became an inevitable accompaniment. The places at which a reduction division might, theo- retically, become established in the life cycle are presented in diagram and described. A comparison between the life cycles of plants and animals is also illustrated by a diagram. ScHAFFNER believes that in the higher animals the condition appears to be similar to that found in Fucus. The significance of a transverse division of chromosomes in interpreting the phenomena of MENDEL’s law is illustrated and discussed. — CHARLES Migration of salts— In an extensive investigation of the content of nitrogen, phosphoric acid, sodium, and potassium in cultivated plants, both field and pot grown, at different periods of their development, it has been found?9 that in different plants the maximum absorption is completed at different periods, bar- ley, spring wheat, peas, and mustard attaining this maximum at flowering, while potatoes reach it at maturity. These substances do not remain at a maximum, but in the plants other than potatoes and with the exception of phosphoric acid, migrate back, in great part, to the soil; this seems to depend on the amount of _ a given substance available, being greater san say, potassium is lacking than if the appropriate materials are all supplied —C. R. B. Anatomy and affinity — Another hese SARTON, has attempted to ascer- tain how much help is to be had from histology in determining the validity of Jordanian species as contrasted with Linnean.3° He studied allied plants, sub- mitted them to cultivation under diverse conditions and then examined their structure. In some cases there were constant anatomical characters distinguish- ing apparently closely allied forms. On the other hand the characters were as often elusive and evidently directly adaptive. Plants long cultivated in the Jardin des Plantes and at Fontainebleau showed no anatomical differences from wild ones of the same species. Nor were there differences between the varieties having different colored flowers.—C. R. B. Scotch moors.—T he succession of plants in the moors of the Scottish southern uplands has been studied by Lewis.3! He finds that in all the localities visited the peat “shows a definite stratification of plant remains, indicating a swing from woodland to heath and moss, and again to woodland. In some districts, an arctic plant-bed is interposed between the lower and upper woodland: beds.” The- vegetation changes are probably correlated with climatic changes at the 29 WILFaRTH, H., R6mer, H., and WIMMER, G: Veet die Nahrstoffaufnahme der Pflanzen in verschiedenen Zeiten ihres Wachstums. Landw. Versuchsstat. 63: 1-79 pls. 3. 1905. 3° Sarton, A., Recherches expérimentales sur l’anatomie des plantes affines- Ann. Sci. Nat. Bot. IX. 2 : 1-115. pls. I4. 1905 3t Lewis, F. J., The plant remains in the Scottish peat mosses. Pt.J. The Scottish southern chee Trans. Roy. Scc. Edin. 413:699-722. aintentaleteehinieeati 1906] CURRENT LITERATURE 225 close of the glacial period. He concludes also that the differences in the basal deposits of these moors as compared with those of the higher Cross Fell district (upon which he reported earlier) indicate the relative time of origin.— E. N TRANSEAU. Aberrant chromosomes.—The discovery of chromosomes of different sizes in the same nucleus in plants suggests that the attention of botanists be called to the terminology just proposed by MontcomeEry for aberrant chromosomes in Hemip- tera.3? The term chromosomes is retained when ‘all the chromosomes of a nucleus are alike; when they are unlike, the name autosoma or autosome is applied to a chromosome of the usual form, and allosoma or allosome to an aberrant chro- mosome. Unpaired allosomes are monosomes, and paired allosomes are diplo- somes.—CHARLES J. CHAMBERLAIN. Iron-algae.—After observation in the field and a study of cultures, GaIDUKov%3 concludes that a Conferva found by him in overflow pools of the Ocka river near Rjasan accumulates iron oxid from the waters, just as other algae do calcium carbonate or silica. He thinks such iron secretion not peculiar to the bacteria, but characteristic of many organisms, not as a necessary life- process, but as an adaptive one. In the present case it seems to be protective to the akinetes, which, eos down by the iron oxid, sink to the bottom and so pass the winter.—C. R. B Photosynthesis and reste. Poxiacct34 announces that electric energy, when it does not exceed a given intensity, promotes very much the formation of starch in leaves, and that this effect is greater with a continuous current passing directly into the interior of the organs. Electrified leaves almost deprived of light in some cases showed starch formation, when, in the same illumination, unelectrified leaves did not. In view of the recent English work on photosyn- thesis these conclusions should be received with reserve— C. R. B _ Formation of proteids.—MonTEMARTINI’S is attacking this much investigated problem. His first paper clears the ground, records once more a good part of the extensive bibliography, and details two experiments, which lead to the con- clusion that the production of proteids is greater in light than in darkness, and greater in light and air minus CO, than in light ‘and normal air. Likewise it is fivefold greater in the day than in the night, and he proposes to analyze the relation of light to this result in his later experiments.—C. R. B 32?MontTGoMERY, THos. H., The terminology of aberrant chromosomes and their behavior in certain Hemiptera. Science 23: 36-38. 1906. 33GarDUKov, N. , Ueber die Eisenalge Conferva und die ang aoa des Siisswassers in algemeinen. Ber. Deutsch. Bot. Gesells. 23 : 250-253- 1905. 34Pottacct, G., Influenza dell’ electricita sull’ assimilazione pastels Nota preliminare. Atti I t. Bot. Pavia II. 11: 7-10. 1905. SIL Da L., Primi studi sulla formazione delle sostanze albuminoidi nelle piante. Atti R. Ist. Bot. Pavia II. 10: 1-20. 1995. 226 BOTANICAL GAZETTE {MARCH Chemotaxis of sperms of Equisetum.—Liprorss,°° to avoid anticipation by SurBata, has made a preliminary announcement of his discovery that the sper- matozoids of Equisetum are markedly chemotactic toward solutions of malic acid especially, and also to maleic acid and its salts. Only indifference is shown to solutions of fumaric acid or of its salts. The threshold concentration of malic acid he finds to be about M/1oo00. Aerotaxis, which had been previously observed in the case of Marchantia spermatozoids, could not be demonstrated.—RaymonD H. Ponp. ‘ Welwitschia.—Tumboa mirabilis is so little known that any fresh observations are welcome. PEARSON37 succeeded in securing material showing the develop- ment of microsporangia, microspores, megasporangia and megaspores. Observa- tions were made upon the habit, habitat, and climatic conditions. It is probable that the plant is partially, if not wholly, insect-pollinated, and that the processes of fertilization and maturation of the seed take place more rapidly than in other gymnosperms.—CHARLES J. CHAMBERLAIN. The cycadean integument.—This is discussed in a recent paper by Miss Stoprs,38 who takes this occasion to compare the structures of the cycad ovule with those of the fossil Lagenostoma. The single integument of the living cycads is regarded as a double structure representing two integuments of some ancestral fo The plane of fusion of the two integuments has been between the inner and outer layers of the stony coat, or between the stone and the outer flesh. —CHARLES J. CHAMBERLAIN. A rust-resistant cantaloup.— Birnn3° finds that the Pollock strain of canta- loups is resistant to the rust or blight which is a common and serious disease in the Rocky Ford district of Colorado. This resistance he found was transmitted through seed selected from resistant plants, and hence seed selection becomes a very practical method of controlling this destructive disease wherever it may occur. The disease is due to the fungus Macrosporium cucum2rinum E. & E.— E. Meap Witcox. 3°LipForss, B., Ueber die Chemotaxis der Equisetum-Spermatozoiden. seg Deutsch. Bot. Coit 23: 314-316. 1905. 37PEARSON, H. H. W., Some observations on Welwitschia mirabilis Hooker. Abstract of a communication to the Royal Society of London, Nov. 23. 1905- 38SToPEs, MARIE C., On the double nature of the cycadean ne pee of Botany 19:561-566. 190 5. 39BLINN, P. K., A rust-resistant cantaloup. Bull. Col. Agric. Exp. Stat. 104: I-15. pls. I-IO. 1905. by oe ET he OS NEWS. Dr. JouNn W. HARSHBERGER is delivering a course of ten lectures on North American trees before the Wagner Free Institute of Science in Philadelphia Dr. Pear OLsson-SEFFER has left Leland Stanford University and bis ac- cepted a position in connection with an experiment station in Mexico, devoted to the investigation of the growing of rubber plants. His address, which corre- spondents are requested to note, will be La Zacualpa Botanical Station, Escuintla, Chiapas, Mexico. THE ANNOUNCEMENT for 1906 of the Lake Laboratory, maintained by the Ohio State University at Cedar Point, on Lake Erie, has been issued. The only instructor in botany for the season is Prof. MALcoim E. STICKNEY, assistant pro- fessor of botany, Denison University. ee, the courses are limited to one in general botany and one in ecology. Th opens June 25th, and closes August 3d. THE MINNESOTA SEASIDE Station on the Straits of Fuca, Vancouver Island, opens its doors for the sixth annual session, July 8, 1906. Owing to the low rates to the Pacific coast which will be in force, this promises to be an impor- tant year in the history of the Station. Those contemplating marine study and research are invited to write to Professor Conway MacMillan, University of Minnesota, Minneapolis, for the illustrated announcement of the Vancouver Island Laboratory-Camp. Dr. ALBERT SCHNEIDER has resigned his position as professor of botany, pharmacognosy, and materia medica at the California College of Pharmacy and has been appointed pathologist and physiologist of the Spreckels Sugar Com- pany. Dr, Henry B. Carey, formerly assistant to Dr. SCHNEIDFR, has been elected to fill the vacancy created by the latter’s resignation. DR. SCHNEIDER is now giving his entire time to the investigation of the so-estied California seas beet blight, which has been the cause of great losses to California beet grow NUAL announcement of the Marine Biological Laboratory at WL Hole, Mass., shows that the laboratories will open on July 5th, the regular courses of instruction extending from that date to August 16th. The depart- ment of botany for this year will be manned by Dr. Grorce T. Moore, of Washington, D. C., and Dr. James J. Wotre, of Trinity College, N.C. Miss J. Macrae will act as collector. Correspondence regarding botanical courses should be addressed to Dr. Moore at the Cosmos Club, Washington. THE Association internationale des Botanistes has shown commendable activity not only in the conduct of the Botanisches Centralblatt but also in arranging for a supply of pure cultures of fungi and alge. Now it further announces a long list 227 228 BOTANICAL GAZETTE [MARCH of places from which it is ready to supply material for demonstration or investi- gation to members of the society. The list is too long to be republished, but it » is evident that one can secure working materal from a wide range of localities. Correspondence relating to such material should be addressed to the secretary, Professor J. P. Lotsy, Leyden, Holland. BEGINNING with January 1, 1906, the form of the publications, which in the past have appeared as bulletins of the Bureau of Government Laboratories in the Philippines, will be changed to a journal to be known as the Philippine Journal of Science. This publication will include original articles by members of the staff of the Bureau of Science, as well as scientific papers submitted for _ publication by other officials of the Philippine government and by individuals not officially connected therewith. The journal will include researches in ieee zoology, chemistry (including physiological chemistry), serums and pro- phylaxis, mineralogy, geology, paleontology, mining, and mineral resources. The journal is to review work which is being accomplished and to present such original results as are obtained. The subscription price is $5 (U. S.) per year. It will be possible to secure reprints of any particular series of the articles at reduced prices. The journal will be edited by Dr. Paut C. FREER, director of the Bureau of Science, with Dr. RrcHArD P. Srronc, chief of the biological laboratory, and Mr. H. D. McCaskey, chief of the division of geology and mining, as co-editors. IN THE SUMMER of the present year a permanent Station for the study of arctic science will be established on the south coast of Disco Island in Danish West-Greenland. The cost of the foundation is defrayed by a gift from Mr. A. Hotcx, of Copenhagen, and the Danish government has promised an annual grant of 10,060 kroner ($3000) toward its maintenance. A laboratory, equipped with appliances and instruments especially for biological researches, will be attached to the Station, and for the present two work-places will be furnished for visiting naturalists. The visitors will have the free use of the instruments, seaeay outfit, and library of the Station. Lodging will be free and a small e will be charged only for board. The first visitors can be received in 1997, oa notices, ,inviting application, will be issued in due course. A library of arctic literature is to be founded at the Station and to be made as pan as possible, but on account of the limited resources of the Station and the vastness of the literature, only a small proportion of it can be purchased. The Director of the Station, M. P. Porsitp, asks botanists to be good enough to come to its assistance by giving to this library works on arctic and antarctic nature, and especially on arctic biology. The publications of the Station will be sent in return, and the Station will be glad to render any service in its power. Up to May 1 Director Porsirp may be addressed at Copenhagen S., Denmark. ‘ ee ee see Ne EE NERVOUS DISORDERS The nerves need a are supply of in apa to hebiok -them steady and deficiency of the et ie ts ais alow acc of nervous tone, indi- cated by exhaustion, sot eters head- ache or ‘ieee ia. Horsford’s Acid Phosphate (Non- Alcoholic.) furnishes the pad Shp in @& pure and abundant form. It supplies the sate cells w. ith. health-giyi ie life force, repairs waste, restores the stren and induces restful win without the use of anders ous drugs. An Ideal Tonic in Mecvods Distases, If your druggist can’t supply you we will send a iat size me batter prepaid, on receipt of 25 cents. 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MEDICAL OPINIONS OF BUFFALO LiTHiA WATER “All the Argument Necessary.” The International Journal of Surgery, August, 1905, under the heading - Shee lian says: ‘‘In the treatment of ae eyed . the e arene. et to all forms of medication. ide seh in whic to administer Moreover, BUEFALO LITHIA WATER ;; iM. = the Cystitic patient, as it is not only a pure solvent, but has the additional virtue of contadaleny substantial quantities * és, the Alkaline Lithates. Patients should be encouraged to take two quarts per per day, if panes they can, and the relief they will obtain will be all the argument PeCeRRALY. after first day or so.’ “The Results Satisfy Me of Its Extraordinary oa" Dr. Jas. Patt, of New Orleans, E Ex-President of the State Board of Health oe Louisiana, Pa oe bia ches of the | kidneys and **Lhave p ee y passages : Gouty subjects, in setaudttnies and in irritable satan of the Bladder and Urethra in females. The results satisfy me Ae its OPNESE Taine ina en of cases usually most difficult to treat.”’ “I Have Witnessed Decided BSenctictal wesidis: aes Its Use.” ia Wm. B. Towles, M. D., r rey Professor of Anatomy and Materia | Midian. _ the Sh Sntversity of Vir are marked in cesnng a beige - ginia: * The effects of ¢ BUFFALO pearance | of Albumin from the — be Rei and in Certain a of — Ss Disease £ hav e witnessed decided be nefi tis results from its use. e “PROPRIETOR 5 BUFFALO LITHIA SP O insure a healthful pom and prevent sick- Holds America’s Highest Prize ness, purify the cellar, closets sinks, dra dusty or damp corners ae pe sracks—nooks germs may emai, with Walter Baker & Co.'s THE HOUSEHOLS than Se cents a eh : =* pt * we ~ Jes a : na ° * 32am. orn. f aa a 63 65 67 E. W. OLIVE, DEL, HELIOTYPE CO., BOSTON. OLIVE on EMPUSA. 1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 261 Fic. 54. A stage in which the daughter-halves appear to be pressed upon by turgescent, vacuolar middle portion 55- A partly polar view of an aidiaidy placed nucleus, showing about 16 fibrillar radiations extending from the centrosomes. 1G. 56. A late condition in which the daughter-nuclei have just been sepa- rated from each other by a cytoplasmic constriction. The nuclear contents occupy a pseudo-synaptic position. Fic. 57. Showing in the upper portion of the hypha a resting nucleus and in pi gi a late stage of nuclear division G. 58. A double cytoplasmic constriction cm taken place so that a vacu- ole a with nuclear sap is left between the two daughter-nuclei. Fic. 59. A telophase condition. Fic. 60. Another telophase condition. Fic. 61. Probably a very late telophase, in which the dark rim of the center has survived, whereas the achromatic centrosome-portion has disappea 1G. 62. A late condition of nuclear division which is characteristic of ate gated cells. Fic. 63. A somewhat later stage, also from an elongated cell. Fic. 64. A younger stage, in which the nuclear membrane has been indented at the ends of the nucleus distally from the poles. Fic. 65. An elongated nucleus in division, in which one pole shows an inden- tation. Fics. 66, 67, Empusa aphidis. Xt1500. Fic. 66. A poorly stained mney showing the division character- istic of the prec cells of this Fic. 67. An earlier stage of ticato BIOLOGICAL RELATIONS OF DESERT SHRUBS. II. ABSORPTION OF WATER BY LEAVES. V. M. SPALDING. DurInc a study of certain shrubs growing in the vicinity of the Desert Botanical Laboratory near Tucson, Arizona, it has been found that the leaves of some of them absorb water, while those of others do not. Although leaf absorption is treated by leading physi- ologists as a matter of indifference, or at any rate of secondary importance, it has seemed worth while to inquire whether differences of habit in this particular, on the part of these desert plants, may not be correlated with other characteristic peculiarities; if so, even if the fact should turn out to be of small importance physiologically, it may be significant from a biological point of view. Our knowledge of leaf absorption as yet is fragmentary and uncertain. For the general subject it is quite unnecessary to cite the voluminous and contradictory literature. DANDENO’ has given a useful historical résumé, reference to which and to paragraphs in BURGERSTEIN’S more recent work? is sufficient for the present purpose. In regard to various highly modified plants, however, the case is quite different. ScHurmper has made such detailed observations of certain epiphytes as to leave no doubt that they - normally absorb large quantities of water through their aerial parts, and that this is a distinct physiological advantage, or even neces- sity. In view also of investigations cited by BURGENSTEIN it becomes necessary to accept the fact of leaf absorption in the case of various other plants. As for the plants of arid regions, the evidence has been less con- clusive than could be wished. VotkENs, in his classical work, describes various special structures by means of which, presumably, many of the plants of the Egyptian-Arabian desert take up dew * DANDENO, J. B. An investigation into the effects of water and aqueous solutions of some of the common inorganic substances on foliage leaves. Trans. Can. Inst. 75230. 1908. 2 BURGERSTEIN, A., Die Transpiration der Pflanzen. 1904. Botanical Gazette, vol. 41] [262 —S 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 263 through their leaves, but the woody species growing in the arid territory of the southwestern United States are so different in their habits and in their environmental relations as to preclude the settle- ment of the question for them, even within the bounds of probabil- ity, in any other way than by direct observation and experiment; in fact they are found, as regards-leaf absorption, to differ widely among themselves. The object of the present paper, therefore, is the presentation of such facts as have been determined for a lim- ited number of species indigenous to southern Arizona. In this region existing physical conditions give to the question special interest. As is well known, precipitation is meager, except at relatively high altitudes, and is distributed throughout the year, with no distinctively rainy season. The rainfall, moreover, is extremely uncertain, and for months at a time is often so slight that it does not wet the soil for more than a few centimeters, an amount of precipitation likely to be of very little positive advan- tage as far as root absorption is concerned. Under such circum- stances, in which delicate adjustment is the condition of survival, it would seem that plants capable of leaf absorption might have a distinct advantage in times of prolonged drouth, during which occa- sional showers occur which are too light to penetrate the soil. As will be seen, however, only a limited number of species appear to enjoy this advantage to an appreciable extent. Nearly all of the species selected for investigation grow in the immediate vicinity of the Desert Laboratory. A single one, Hola- _cantha Emoryi, which seems not to be indigenous here, was obtained from the grounds of the University of Arizona. The following classification of the plants employed into biological groups is pro- Visional, but will serve to direct attention to the very diverse eco- logical history of the species now growing together in this region. BIOLOGICAL CLASSIFICATION OF PLANTS STUDIED. I. Shrubs, with relatively slight modification of form and struc- ture, their habits plainly indicating mesophytic origin. Celtis, Covillea, Lycium. II. Shrubs or small trees, more conspicuously modified, but retaining manifest traces of mesophytic habits. Parkinsonia, Pro- sopis, Acacia. 264 BOTANICAL GAZETTE [APRIL III. Woody or partly herbaceous plants, exhibiting peculiar modifications of distinctly xerophytic types. Fouquieria, Hola- cantha, Koerberlinia, Zizyphus, Atriplex. IV. Plants of the most pronounced xerophytic character. Opun- tia, Cereus, and other cacti. V. Plants adapted by habit, rather than structure, to desert conditions. Sphaeralcea and many other half-shrubby or more or less herbaceous forms. Of the species employed in the experiments, Celtis pallida is a shrub, growing commonly to the height of one to one and one-half meters on the laboratory hill, where it is rather abundant. It holds its foliage so well that it might be ranked as an evergreen, though it suffers to some extent from the effects of frost. Its leaves are rough-hairy, thin but firm in texture, and conforming in general to the generic type. Covillea tridentata, the well-known creosote bush, is the most abundant woody species of this region. Its small coriaceous leaves, presented more or less edgewise to the sun and covered with waxy varnish, are well protected against excessive transpiration. Lycium Berlandieri is a small shrub, more than a meter in height, of frequent occurrence on rocky exposures. These species, of the three genera named, while well adapted to their habitat, exhibit characters far less conspicuously xerophytic than those of many of the plants with which they are associated. Coming to the second group, Parkinsonia Torreyana attains the dimensions of a small tree, and is conspicuous by reason of its green bark, from which it has the common name of palo verde. Though a denizen of the desert, it is not a dry ground form, but frequents low places, where more water is available than on the mesa or even on the adobe soil of the hills, where Parkinsonia micro- phylla, a related species, does well. Prosopis velutina, the mes- quite, grows chiefly in low ground, within reach of abundant water, but it also occurs, though scattering and undersized, on the adobe soil of rocky hills. Like the palo verde and many other legumi- nous plants, the leaves of the mesquite exhibit in their structure and position excellent adaptations for the prevention of excessive transpiration. Acacia constricta, of similar distribution, occurs on the mesa and also on rocky upland. It is a vigorous shrub, 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 265 one or two meters in height. These several species of Parkinsonia, Prosopis, and Acacia thrive well under the rather severe conditions to which they have become accustomed; they all retain, however, manifest traces of mesophytic habits, particularly in their choice of habitat. The species assigned to the third group, among which are Fou- quieria splendens, Holacantha Emoryi, Koerberlinia spinosa, and Zizyphus lycioides, present more striking modifications of form and structure than do any of the members of the preceding groups, and, though differing greatly among themselves, agree in possessing such conspicuous adaptations to xerophytic conditions as easily to rank next to members of the following biological group. The cacti are commonly taken to represent the extreme type of xerophytes, but notwithstanding various striking features com- mon to members of this order, there are essential differences of habit and adaptation, even between closely related species, ren- dering it quite impossible to generalize from the study of “typical forms” in the investigation of biological problems presented by them. The half-shrubby and herbaceous plants are much like those of other regions, exhibiting as a rule no structures that would be thought of as distinctively xerophytic, but accommodating them- selves to desert conditions by their habits, especially such as enable them to take advantage of periods favorable for rapid development and production of seeds. By way of first ascertaining whether any of the plants of these several groups absorb enough water through their leaves or inter- nodes to be readily detected by weighing, the following method was employed: A small branch with leaves functionally active, though often showing plainly the effects of long drouth, was severed and the cut end immediately covered with vaseline. In a few instances, which are specified, branches without leaves were used. The branch was then weighed and directly afterwards immersed in water, except at the cut end, for a definite time, usually about three hours. At the end of this period, after exposure to the open air long enough to be certain that the surface was fully dry, the branch was again weighed, and the increase of weight, if any, was 266 BOTANICAL GAZETTE [APRIL taken to represent closely the amount of water absorbed, though, owing to loss during the operation of drying the surface, the amount absorbed must often have been rather greater than the increase of weight indicated. In the first preliminary set of observations, a pair of large balances, weighing satisfactorily to ten milligrams, was employed; but in subsequent experiments quantitative bal- ances were used, the weighing being made to a milligram in each case. Changes during the process of weighing rendered it as useless as it was unnecessary to attempt a higher degree of accuracy. Inspection of Table I shows that leafy shoots of Celtis, Covil- lea, and Lycium, by immersion in water for three hours, gained 1.9 to 5 per cent. of their original weight; Atriplex in a little longer TABLE I. PRELIMINARY TEST OF CAPACITY FOR ABSORPTION. November 1904. Species Date 7 Time baton ig = esa Covillea tridentata (1)......... Nov. 1 10:12 A, M. 18.540 ; : 1:12 P.M, 19.250 | 3.8 gain Covillea tridentata (2) ........ i 9 10:55 A.M. 8.235 : : 1:56 P.M. So6ca ) fossa 3s RMN IME NOM ss 0g 9 ld san wc 5 I 10:57 A.M. 24.920 : . 10:41 A.M. 8.650 ’ ; 1:40 P. M. 8.670 | 0.2 “ Encelia farinosa..... 2.05556... < 9 12:00 M 5.010 3:00 P.M Go PaaS 3:18 P.M 5-000 Sphaeralcea pedata........... meer ae 12:07 P.M 1.120 3:12° P.M i.e besa 3-21 P.M 1.110 2 sinscmmoneas ini 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 267 period gained 5.1 per cent.; Parkinsonia, Prosopis, and Acacia gained 0.4 to 1.5 per cent.; iil Zizyphus, Fouquieria, and Koer- berlinia, all without leaves, showed almost no appreciable gain. Species of Sphaeralcea and Encelia gained in weight 5.4 and 6.2 per cent. respectively, but promptly lost all they had gained by a few minutes drying. It will be noticed that of the plants employed in this prelimi- nary work those without leaves absorbed no water to speak of, while those in leaf fell into two categories, those absorbing and those not absorbing water in quantity. The experiment, there- fore, pointed to leaves rather than internodes as agents of absorption, and indicated, apart from Sphaeralcea, Encelia, and the peculiar Atriplex, only the woody species belonging to the first group as likely to prove capable of absorbing much water. Starting with the suggestions derived from these facts, a more careful and detailed study was undertaken. Cut shoots were still employed for a time, though it was understood that confirmation of results would necessitate the use of entire plants, and these were, as a matter of fact, employed to a large extent in the later work. Care was exercised in the selection of material, and in each case its source and any conditions liable to affect results were noted. GROUP I. Celtis pallida. Four specimens of this species were selected, all in good condi- tion, though apparently not as active physiologically as they would have been earlier in the year. Numbers 1 and 2 were fresh shoots, while numbers 3 and 4 were small branches taken from older bushes. Those numbered 1 and 3 were cut so as to include a large leaf sur- face as compared with the other two. In every case the cut ends were covered at once with vaseline, and the first weighing was made as soon as practicable after bringing them to the laboratory. They were then wet at frequent intervals for a little more than three hours, and; after drying the surface, were weighed again, after which they were left in the laboratory to dry until the next day, when the same steps were repeated. Finally they were immersed in water over night and again weighed. 268 BOTANICAL GAZETTE [APRIL TABLE II. CELTIS PALLIDA. December 1904. No.| Date Time bo nvcmcig ges Period of treatment ~ 1 | Dec. 19 | 10:44 A.M. | 1.834 £255 P.M. | 1.853 | 2 gain After wetting mt 3 iy Ir min. 20 | 9:48 A.M. | 1.516 [18.2 loss ng tg hrs. 53 m 2250 P.M: | 1.997 )| 5 - gain “ wetting nearly Ma hrs. 22 min. ar || 0535 A.M. | ¥.895 (7 3 ca 25 2 Ig | 10:54 A.M. | 1.392 2:02 P.M. | 1.410 | 1.3 gain After wetting nearly 3 hrs. - min. 20 | 9:57 A.M. | r.192.|15.5 loss drying t9 hrs. 55 mi 2:25 P.M. | 1.238 | 3.9 gain oh wetting nearly 4 hrs. 28 min. 21 | 10:52 A.M. | 1.440 |16.3 gain : 27 3 ¥ | 12213 A.M. | 2.137 2:20 P.M. | 2.185 | 2.2 gain After wetting nearly 3 hrs. A min. 20 | 0:07 A.M. | 1.734 |20.6 loss 1g hrs. 47 m 2:38 P.M. | 1.870 | 7.8 gain * wetting seed 7 hrs. - ‘min. at | 11:05 A.M. | 2.182 |16.7 “ rs 7 4 IQ | 11:25 A.M. | 1.505 - 2°30 -P.M..| ¥.5§21-| 4.1 gain After pata ad 4 ma 6 min. 20 | 10:16 A.M. | 1.351 |11.2 loss Ts. 45 m 2:46 P.M. | 1.387 | 2.7 gain 3 wetting ‘nearly = hrs. 30 min. SE.) 067564 Me. | oeaS ese - 29 Inspection of Table IE shows: 1. That all four specimens absorbed water very slowly just after they were freshly cut, and that the rate of absorption was greatly increased after they had lost weight by remaining over night in the dry air of the laboratory. 2. The rate of absorption showed a correlation with extent of leaf surface, being considerably greater in the two specimens with large extent of leaf surface than in the other two. 3. The weight lost by drying for a given period was nearly or quite regained when the leaves were given a full supply of water for a corresponding length of time. The capacity of this species for leaf absorption, under the conditions described, is thus fully demonstrated. Its deportment in the seedling stage, which offered for experiment perfectly fresh and unmutilated material, will next be considered. Seedlings of Celtis pallida were grown from seeds sown Novem- ber 14, 1904. When used for experiment in January and February, 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 269 1905, they were all in healthy condition, and when taken up were found to have fine vigorous roots. In addition to the cotyledons, which were still capable of photosynthesis, each seedling had two or three perfectly healthy green leaves that had attained the length of about one centimeter. The seedlings were transplanted into earth contained in glass vials of convenient size for accurate weigh- ing, sheet rubber being used to prevent evaporation from the soil. In the case of seedling number 1 the earth was very moist when the rubber was adjusted, and it was found that this plant, which was transpiring vigorously, showed almost no capacity for absorption. The case was different with seedling number 2, which was left some five days after transplanting with the soil open to the air, so that it became relatively dry before the rubber was adjusted. The coty- ledons of number 1 were removed, their place of attachment being carefully covered with vaseline; number 2 had one large cotyledon which remained in place during the experiment. These details are necessary to an understanding of the different behavior of the two seedlings as shown by Tables III and IV, which cover the period from January 21 to February 1, at which latter date the experiment _ Was concluded. It is seen that both seedlings transpired regularly and largely, but that number 1, in spite of the fact that its transpiring surface had been lessened by the loss of its cotyledons, exhibited a decidedly higher rate of transpiration than number 2, which was in drier TABLE Iii. CELTIS PALLIDA. SEEDLING No.1. January 1905. Date Time Weight in! 7 o¢s or gain Conditions grams Jan. 21 | 1:19 P.M. | 26.256 seit ; 4:05 26.229 | 0.027loss_ | After standing in dry air 23 | 10:15 A.M. | 26. PoE 5, ed Ps fe yet, Mer, 2:50 P.M. | 26.061 | 0.033 “ ~ Pw ee 25 10:50 A. M. 25.936 0.125 ce “ce “ce “ec oe “cc 26 9.50 2 ;.870 0.066 “c ee &e tS te ee 27 9:28 25.806 0.064 “ “ “ (2 ied | eae 2:53 P.M. | 25.810 | 0.004 gain After wetting i 3:41 25.803 | 0.007 loss standing i in dry air 28 | 11:02 A.M. | 25.762 | 0.041 “ x gulp eae ba 13:33 P.M. | 25.764 | 0.002 gain ns ng f 2:58 25.759 | 0.005 loss ie) cia divaie 270 BOTANICAL GAZETTE [APRIL TABLE IV. CELTIS PALLIDA. SEEDLING No. 2. January and February 1905. Date Time ging Loss or gain Conditions Jan. 30 | 10:09 A 22.638 1:10 P.M. | 22.630 | 0.008 loss After standing in dry air 3:40 22.626 0.004 “< “ce ““ “ «@ 30)" O245 A: Me |) 22659 | 0.027 gain “wetting 12:45 P.M. | 22.628 | 0.025 loss ‘** standing in dry air gi 22.634 | 0.006 gain “wetting Feb. 1 | 9:48 A.M. | 22.592 | 0.042 loss ast ng in dry air 9:50 0.027 Weight of plant above ground 2:08 P.M. | 0.045 | o.o28 gain | After wetting - soil, an interesting result in harmony with earlier experiments, showing the direct relation between available soil water and rate of transpiration.3 On the other hand, while the quantity of water absorbed by: number I was so meager as to be negligible, that absorbed by num- ber 2 was much more, in one case almost exactly 100 per cent. of its own weight, i. e., of the part above ground when it was after- wards severed from the root. Number 2, although apparently perfectly healthy while the work was in progress, seems neverthe- less to have reached a condition in which the diminished supply of water from the soil was followed by a marked acceleration of leaf absorption, while in the case of number 1, growing as it was in moist soil, no such compensation was made or required. Of interest as bearing on the validity of determinations of absorp- tion by the use of detached shoots is the fact that while seedling number 2, after it had finally been cut off at the surface of the ground, absorbed in a few hours its own weight of water, it had done pre- cisely the same thing before mutilation, only in longer time. It may well be that a detached shoot, cut off from its normal source of water supply, will absorb more rapidly through its leaves than the same shoot, which, while attached, is supplied, even inade- quately, from the soil; but this difference plainly does not justify the degree of discredit that has been thrown upon evidence derived from experiments with separated parts of plants. Three other seedlings of Celtis pallida were treated like the preceding ones, except that the observations were not begun until 3 SPALDING, V. M., Soil water in relation to transpiration. Torreya 5:25. 1905- 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 270 the plants had been some weeks in the vials to which they were transplanted, and weighings were made during a longer period and with more complete records as to soil conditions, health of seed- ling, etc. Their records as to weight are given in Tables V, VI, and VII. All of these seedlings were in a healthy condition and apparently capable of entirely normal development. The small extent of surface through which absorption and transpiration took place renders the consistency of the results all the more striking. In every case water was absorbed when it was presented to the leaves and internodes, and transpiration was resumed as soon as their surfaces were dried. TABLE V. CELTIS PALLIDA. SEEDLING No. 3. February 1905. Date Time Weight in Loss or gain Conditions grams Feb. 14 | 11:20 A.M. | 23.084 Weight of vi a and outfit 2:25 P.M. | 23.091 | 0.007 gain After i immersion raped water 15 -M. | 23.044 | 0.047 loss standin aii dry ai T1755 23-053 | 0.009 gain 5 frametsioa! in rain qa 16 | 11:42 A.M. | 23.007 | 0.046 loss “standing in dry air 17 | 2:11 P.M. | 23.046 | 0.039 gain immersion in rain water 20 | 10:23 A.M. | 22.895 | 0.151 loss ‘standing in dry air 2:51 P.M. | 22.916 | 0.021 gain n immersion in rain water 21 | 10:28 A.M. | 22.0932 s0x6., rs 12:00 M. 22.907 | 0.025 loss 8 standing in dry air 3:02: Pi M. | 22.887 | 0.020" © se ae 22 {| 12:08 22.845 | @.052 * - - : T2:145 22.828 Weight oe — cotyledon act 22.8 0.007 gain | After immersing in rain water a? a O. oe Weight ar cut is pares of earth 2228 22.799 4 , rubber, and ea i 25 1:58 0.015 * of plant above ground, air- dried in laboratory When the first weighing was made, February 14, seedling no. 3 had two cotyledons, still attached, and three foliage leaves. The cotyledons showed some indications of drying. The earth in the glass vial in which the seedling was growing was becoming rather dry, but still contained sufficient water to maintain a transpiration current for a week and probably longer. On February 16, the note was made “one of the cotyledons drying, curled, and getting stiff; the other paler than the foliage leaves, but still flexible, other- wise the seedling is in good condition.” On February 22 the cotyledons were removed and the subse- ae | { 272 BOTANICAL GAZETTE [APRIL quent deportment of the seedling indicates that their failing condi- tion previous to removal may be disregarded as not materially affecting the results. As the table shows, the weight of the whole plant above ground, including cotyledons, was less than the weight F of water transpired in 19 hours (Feb. 14-15) and also less than the gain of weight by absorption in 26 hours (Feb. 16-17), a conclusive proof of the relatively large quantities of water absorbed and trans- pired by this seedling during the period of experimentation. The facts regarding seedlings 4 and 5 are so fully set forth in Tables VI and VII as to render further explanation unnecessary. | TABLE VI. | CELTIS PALLIDA. SEEDLING No.4. February and March 1905. Date Time braces in| Loss or gain Conditions Feb. 14 | 3:30 P.M. | 20.130 Weight of plant and outfit I5 | 10:02 A.M. | 20.090 | 0.040 loss After spent in dry air 12:47 P.M. | 20.093 | 0.003 gain immersion in rain water 16 | 12:00 M. 20.031 | 0.062 loss by saul in n dry air 17 | 3:11 P.M. | 20.051 | 0.020 gain ‘* immersion in rain water 20 | 10:46 A.M. | 19.821 | 0.230 loss «standing in 3:19 P.M. | 19.825 | 0.004 gain eC Sats mersion in rain water 21 | 10:54 A.M. | 19.830 | 0.005 “ 4 12:11 P.M. | 19.813 | 0.017 loss = standing i in dry air 22° tasdo 19.696 | 0.117 “ os 3:08 19.707 | 0.011 gain a mersion in rain water March2 | 10:04 A.M | 19.080 | 0.627 loss is saad in dry air 2:04 P.M. | 19.085 | 0.005 gain a a in rain water 2225 19.080 | 0.005 loss “* standing in 2:30 0.070 Weight rakes cut Biss at surface of rth TABLE VII. CELTIS PALLIDA. SEEDLING No.5. February 1905. Date Time igo Loss or gain Conditions Feb. 20 | 11:16 A.M. | 20.207 Weight of plant and outfit 3:28 P.M. | 20.204 | 0.003loss | After wetting with rain water (not immersin: 2I | If:og A.M. | 20.219 | 0.015 gain | After immersion in rain water 12:15 P.M. | 20.214 | 0.005 loss af standing in dry air ~ 22 12:34 20.172 0.042 oe ce té- Gt ce 3:18 20.178 | 0.006 gain in rain wate 3*33 0.025 Weight mye inky ean off at mates of 2G.) @txe 0.012 Weight after air drying 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 273 Covillea tridentata. The specimens of creosote bush selected for experiment were taken from four different sources, for the sake of securing material as different as practicable in regard to the amount of water in the tissues. Number 1 was from a bush growing near an irrigating ditch, where it had been abundantly supplied with water. Its leaves were large, dark green, and fresh, and numerous flower buds had been formed. Number 2 was from a plant growing on the mesa a few rods distant. Its leaves were smaller and lighter green, and in comparison with number 1 it was plainly a dry ground form, though it did not have the appearance of having suffered to any great extent from lack of water. Numbers 3 and 4 were from plants growing on the mesa, near the foot of the laboratory hill, where in a dry time the Covillea, the only shrub that keeps alive there, shows the effects of drouth very badly. Their leaves were still smaller and paler in color, those of number 4 especially, indicating by their minute size and other peculiarities a plant that had long lacked a sufficient supply of water. The contrast between this and the first member of the series was very striking. It should be stated, however, that none of the specimens were in quite so dried-up a condition as those employed early in November before the Decem- ber rains, which though meager—o.82 inch (21™™) thus far— had freshened vegetation to some extent. The dried-up leaves that were dying in November had been shed, and the leaves remain- ing on the bushes when the experiment was conducted, late in Decem- ber, were apparently in a vitally active condition. It will be noticed by reference to Table VIII that, precisely as in the case of Celtis, all the specimens of Covillea gained very little in weight as the result of wetting soon after they were cut. Num- ber 1, from the irrigated bush, gained least, and number 4, from the dry ground plant, gained most. After prolonged drying and again wetting, the gain was much greater than before, the greater gain in each case being made by number 4, which, as already stated, was from the most distinctively dry ground form. The deportment of number 1, from the robust, well-watered bush, is instructive, especially as it may throw light on the question as to whether leaf absorption is a normal process that takes place 274 BOTANICAL GAZETTE [APRIL TABLE VIII. COVILLEA TRIDENTATA. December 1904. No.| Date Time bears ie eo Samy Period of treatment 1 | Dec. 26 | 11:05 A.M. | 2.529 2217 P:M. |2.5383 | 6,4 gain After wetting ay 3 na 12 min. 28 | 10:32 A.M. | 1.957 {22.9 loss drying 4 2:18 P.M. | 1.996 | 2.0 gai = wetting nearly 3 hrs. e min. 29|| IorSo A.M. | 2.388 | 9.6 “ es 3 2 26 | 11:12 A.M. | 2.434 : 2:35 P.M. | 2.453 | 0.8 gain | After wetting nearly 3 hrs. 23 min. 28 | 10:40 A.M. | 2.234 | 8.9 loss ** drying 44 hrs. 05 min. : 2733 PM. | 2.258.| 1.1 gain “wetting nearly 3 hrs. 53 min. 20: | FR203 Ai M. | 2-416-| F.0 “* if oo 2. ga * 3 26 | 11:21 A.M. | 2.261 2:58 P.M. | 2.281 | 0.8 gain After wetting nearly MA hrs. 37 min. 28 | 10:48 A.M. | 1.980 [13.2 loss drying 43 hrs min. 2:45 P.M. | 2.020 | 2.0 gain sa wetting nearly 3 “hrs. : min. 29 | i1:14 A.M. | 2.225 a es © 4 26 | 11:28 A.M. | 2.646 3:03 P.M. | 2.688 | 1.6 gain After wetting nearly 3 hrs. 35 min. 28 | 10:55 A.M. | 2.357 |r2.3 loss drying 43 hrs. 52 min. 2:56 P.M. | 2.415 | 2.5 gain ‘“* wetting nearly 4 hrs. 1 min. 2Q | 11:23 A.M. | 2.766 |14.5 “ is ee = ae ae under natural conditions. This shoot, with its large, fresh, turgid leaves, lost water by drying approximately twice as rapidly as did _ those from dry ground, with their much smaller leaves and firmer tissues, and on subsequent wetting absorbed far less than the latter in proportion to previous loss. Unlike these, moreover, the leaves of the first specimen, in the course of alternate drying and wetting, lost their fresh look and became discolored. The impres- sion was received that this specimen, taken from a perfectly fresh plant and requiring no additional supply of water, suffered patho- logical changes in the course of the treatment to which it was sub- jected, while the others, coming from dry ground plants in need of water, absorbed it as by a perfectly normal process. Even these, however, were not in a condition for rapid leaf absorption when first cut, their gain per cent. being decidedly less for a given period than that exhibited by individuals of the same species during observations made before the December rains. In brief, the experi- ments of December 26-29, in connection with those of November 1-9, indicate on the part of the creosote bush marked capacity for 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 275 subaerial absorption after protracted drouth, but more limited capacity for such absorption, even if artificial drying is resorted to, when it is receiving a better supply of water. Lycium Berlandieri. At the time when the observations on Lycium were made, late in December, most of the summer leaves had fallen and fresh ones, following recent light rains, were only beginning to appear. Con- sequently it was difficult to secure entirely satisfactory material, but a few specimens were finally obtained for experiment which were in a normal and active condition. The leaves of this species are small, only about one centimeter in length, but otherwise the plant gives the impression, as already stated, of having retained up to the. present time distinctively mesophytic tendencies. The rapidity with which the leaves were transpiring was at once obvious when weighing was undertaken, and, as in cases previously cited, absorption was found to take place extremely slowly while the leaves were still fresh. Even after drying, water was absorbed in no case as rapidly as it had been lost. Thus number 2 lost 3 per cent. of its weight by drying three and one-half hours, and gained after- wards by wetting nearly four hours 1.9 per cent. Of the actual capacity of this species for subaerial absorption the experiments leave no room for doubt; but the.specimens employed deported themselves much like the well-watered Covillea, except that no suggestion of pathological change in the course of the treatment to which they were subjected was noted. When gathered they were simply in the condition of fresh, actively transpiring plants, TABLE IX. Lycrum BERLANDIERI. December 1904. No.| Date Time Iyer ge sl i Period of treatment 1 | Dec. 28 | 10:15 A.M. | 3.065 1:47 P.M. | 3.076 | 0.4 gain After ee aes? nearly 3 3 32 min. AQ: NOTE E Ac M+[ 32235. |-5 22” paris 20 hrs. 28 m 1:42 P.M. | 2.956 | 8.7loss | ‘“‘ drying 3 hrs ‘ 3:25 3-002 | 1.6gain; “ wetting cue I ae 43 min. 2 28:).107 92 A: Mi. | si ¥a9 E°G0 POM. |-3.028 | 370 loss After drying 3 hrs. * min. 29: | 10:00 A,M. | 2.602 |. 7-5 ° 1:53 P.M. | 2.855 | 1.9 gain “wetting nearly : re 53 min. 276 BOTANICAL GAZETTE [APRIL which apparently could: derive no advantage from an additional supply of water presented to their leaves. The record of these three species of Celtis, Covillea, and Lycium has been given at length, on account of the importance of estab- lishing beyond doubt the fact that in these plants, which have been taken to represent desert species that retain in structure and habits obvious indications of mesophytic origin, leaf absorption certainly takes place, and apparently as an entirely normal process. We have next to deal with a group of species genetically related, which deport themselves quite differently from members of the first bio- logical group in regard to leaf absorption. As representatives of this second group, species of Parkinsonia, Prosopis, and Acacia were selected, all belonging to the Leguminosae. The record of experi- ments and their results is such as to admit of statement in few words. GROUP II. Parkinsonia Torreyana. The specimens of palo verde employed in this work were seed- lings some two months old. One was cut about thirteen hours, the other (number 2) an hour and a half before weighing. After weighing an attempt was made to wet the leaves by repeatedly immersing the seedlings in water. The experiment might fairly have been dropped at this point, since, as it was found impossible to wet them, leaf absorption could hardly be thought of; but as there remained a possibility of some slight absorption where drops of water collected on the surface of the youngest parts, the attempt was continued with number 1, which was repeatedly immersed during a period of something over three hours. As seen from Table X this seedling, so far from gaining by absorp- tion of water presented to it, actually lost 1.8 per cent. of its weight in three hours and thirteen minutes, its surface having remained almost entirely unwetted, so that loss of water was possible during the whole, or nearly the whole, of this period. Seedling number 2 was allowed to dry, after an unsuccessful series of attempts to wet its surface. Its loss of weight, as might be expected, was greater than that of number 1. If these results are compared with those of November 1 and 9, derived from similar experiments with shoots of Parkinsonia micro- 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 277 ABLE X. PARKINSONIA TORREYANA. SEEDLINGS. December 1904. . Weight in|Loss or gain Boa No.| Date | Time | erin per cent. Conditions t | Dee. 31 | -g232 AM. | 0.901 12:45 P.M. | 0.944 | 1.8 loss | After repeatedly immersing in water 2 9.37 A.M. | 0.886 gtd drying, aera! be gees at- 5 tempts to i the s 12°52 P.M. | Ovd55 | 3.5408 phylla and P. Torreyana, the conclusion must be drawn that the species of Parkinsonia represented here either absorb no water, or at most an exceedingly small quantity, through their leaves. Before making the experimental test it was thought that the fresh, rapidly transpiring leaves of seedlings might exhibit a capacity for absorption not shown by those of older plants, but this has not proven to be the case. Prosopis velutina. Work on the mesquite was carried on at intervals for a number of weeks in January, February, and March, the material first em- ployed being obtained from mature specimens, while in the later experiments seedlings were used. Of specimens taken from mature plants only the leaves were immersed in water. In some cases the upper surface resisted wetting, while in others both surfaces were easily wetted. This was followed, as indicated by some increase of weight, by absorption of water in limited quantities. The seed- lings which were employed in subsequent experiments remained unwetted in all cases when they were immersed in water, and in spite of the fact that two of the specimens had been left to dry as much as forty-two hours and showed the effects of this treatment before im- mersion, there is no evidence that they absorbed any water whatever. It is apparent, then, that as long as the leaves of the mesquite are perfect and resist wetting they absorb no water, even after dry- ing for some time, but that they may absorb more or less after they have become old and can be wetted. It is very questionable, to say the least, whether in the latter case this process has any physi- ological significance. It would seem that in the mesquite, as in the palo verde, adaptations to xerophytic conditions have been carried so far in the direction of preventing excessive transpiration that leaf absorption, as a normal process, does not take place. 278 BOTANICAL GAZETTE [APRIL Acacia constricta. A series of experiments with this species was carried out, but it was found unfavorable for exact results, owing in part to the fact that its leaflets become tightly closed after wetting, thus rend- dering it difficult to secure perfect drying of the surface without overexposure and consequent uncertainty as to the true weight. Accordingly, the conviction that the data obtained were unreliable led to their rejection. For this second group, therefore, we are restricted to the positive results obtained from Parkinsonia and Prosopis, which exhibit either no capacity or very slight capacity for leaf absorption, so long as the leaves are in perfect condition and normally active. . GROUP III. This third group includes representatives of a number of genera much modified in form and structure, and differing among them- selves in their methods of meeting desert conditions. Several of these are more commonly seen without ‘than with leaves, photo- synthesis then taking place in their green shoots; while others, more dependent on leaf activity, are commonly in a leafless condition during a large part of the year, pushing out new leaves promptly when conditions are favorable, and dropping them again when they become adverse, .as is seen particularly in the case of Fouquieria. Holacantha Emoryi. Of this peculiar shrub a small branch with leaves was. cut and left several hours to dry. At the end of this time it was still fresh, with no indication of wilting. After weighing it was wet for two hours and thirty-nine minutes, after which it was weighed again, the weight remaining unchanged. Leaving the shoot now to dry until the next day, and then wetting it for four hours and twelve minutes, ' there was a gain in weight of only o.5 percent. Part of the same shoot, destitute of leaves, was treated in the same way, and, after wetting four hours and eleven minutes, also showed a gain of barely ©.5 per cent. of its former weight. These results indicate on the part of this species capacity for leaf absorption so inconsiderable that it may be neglected. Koerberlinia spinosa, a closely related species, agrees with Hola- ‘cantha as far as observations have yet been made. Only leafless 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 279 branches have been obtainable, but these, as in the preceding species, are green, and for a large part of the year the plant has no other organ of photosynthesis. So far, then, as present evidence goes, absorption through leaves or internodes is not to be predicated of either of these plants. Zizyphus lycioides. After the preliminary experiments already recorded, very little satisfactory material for the study of this species was obtainable, as the plant cast its leaves and remained bare until after the period of study was concluded. From observations made early in the year, however, it appears that leafless shoots of Zizyphus do not absorb water in appreciable quantity, but that leafy shoots have considerable absorptive capacity, indicating that it is the leaves and not the internodes through which absorption takes place. Fouquieria splendens. Leafy shoots of the ocotillo, as shown by Table XI, absorb con- siderable water when wet for some time after drying. As in various other cases, the loss of weight on drying the shoots after wetting is considerably more rapid than the preceding gain by absorption. TABLE XI. FOUQUIERIA SPLENDENS. January I905. No.| Date Time Weight in/I per pl as Conditions I} Jan. 26 | 12:00 M. 1.614 In each case loss followed drying and gain followed wetting the sp-eci mens during the periods indicated in the time column. 27 | 10:00 A.M. | 1.582 | 1.9 loss 28 | 11:40 1.664 | 5.2 gain 3:20 P.M. | 1.598 | 4.0 loss 3° | zio2 Ta§I6. 500" 3:28 1.530 | 0.8 gain 4:09 1.523 | 0.5 loss 31 | 4:15 1.490 Reb. = £532 1.585 2 | Jan. 26 2:53 POM. 4s2t4 27 | to:1§ A.M. | 4.112 | 2.4 loss 28 | 11:45 4.059 | 1.3 loss 30 | 2:04 P.M. | 4.363 | 7.5 gain 3:04 4-255 2.5 loss 4:02 4.197 | 1.4 Biol tae ACM: | S461, rea: 77 2:45 P.M. | 3.840 | 2.1 gain 3245 3-798 | 1.1 loss 280 BOTANICAL GAZETTE [APRIL It appears, then, that for this biological group, which includes a number of plants for the most part unrelated systematically, no general rule can be formulated regarding leaf absorption. The experiments go: to show that Holacantha and Koerberlinia hardly absorb at all, while Fouquieria is certainly capable of absorbing considerable quantities of water. GROUP IV. The only representatives of the cacti that have been studied thus far are two species of Opuntia, O. Engelmanni and O. ver- sicolor, A number of specimens of each species were selected after a prolonged drouth, the results of which were plainly seen in their much shrunken condition, very favorable, it would seem, for the demonstration of absorption if this ever takes place. As the mate- rial was rather bulky the large balances were employed, a terminal joint in each case being severed and weighed; but there is no reason to doubt the substantial accuracy of the results. As shown by Tables XII and XIII, Opuntia Engelmanni did not in any case gain more than 0.6 per cent. of its original weight, even when immersed in water upwards of 45 hours, and it is safe to conclude from this result, drawn from experiments with a num- ber of specimens, that the species in question does not normally absorb any considerable quantity of water in this way. Opuntia versicolor, on the other hand, treated in the same manner, showed TABLE XII. OPUNTIA ENGELMANNI. December 1904. No.| Date Time Weight in| pe st ocieed Conditions 1 | Dec. 3 | 10:15 A.M. | 82.360 In each case gain followed wetting ad loss followed drying rs, = oan indicated a the 1:30 P.M. | 82.850] 0.6 gain §.| 18:00 A.M. | 83: 360] 0.6 2 6 | 10:12 A.M. | 72.085 I:10 P.M. | 72.470] 0.5 gain 3:05 72.080] 0.5 loss 3 10:39 A.M. |131.750 1:33 P.M. |132.290) 0.4 gain 3:20 131.760] 0.4 loss ra . 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 281 TABLE XIII. OPUNTIA VERSICOLOR. December 1904. . | Weight in| Loss or gain’ de No.| Date Time Pisano per cent. Conditions 1 | Dec. 3 | 10:30 A.M. 116.390 1355 P.M. |16.770 | 2.3 gain | See Table XII f | FE Ae. [16.580 7|o21 7? 2 6 | 9:55 A.M. |10.640 12:14 P.M. /10.850 | 2.0 gain 3:02 10:630 | 2.0 loss 3 10:30 A.M. | 8.115 12:35 P.M. | 8.360 | 3.0 gain 3:10 8.115 | 2.9 loss a gain of 2 to 3 percent. The specimens most shrunken with drouth were found to absorb water most rapidly. The rapid loss of water and the curiously close correspondence in each case between the percentages of gain and loss, suggest that in this species it is merely the tubercles that act as organs of absorp- tion, and notwithstanding the fact that the water absorbed is so promptly given off in a dry atmosphere, it appears probable that in a period of frequent light rains the continued absorption of water by the tubercles is precisely the necessary preparation for the develop- ment of the young shoot which presently follows. Meantime the different deportment of these two species of Opuntia as regards amounts of water absorbed, corresponding as it does with a marked difference in size of their tubercles, suggests the desirability of a more extended comparison of these structures in different cacti with reference to their marcas for absorption and the physiological value of the process. GROUP V. A discussion of the annuals and partly herbaceous perennials that have been referred to a fifth biological group, many of which, structurally at least, are not to be thought of as characteristic desert plants, does not fall within the limits of the present study. As already stated, as many of them as have been studied agree in promptly absorbing water when it is presented to their leaves and internodes, which, however, is given off so rapidly in dry air that it hardly seems possible that its absorption is of any utility. Cy. Table I, Encelia and Sphaeralcea. 282 BOTANICAL GAZETTE [APRIL SUMMARY. From the preceding observations and experiments, in which woody plants were chiefly employed, it has been seen that certain species of desert plants of southern Arizona absorb water presented to their leaves and internodes, while others do not. The species represented in the vicinity of the Desert Botanical Laboratory may be divided into several biological groups, based primarily on the water relation, of which leaf absorption is a phase. Thus, in the first group, including shrubs, which retain well marked mesophytic tendencies, leaf absorption is characteristic. Members of the second group, more distinctively xerophytic in various structural particulars, are incapable of leaf absorption during their period of normal activity. The third group, decidedly xerophytic, but including species of widely different structure and habits, exhibits corresponding differences in regard to subaerial absorption, which takes place in some of its representatives and not in others. The fourth group, including cacti which are assumed to represent the extreme type of xerophytes, also exhibits interesting differences in size and structure of the tubercles by means of which water is absorbed. Finally, members of a provisional fifth group, which in habit and structure are nearer than any other others to the meso- phytes of moist temperate regions, absorb water largely, but very quickly give it up again. It may be doubted, perhaps, whether this classification, based on biological relations, has in itself any permanent value, but mean- time it serves to express and emphasize what is apparently no mere theoretical conception, but a simple historical fact, namely, that differences of habit on the part of these desert plants, as well as the structural adaptations with which they are correlated, have become established step by step together, during the long period of geographical changes through which the land they now occupy has been passing. A discussion of the physiological significance of the facts which have been brought out does not fall within the province of this pay er. DESERT BOTANICAL LABORATORY, son, Arizona. NEW SPECIES OF CALIFORNIAN PLANTS. ALICE EASTWOOD. (WITH TWO FIGURES) ’ Zygadenus exaltatus, n. sp.—Bulb large, pear-shaped, covered with brownish, membranous coats, about 1o°™ long, and half as wide: radical leaves forming a conspicuous bunch sheathing the base of stem, 6°" or more long, 2°™ wide, veins prominent, midrib conspicuous on lower part of leaf, less so above, glabrous except for the short, rather thick cilia on the margin: stem tall and stout, hollow, 7-8 high, 1°™ in diameter at base, leafy to the inflorescence: upper leaves merging into the bracts, lower cau- line with broad clasping base, 3-44" long and as broad as the radical leaves: inflorescence paniculate, 2-3°" long, the upwardly spreading branches varying in length at different stages of devel- opment, the lower branches generally bearing only staminate flowers, the perfect flowers principally borne on the main stem above the branches; staminate racemes with peduncles shorter than the bracts; bracts attenuate; bractlets white, membranous, longer or shorter than the slender pedicels, ovate-attenuate: perianth 17™™ in diameter, outer divisions sessile, elliptical, obtuse, the gland 2™™ from the base with parallel veins below forming a margined claw, veins on the upper part proceeding from the teeth of the gland, branching; inner divisions of the perianth with claw 2™™ long, 1™™ wide, the gland oblong, obtuse, veins asin the outer division; filaments broaden- ing towards the base, 5™™ long, attached to base of perianth, anthers oblong, becoming explanate in age: fruit becoming 2°™ long, includ- ing the rostrate styles, tapering a little more at base than at summit. Type collected by F. E. Blaisdell at Prindle’s ranch, above Mokelumne Hill, Calaveras County, California, at an altitude of 425™, in April 1900 The other specimens in the Herbarium of the California Academy of Sciences are Amador, California, May 1886, M. K. Curran ae with root); Soda Creek, Tulare County, C. A. Purpus, June 1896, no. 1549; Hindeman’s Trai] gis Coyote Pass, Tulare County, California, July 19, 1903, collected by myself. I also saw it on the trail on the way to Little Kern. In habit of growth and 283] {Botanical Gazette, vol. 41 284 BOTANICAL GAZETTE [APRIL size of bulb it is related to Z. paniculatus, but it has flowers much larger and all the floral organs differently shaped. It is probably the largest species of Zyga- denus known. It is probable that there is no Z. paniculatus on the western slope of the Sierra Nevada. “ Silene deflexa, n. sp.—Stems several from a creeping root- stock, about 24" high, erect, glandular-puberulent especially above: radical leaves spatulate, 1-2°" long including the margined petioles, rather thick in texture, obtuse; cauline leaves 3-4 pairs, the uppermost very small, not more than 3™™, the lowest oblan- ceolate to oblong, obtuse, 2°" long, connate-clasping and nodose at base: flowers solitary in the lower leaf-axils, pedicels erect and close to stem, flowers curved-deflexed in anthesis, erect in fruit; terminal flowers few, cymose, pedicels capillary, 7-12™™ long, generally shorter than the flowers; calyx narrowly cylindrical in flower, enlarging and breaking apart with the expanding capsule, ro-ribbed, 9™™ long, divisions rounded at apex, oblong, some- times uneven, membranously margined, 1.5™™ long; petals with the claws united tc the stamens and the stipe of the ovary at base, woolly, gradually enlarging to the blade, not auricled, blade 4-cleft into linear lobes, the two middle 2™™ long, o.5™™ wide, the two lateral narrower and shorter, appendages oblong, reaching the base of divisions, retuse with one side pointed; stamens and styles appar- ently not surpassing the petals, the latter three and the upper thick part of the ovary splitting into three valves, lower part of ovary thin cylindrical, all together 4™™ long; stipe 1™™ long and almost as thick: seeds suborbicular, brown, strophiolate. Type collected in the Hudsonian Zone above the nies Canyon Creek, Trinity County, California, by Vernon Baily, August 25, It is related to S. Lemmoni, but differs in the foliage, eS shorter filaments and styles, the close inflorescence, and the differently shaped floral organs. It really resembles that species only in having the flowers pendent and the blades of the corolla with four divisions. ’ Silene lacustris, n. sp.—Cespitose from+ creeping rootstocks, slender, erect, 1-1.5 high, glandular-puberulent throughout especially the inflorescence, flowering from the lowest axils: radical leaves narrowly oblanceolate, acute or obtuse, tapering to a long margined petiole, all together 2°™ long; cauline leaves 2-3 pairs, linear, obtuse, connate-clasping at base, 1-2°™ long, 1-2™™ wide: ing, ee ee 1906] EASTWOOD—CALIFORNIAN PLANTS 285 calyx broadly cylindrical, 1°™ long, thin, with ten purple nerves, the divisions rounded, 2.5™™ long, 2™™ wide; petals with claws woolly and cohering round the thick stipe, gradually broadening to the membranous laciniate auricles, 4™™ at top, blades purple, 2-cleft with rounded divisions, the lateral teeth short or none, the appen- dages bifid and laciniate, 1™™ long; stamens and styles exserted; ovary oblong, 4™™ long: fruit unknown. Type collected by the author at Monarch Lake near Mineral King, Tulare County, California, July 21, 1903. This species belongs to the group of high mountain Silenes, including S. Grayi, S. Watsoni, and S. Suksdorfii. In appearance and shape of leaves it most closely resembles the first, but differs from this and the others in the broader auricles, and the bifid, laciniate appendages of the corolla. ‘ Silene pacifica, n. sp.—Perennial, with thick woody rootstocks; stems generally several, erect, 4.5°" high, viscid throughout, espe- cially the inflorescence, nodes prominent: leaves rather thin, radical and lower cauline oblanceolate to obovate or elliptical, tapering at base and decurrent on the long petiole; blade 5-6°™ long, 2-3°™ wide, slightly ciliate, sparingly pubescent, obtuse or acute; peti- oles margined, dilated and sheathing at base; cauline leaves con- nate-clasping at base, 4-5°™ long, the uppermost leaves lanceolate, sessile: flowers verticillate in the axils or cymose on short peduncles, pedicels 0. 5-2°™ long, the longest equaling the longest floral leaves: calyx truncate at base, tubular, becoming somewhat turbinate with the enlarging capsule, very viscid, prominently green or pur- plish veined, the divisions deltoid, obtuse or acute, 4™™ long, green with white or purplish membranous margins, entire calyx 1.5°™ long; corolla claret color, the claws of the petals white or tinged with claret, 1.5°™ long, attenuate at base and broadening at trun- cate summit to 4™™, exserted 5™™, blade deeply cleft, each part entire, laciniate, or bilobed, the prominent divaricate teeth on each side almost as long as the divisions, more than 1™™, appendages laciniate-dentate, 1. 5™™ long; stamens monadelphous at base, encircling the stipe and pubescent, glabrous above, varying in length: pod 11™™ long, the tips of the three valves stellately reflexed and often splitting into five or six over the calyx when the seeds are ripe; stipe stout: seeds light brown, slightly pitted, reniform, 2™™ wide. 286 BOTANICAL GAZETTE [APRIL Type collected by the author along the south side of Rodeo Lagoon, not far from the ocean, Marin County, California, July 4, 1905. The species seems to be isolated, as the number of plants is small and it is not elsewhere to be found in the region. It is also in danger of extermination on account of the improve- ments that are now going on in the vicinity of the military post This is most closely related to S. grandis Eastwood Sin Bodega Head, likewise a maritime species. It differs in having claret-colored flowers, a differ- ently shaped calyx, the simpler inflorescence, thinner and different leaves, and entire lack of the velvety pubescence so noticeable on S. grandis. A smaller and more slender plant. “Horkelia mollis, n. sp—Stems several, ascending from the sheathed caudex, red-purple, villous with fine silky spreading hairs, about 24" in height: radical leaves 6-9°™ long, less than 1°™ wide, the petiole less than half the entire length, often with a few scattered simple leaflets near the base; leaflets crowded towards the top, pinnately divided but apparently pedate on account of the lower divisions surpassing the upper, the divisions lincar-spatu- late, 3-4™™ long, finely villous; stipules adnate for 8™™, the free tips filiform-attenuate, about 4™™ long, villous; cauline leaves similar but with petioles becoming shorter as they ascend, stipules often incised and always broader than those on the radical leaves: flowers corymbose-capitate, terminating the stems, a few solitary ones or few-flowered clusters in the axils of the upper cauline leaves; hypanthium campanulate, 5™™ long, the bractlets linear, about as long as the subulate sepals; petals yellow, the blades broadly spatulate, 1™™ wide, a little longer, slightly shorter than the linear claw; stamens 15 in three rows; ovaries 5-20, glabrous, the slender styles tuberculate at base. The type is 4405 of Carl F. Baker’s distribution, collected by Culbertson July 19, 1904, at Hockett’s Meadows, Tulare County, California. In the her- barium of the California Academy of Sciences are specimens of the same, col- lected by the author along Volcano Creek in the same region, July 17, 1903. This species probably is most closely related to Horkelia campestris (Jones) Rydberg. A comparison with a duplicate of the type of the latter shows H. mollis to be a larger, more villous plant, the appendages of the hypanthium longer, the divisions more pointed, the petals more exserted and with blades orbicular and claws more pronounced. In general the flowers are larger. v STYRAX CALIFORNICA fulvescens, n. var.—Shrub a meter or so high, with stiff divaricate branches; older stems gray-black, i i ; : 1906] EA STWOOD—CALIFORNIAN PLANTS 287 younger white or tawny with dense stellate tomentum: leaves orbic- ular-cordate, the apex obtuse or abruptly acuminate, generally slightly longer than broad, 3-6°™, both surfaces stellate-tomen- tose, the upper less than the lower, the fulvous hairs often outlining the veins on lower surface; petioles 5~1o™™ long: flowers 1-3, cymose, pendent, the pedicels as long as the peduncles; calyx cam- panulate, cuneate at base, the margin truncate but marked with 5-6 short obtuse scattered teeth, densely clothed with white or rufous tomentum; stamens 12, almost equaling the petals, attached almost the entire length of the corolla tube, filaments glabrous, ribbonlike, anthers with cell divisions white, the connective yellow, thick; style thick, broadening at base, lower half tomentose, stigma 2-lobed, surpassing the corolla. The type of this variety was collected by the author May 17, 1904, near the Painted Cave Ranch in the Santa Inez Mountains back of Santa Barbara, California. Mr. T. S. Brandegee collected the same in the same mountains probably near San Marcos Pass in 1888. There is a specimen also of what seems the same collected by J. G. Lemmon near San Bernardino, May 1878. Near the head of Mission Creek a second collection was made by the author. This bush grew in the shade and was taller and less rufous than the others on the open hills. This differs from the typical S. californica in the broader, rounder leaves, heart-shaped at base, the much denser stellate tomentum, and the general prev- alence of rufous ha.rs especially on the calyx. v Diplacus calycinus, n.sp.—Suffrutescent, viscid-arachnoid through- out, the young stems light brown, branching diffusely: leaves elliptical to oblong, narrowed at each end, apex obtuse, base cuneate, margin revolute, entire or somewhat sinuate-denticulate, upper surface glab- rous, often viscid, lower tomentose and viscid, 2-6°™ long, 1-2°™ wide; petioles very short, revolutely margined, woolly at junction: flowers axillary, the peduncles 5~7™™ long; lower part of fruiting calyx cylin- drical, 2°™ long, 5™™ in diameter, 5-ribbed, upper half dilating ab- ruptly to thrice the diameter of the lower, with 5 strongly keeled almost equal divisions 7”™ long, 3™" wide at base when folded, 1™™ at the rounded apex, total length of calyx 3.5°™; corolla light yellow, the tube curved, uniformly slender for 1.5°™, dilating above, the divisions having a spread of 1.5-2°™, exserted from the calyx. 288 BOTANICAL GAZETTE [APRIL This was first collected by Mr. T. S. Brandegee in Kaweah Canyon, Tulare County, California, July 26, 1892. The type is 4407 of C. K. Baker’s distri- bution collected by Culbertson in the south fork of Kaweah River, 1800™ alti- tude, July 22, 1904. This species is distinguished from allied species by the peculiar foliaceous calyx described above. The corolla in the dried specimens cannot be satis- factorily described, as in both collections the specimens are a little old. “ Orthocarpus Copelandi, n. sp.—Stems about 1% high, simple or divaricately branched, minutely scabrous with short, curved hairs: lowest leaves narrowly linear-lanceolate, obtuse, 3-4°™ long; upper on main stem as long but twice as broad; uppermost on branches falcate, alternate or opposite: spike short and dense; lowest bracts green, the middle division like the broadest leaves, the lateral divisions spreading and very slender, about one-third as long as the middle; upper bracts shorter and broader, ellip- tical, rose-tipped: calyx thin and membranous, becoming globular- inflated, pink with green ribs, cleft half in front, deeper in the back, villous. with short gland-tipped hairs, 7™™ long, 4™™ broad, with divisions triangular attenuate; corolla minutely glandular, 13™™ long, galea straight, obtuse, rose-color, ciliate, 6™™ long, lower lip yellow, the three sacs inflated somewhat, 5™™ long, middle tooth much larger than the other two: capsule bright brown, 5™™ long, 3.5™™ wide, obovate with obcordate apex, with few (appar- ently only two) seeds.—Fic. 1. Collected on Mount Eddy, August 18, 1903 at an altitude of 2130" by Dr. Edwin Bingham Copeland, in whose honor it is named. It is a beautiful species related to O. imbricatus and that group which contains so many closely related species. H.E. Brown’s number 449 from the north side of Mt, Shasta is the same but very immature. v Veronica Copelandi, n. sp.—Perennial from slender, running root- stocks, about 1% high, simple, glandular-villous throughout: leaves five or six pairs, crowded on the lower part, sessile, oblong-ellip- tical, entire, acute, veinless, 1-1.5°™ long, 4-8™™ wide: racemes sometimes becoming 8°™ long, 5-15-flowered, the highest leaves often with one or two axillary flowers; bracts lanceolate, the lowest opposite, others alternate, shorter than the pedicels; peduncles 1-2°™ long, sometimes scarcely apparent; pedicels filiform, 5™™ long, a small bractlet immediately below the calyx appearing like another PPT — 1906] EASTWOOD—CALIFORNIAN PLANTS Fic. 1.—Orthocar pus Copelandi Eastw. 289 290 BOTANICAL GAZETTE [APRIL sepal: sepals 4, oblong-ovate, obtuse, 3™™ long; corolla purple, glabrous, g™™ across, the three larger divisions orbicular, entire, 4™™ in diameter, the smallest ovate-obtuse, 3™™ wide; stamens exserted, 4™™ long, filiform, anthers obtuse and obtusely sagit- tate at base, 1.5™™ long; stig- ma exserted from the opening bud, obscurely bilobed, style 7mm long, filiform at base, flattening and slightly broad- ening towards the apex: cap- sules becoming almost twice as long as the calyx divi- sions, broadly oblong, 5™™ long, 3.5™™ wide, emarginate, the lobes and sinus obtuse; style persistent.—FIGc. 2. This was collected on Mount Eddy at an elevation of 2500™ by Dr. Edwin Bingham Copeland, August 18, 1903, distribution of C. E. Baker, 1903. no. 3931. It is near to V. Cusickii Gray, differing in pubescence, shape of leaves and sepals, and a larger and more open- spreading corolla. v Erigeron decumbens, n. sp. —Stems several, from slender creeping rootstocks, decumb- ent or ascending, 1-1.5™ high, scabrous and somewhat canescent with short appressed hairs which are glandular at base (under a lens): leaves oblanceolate to spatulate, sessile, obtuse, apparently veinless, 5-15™™ long, 3-5™™ wide, with pubescence similar to the stems: heads few, rayless, 7™™ high, terminating short branchlets, which are leafy near the junction with the stem and have a few scattered minute bracts on the upper part; scales of the involucre in four series, glandular-puberulent, outer ones small, reflexed-spreading, inner green-tipped, ribbed, membranous at base, FIG. 2.—Veronica Copelandi Eastw. | | | 1906] EASTWOOD—CALIFORNIAN PLANTS 291 linear-lanceolate, acute, 6™™ long: corolla yellow, tubular, abruptly narrowed 1™™ above the base, the border consisting of five short, obtuse, incurved teeth; style branches exserted, the hairy tips very short: akenes slightly hairy at top; pappus simple, barbellulate, as long as the corolla. Collected by Dr. Edwin Bingham Copeland on Mount Eddy, Siskiyou County, California, at an altitude of 1400™, August 17, 1903. It belongs to the group which includes E. miser Gray, as well as many species described by Dr. E. L. Greene in Flora Franciscana, p. 394; but it agrees with none. Erigeron Copelandi, n. sp.—Cespitose from an underground, branched caudex, covered with black, scale-like, imbricated bases of old petioles: radical leaves spatulate, subcanescent, with closely appressed very short pubescence; petioles equaling or longer than the blades, together 1-3°™ long, 4-8™™ wide, the petioles dilated and closely imbricated at the reddish-purple base: stems 1-flowered, 5-10" high, sparsely leaved with narrow linear or linear-oblan- ceolate leaves 5-10™™ long, becoming minute and bract-like on the glandular-puberulent upper part which is like a peduncle: heads about 6™™ high exclusive of the numerous, very narrow, lilac to violet rays, which are 5™™ long; scales of the involucre in three series, glandular-puberulent, the outermost shorter, clothed with some scattered hairs, innermost linear-attenuate, sparsely ciliate, green-ribbed, membranously margined, about 5™™ long; disk flowers numerous, yellow, 2.5™™ long, narrowed 1™™ above the base, glandular on the lower part, the border of five short acute incurved teeth: pappus upwardly barbellulate, simple, that of the ray flowers shorter than that of the disk, none as long as the corolla; akenes clothed with upwardly spreading hairs; stamens exserted in some flowers, pistils in others; fertile and sterile flowers in the same head, ray-flowers sterile. Collected on Mt. Eddy, Siskiyou County, California, at an altitude of 1250™ _ by Dr. Edwin Bingham Copeland, in whose honor it is a pleasure to name this pretty plant. It is related to E. pygmaeus Greene and others of that group, but differs from all in caudex, pubescence, leaves, and heads. Y Chrysopsis gracilis, n. sp—Stems slender, simple, 3°" high, loosely and sparingly villous-arachnoid, terminated by 2-4 cymose heads: leaves thin, linear-lanceolate, narrowly acuminate, 3-4°™ 292 BOTANICAL GAZETTE [APRIL long, 6™™ wide, sessile, the upper surface somewhat dotted, lower surface arachnoid (under a lens): peduncles with pubescence like the stem but also somewhat viscid, bracts few, narrowly linear: involucral scales in about 5 ranks, the outer narrowly linear- atten- uate, the others lanceolate, acute, tipped with a green and glandular spot, below yellowish, chartaceous, keeled, the innermost some- times tinged with purple and considerably surpassing the others; heads with about 15 flowers, rayless: corolla straw color, about as long as the pappus, trumpet-shaped, gradually narrowed to the base, border with acute teeth, 1™™ long; style branches filiform, exserted, twining around each other at base; pappus thick, with an outer shorter row, barbellulate; akenes flat, villous, white. Collected on Mount Eddy at an elevation of 2225" by Dr. Edwin Bingham Copeland, August 17, 1903. This comes very near C. Breweri Gray, of which it may prove to be only a variety. It differs however, in the simple instead of much branched stems, more finely arachnoid pubescence, and leaves of different outline. The invol- ucral scales are the most distinctive; in C. Breweri they are attenuate and not keeled; in C. gracilis they are broader, acute, keeled, conspicuously green- tipped and glandular; the corollas are paler and the pappus not so rough. - Psilocarphus tenuis, n. sp.—Sparingly clothed with long loose white woolly hairs; stems filiform, erect or ascending, 3-5°™ high, with few slender divaricate branches: leaves oblong to elliptical, 5-10™™ long, 3™™ wide, veiny and submembranous, mucronate at apex, the base of the opposite leaves connate-clasping: heads in the forks and at the ends of the branches, the involucral leaves 4, ovate-oblong, folding over and almost concealing the flowers within, texture similar to the other leaves: fertile flowers few or many, completely enclosed by the obliquely-cuneate bracts, these gibbous, veiny, membranous, slightly woolly, 2.5™™ long, the apex orbicularly truncate or concave, the exserted membranous tips brown- ish, conspicuous, generally curved upwards: akenes shortly stipitate, narrowly obovate, 1™™ long: sterile flowers few, the corolla attenuate to the base, divisions reddish-brown. Type collected at Monterey, California, by Mrs. Joseph Clemens, July 1905. What seems to be the same, but too young for certainty, was collected by the author at Bakersfield, Kern County, California, April 4, 1893, and at Kaweah, Tulare County, California, April 27, 1895. 1900] EASTWOOD—CALIFORNIAN PLANTS 293 This seems most distinct from all the other species in having the involucral leaves almost closing over the flowers, the peculiar concave or truncate top to the bracts enclosing the fertile flowers, and in the more veiny and membranous foliage, less woolly pubescence, and more slender habit. ~ Senecio Millikeni, n. sp.—Stems tall, glabrous, hollow, ribbed, paniculately branched, the slender virgate branches leafless in the lower part: leaves linear-lanceolate, narrowed at both ends, with acute apex, sessile base, margin dentate with small uneven obtuse teeth, the lower 12°™ long, 2.5°™ wide, diminishing up- wards: panicle thrysiform, the peduncles and pedicels slender, bracts and bractlets attenuate, equaling or longer than the slender pedicels: heads 1°™ high, bracteate at base, the involucre 5™™ high, with glabrous scales tipped at apex with a tuft of tomentum; rays 6, 3-toothed, 7™™ long, style exserted 3™™; disk flowers 7™™ long, the acute triangular teeth of the corolla slightly gran- ular, stamens exserted but style branches surpassing them: akenes glabrous; pappus soft and abundant, about as long as the corolla. Type collected in Natural Bridge Meadows, Tulare County, California, by Culbertson, Aug. 10, 1904, C. F. Baker’s distribution 4268. It is named in honor of Mr. Culbertson’s assistant. This belongs to the polymorphous group of which S. triangularis was the first described. It differs from all in the narrowed bases of the leaves, the thyr- siform inflorescence, and the smaller heads. SAN FRANCISCO, CALIFORNIA. Pricrerk ARTICLES. NOTES ON NORTH AMERICAN GRASSES. VI. SYNOPSIS OF TRIPSACUM. Tripesacum L., Syst., Ed. 10, 2:1261. 1759. A GENUS of grasses confined for the most part to North America. The type species is 7. dactyloides L. KEY TO SPECIES. Staminate spikelets all sessile or nearly so, outer glume coriaceous; spikes single or 2- to 3-digitate. Section DACTYLOIDEs. Blades 4 to 5°™ wide, pubescent on upper surface. . . Jatifolium Blades mostly less than 2°™ wide Blades 1 to 3™™ wide, involute . . - . + « + floridanum Blades 1 to 2°™ wide, flat Sheaths glabrous, blades sian except sometimes along the midribabove. . . . . dactyloides Sheaths more or less hispid, or sometimes es glab- rous, blades hispid on upper surface dactyloides hispidum Staminate spikelets with one of the pair sessile, the other pedicelled, outer glume membranaceous; pistillate spikes branched, form- ing a fascicle. Section FascrcunaTa. Sheaths hispid . . pilosum Sheaths glabrous daceps ‘oe fowccstiosé: or « Miepid ‘ally at the throat Blades 3°™ or more in width, glabrous . . . fasciculatum Blades 2°™ or less in width, pubescent on upper surface 1.5 to 2°™ wide, flat or folded, culms robust . .lanceolatum 5 to ro™™ wide, more or less involute, culms Ct, . . « . Lemmoni Tripsacum latifolium, n. sp. ge ose vaginis glabris vel apice pubescentibus, laminis amplis, ad 4.5°™ latis, 70°" longis, planis, supra pubescentibus subtus scabris vel glabrescentibus, spiculis steril- ibus geminis sessilibus, 3-4™™ longis, oblongis, obtusis vel breviter acutis. Culm robust, 1°™ in diameter, glabrous; sheaths glabrous or pubes- cent towards apex; blades ample, as much as 7o°™ long and 4.5°™ Botanical Gazette, vol. 41] . [294 de ee 1906] BRIEFER ARTICLES 295 wide, pubescent above, minutely papillate-scabrous or glabrescent be- neath, scabrous-ciliate on the margin; ligule very short, scarcely 3™™ long, fimbriate; spikes 1 to 3, similar to 7. dactyloides but more slender, pistillate section 2 to 3™™ wide, staminate spikelets sessile or nearly so, 3 to 4™™ long, outer glume coriaceous, oblong, rounded at apex, scab- rous, ciliate on marginal keels, rather minutely striate with about ten nerves. ; The type specimen was collected by H. von Tuerckheim at Cubilquitz, Dept. Alta Verapaz, Guatemala, alt. 350", Jan. 1902, no. 8333. The only other specimen I have seen was collected by C. Thieme at San Pedro Sula, Dept. Santa Barbara, Honduras, alt. 500™, March 1887, no. 55958. Both specimens are in the National Herbarium (Herb. John Donnell Smith). The species is well distinguished from the other species with sessile staminate spikelets by its broad pubescent leaves. Tripsacum DActTyLompEs (L.) L., Syst., Ed. 10, 2:1261. 1759.— Coix dactyloides L., Sp. Pl. 2:972. 1753.—Usually glabrous through- out except the upper surface of the blades along the midrib near the base. This and sometimes a considerable portion of the upper surface of the blades may be sparsely pilose. The specimens from Florida and along the Gulf Coast are usually pilose in this way, or occasionally the pubes cence may extend to the young sheaths of the branches. The more pubes- cent forms connect the species with the following subspecies, which occurs in Mexico. The terminal spikes are usually in digitate clusters of two to three, while the axillary spikes may be single. Sometimes, especially in Texas, the terminal spikes are also single (TJ. dactyloides monostachyum) (Willd.) Gray, Man. 616. 1848. T. monostachyum Willd., Sp. Pl. 4:202. 1805. Type locality ‘Carolina meridionali.”” Southern New England to Florida and Texas, mostly near the coast; but extending inland west to west Texas, and north to Nebraska, Iowa, southern Illinois, and eastern Tennessee. If the spike is single the pistillate portion is cylindrical; if the spikes are two or three, the pistillate portions are flattened on the inner surfaces so that all together they form a cylinder, and the lower are more or less peduncled. TRIPSACUM DACTYLOIDES hispidum, n. subsp. —Laminae supra his- pidae; vaginae hispidae vel glabrescentes. The staminate flowers are less chartaceous than is usual in 7. dacty- loides. Mexico and southward. San Luis Potosi, rocky hills, Las Canoas, 296 BOTANICAL GAZETTE [APRIL Pringle 3811 (type); Jalisco, Rio Blanco, Palmer 509; City of Mexico, Holway 8; Lower California, El Taste, Brandegee, Nov. 1, 1902; Trini- dad, Botanical Garden Herbarium 3303; Central Paraguay, Morong 675. This form connects T. dactyloides with T. lanceolatum. In some specimens the upper spikelet of the staminate pair is somewhat pedicelled. T. dactyloides and possibly some of the other species may occur widely distributed in South America. Information on this point is desired. TRIPSACUM FLORIDANUM Porter, Contr. Nat. Herb. 3:6. 1892. PortTEr’s herbarium name was published by Dr. VAsry in his monograph of the grasses of North America. Type locality ‘Florida (A. P. Garber) and Texas (G. C. Nealley);’ duplicate type in National Herbarium. T. dactyloides floridanum Beal, Grasses 2:19. 1896. There are no _ specimens of this species from Texas in the National Herbarium, nor are there any so labeled by Dr. VAsEy; consequently the Texas locality given above is uncertain and is probably incorrect. Our specimens are all from the vicinity of Miami, Florida, Garber 454, June 1877 (type); Pollard & Collins 272, April 1898; Eaton 530, Dec. 1903; Hitchcock, March 1903. Distinguished from T. dactyloides by its smaller size and much nar- rower leaves. TRIPSACUM FASCICULATUM Trin.; Ascherson, Bot. Zeit. 35:521- 1877.—Well distinguished by its ample glabrous leaves, which are as much as 6.5°™ wide and 70% long, resembling leaves of Indian corn (Zea mays L.). Plant glabrous throughout; spikes branched, forming a fascicle; staminate portion slender and more or less flexuous, the spike- lets 5 to 6 ™™ long and broadest near the top. The name first appears in the second edition of SrEUDEL’s Nomen- clator. 2:712, as Tripsacum “‘fasciculatum Trin. Mpt. Mexico. T. dactyloides Schlecht. in Linnaea VI.”” The latter name is a nomen nudum, as is also T. fasciculatum Trin. in Steud. Gram. 1:363, and in Ruprecht, ‘Bull. Acad. Brux. 9:243. The first description appears to be by ASCHER- SON’ in 1877, Bot. Zeit. 35:525, where a specimen from ‘“‘Pr. Hacienda de la Laguna (Schiede)”’ is designated as the type. Fournter, Mex. Gram. 69. 1881, includes the name without description and cites the - following specimens: Hacienda de la Laguna (Schiede 947); Orizaba * ASCHERSON had previously mentioned the species and given a brief description as follows: ‘Diese Art besitzt Blatter von der Breite der Maisblitter, und die zahl- reichen, schlaffen, mannlichen Inflorescenzzweige, deren Aehrchen kleiner als bei T. dactyloides sind, erinnern ebenfalls an Euchlaena.” (Verh. bot. Ver. Pr. Brandenb. 17:79. 1875, in a footnote to an article on Euchlaena mexicana.) a cogs ie rc I ace 1906] BRIEFER ARTICLES 297 (Bourgeau 3138); Mirador (Liebmann 549); Zacuapan pr. Jalapa (Gal- eoltt 5796); Arumbaro (Galeotti 5844). The Bourceau and LirBMANN specimens are in the National Herbarium; also Brade 16174, from Costa Rica. _TRIPSACUM LANCEOLATUM Rupr.; Fournier, Mex. Gram. 68. 1881.— Leaves mostly 1 to 2°™ broad, pubescent on the upper surface; stami- nate flowers 7 to g™™ long, spindle-shaped, often rather abruptly nar- rowed above the middle. Mexico. Sonora, Guadaloupe Cafion, International Boundary Com- mission, 2035; Durango, Palmer 537; Oajaca, Villa alta, Liebmann 547; Lower California, Sierra de San Francisquito, Brandegee 6, Sept. 1899; Jalisco, between Huejuquilla and Mesquitec, Rose 3570. In addition to these specimens in the National Herbarium, FourNIER gives the following: Inter Victoria et Rio Blanco (Karwinsky); Borrego prope Orizaba (Botteri 1213 in herb. VAN HeEuRcK); Mirador (Schaffner); Tacubaya (Schaffner 41 in herb. FRANQUEVILLE); Secus Amnem in her- bosis- pr. Pedregal (Bourgeau 444); Aguas Calientes (Hartweg 252). Liebmann 547 is also cited by FourRNIER and it is upon this specimen that I have based my identification of the species. FoURNIER’s description does not apply in all respects to the plants which I have included under this species. He states that the culms are pilose, which is not true of any of the specimens I have seen. Neither are both staminate spikelets pedi- celled, as he describes. The name first appears in Plant. Haitw: Addenda, p. 347. In the body of the work (p. 28) no. 252 is listed without description as T. dacty- loides ‘‘in saxosis, Aguas Calientes.” In the addenda this is corrected as follows: ‘“‘n. 252 est species a Tripsaco dactyloide distincta, T. lan- ceolata, Ruppr. ex cl. Rupprecht in Litt.”” Fournter (/.c.) cites T. lanceolatum Rupr. in Benth. Pl. Hartw. 247. Under the circumstances I think Hartwec’s no. 252 from Aguas Calientes should be considered as the type of 7. lanceolatum rather than coca gomgse s specimen, the first cited by FouRNIER. FourNIER cites as a synonym of this “7. acutiflorum Rupr. mss. in herb. Petrop.”” Under the rules of the recent code 7. acutiflorum was not published. FouRNIER (/. c. 69) also mentions without description, var. 8B monostachyum from San Luis Potosi (Virta 1447). I have not seen this specimen. TRIPSACUM PILOSUM Scribn. & Merr., Div. Agrost. Bull. 24:6. rgor. —Type locality Mexico. ‘Collected on the road between Colotlan and 298 BOTANICAL GAZETTE [APRIL Bolafios, State of Jalisco, 2841 J. N. Rose, September 7, 1897.’’ Speci- men in National Herbarium. The preceding species, together with this and the following, form a rather closely connected series. The type of T. pilosum is distinguished by the strongly papillate-hirsute sheaths, and the blades pubescent upon both surfaces, but these characters are much less marked in some of the specimens which agree with the type in other particulars. I have referred here the following specimens: Jalisco, Rio Blanco, Palmer 508; Cafion near Guadalajara, Pringle 2623, and hills near Guada- lajara, Pringle 2611; San Luis Potosi, limestone ledges, Tinamel, Pringle 3993; and San Jose Pass, Pringle 3447. TriesacuM Lremmoni Vasey, Contr. Nat. Herb. 3:6. 1892. Type locality, ‘‘Huachuca Mountains, Arizona (J. G. Lemmon).” Type specimen in National Herbarium. T. dactyloides Lemmoni (Vasey) Beal Grasses 2:19. 1896. Plant glabrous throughout except the lowermost sheaths, which are more or less hispid. The leaves are long and narrow, 5 to 10™™ -wide, and in herbarium specimens inrolled at the margins. In addition to the type specimen I have included two Mexican speci- mens, Jaral, Gebirgsthaler, Schumann 1718, and Jalisco, Mountains near Guadalajara, Pringle 2610. These two specimens have the spikes digitate instead of fascicled as in Arizona specimen, but the latter has the lateral spikes in ones or twos.—A.S. HirrcHcock, U. S. Dept. Agric., Washington, D. C. BOOK REVIEWS. Vegetable foods. THis well-known work of MOELLER on this subject," first put out about twenty years ago, has played an important part in connection with the increasing use of the microscope as a practical instrument for recognizing vegetable sub stances in a more or less finely divided state. Many changes have taken place essary. now of the Connecticut eco ral Experiment Station, a pupil of MoELLER, has furnished very important aid in the form of excellent figures as well as text. The scope of the work is in general indicated by the title and those articles here treated are with few exceptions used as food for man or beast, the term food being defined so as to include such articles as flavoring agents as well as tea, coffee, and cacao. Under the appropriate headings those substances are also described and figured which occur as impurities, substitutions, and adulterations. Since it often happens that condiments are also official , many chapters have a strong pharmaceutical interest. A few articles are con- sidered which have their chief significance as coe products, e. g., sandal wood, guarana, cubebs, cola, salep, and ca In the treatment of the individual has. the book is distinguished by a concise and exact statement of the features, gross and microscopic, character- izing the structures concerned, dimensions frequently cited giving definiteness to terms of size. As valuable as the excellent text, are the numerous drawings illustrative of it. A large number are original, many being by Dr. Winton. A bibliog- raphy of the most important articles written on each subject closes the consid- ation. One novel feature among the illustrations is seen in the gross pictures of the leaves discussed. Here a direct print is made on a sensitive surface, using the leaf itself as an opaque object. This method has been successfully used before by a number of authors with various objects and here the result is in general successful. Frequently a very considerable amount of detail has : eect nee — ——T vege Nahrungs- und Genussmittel aus dem fi te und unter Mitwirkung A. L. Winton’s vermehrte Auflage. ous pp- iit sons jigs. 599. Berlin: Tolles” Springer. 1g05. M 18; geb. M 20. 299 300 BOTANICAL GAZETTE [APRIL oe is in the half tone reproductions of these so-called ‘‘autophoto- ‘The revision of this important work has again brought it to the front and promises to continue it as one ies the valuable literary aids to the investigator of pure foods.—Ropney H. ALMOsT simultaneously with the foregoing has appeared in this country a similar compendium by the same team; this time the pupil leads and the master is the collaborator.2 The general plan and purpose of Dr. WinTON’s weighty volume are similar to those of Dr. MoELLER’s. The fact that it is in English will give it a sale that the German book could not hope to attain among the food commissioners and inspectors and the official chemists, to whom at present such a work makes its chief appeal. By reason of the existing agitation in this country on the subject of pure foods and drugs, the enforcement of existing laws, and the imminence of new and more exacting legislation, this publication is peculiarly timely. The botanical features are on the whole reasonably accu- rate, especially the anatomy, which is most fundamental. The definitions in — the glossary are not always above criticism, and accuracy would not have ren- dered them less practical. The illustrations are numerous and good, particu- cited in the bibliography. The arrangement of material, analytic keys, lists of adulterants, and the suggestions as to diagnosis are sure to be of great practical Service in the new paaoraes against sophistication by unscrupulous manu- facturers and dealers—C. R. B. MINOR NOTICES. Cryptogamic flora of Brandenburg.s—This monumental work begins its seventh volume with the first fascicle of the Ascomycetes. Its character and scope are so well known that the announcement of its publication and contents Nn suffice to secure the orders of all who concern themselves with this group. miasci are treated by G. Liypau; Saccharomycetineae by P. LINDNER; Pobsatiein by G. Linpav; Exoascaceae, Erysiphaceae, Perisporiaceae, Macro- thyriaceae, and Aspersiaceac by F. Necrer; Onygenaceae, Elaphomycetaceae, Terfeziaceae, and Tuberaceae by P. Hennincs.—C. R. B. 2 Winton, A. L., The microscopy of vegetable foods, with special reference to the detection of adulteration and the diagnosis of mixtures. With the collaboration of Dr. JoseF MOELLER. Imp. 8vo. pp. xvit+7or. figs. 589. New York: John Wiley & Sons. 1906. $7.50. 3 Kryptogamenflora der Mark Brandenburg, apn 7, Heft x. Pilze. Von P. HENNINGS, G. Linpav, P. LinpNErR, F. NEGER. 8vo. pp. 160. figs. 17. pls. 8. Leip- zig: Gebriider Decanneace: 1905. M1.50. (Not sci hear) 1906] CURRENT LITERATURE 301 NOTES FOR STUDENTS. What is a species?—The many discussions as to what is a species have resulted in a general appreciation of the facts that species are not all of equal rank, that they are distinguished by more or less arbitrary characters, and that although many species are real natural groups of individuals, many others are simply arbitrary groups, associated for the sake of convenience. After review- ing the various methods of distinguishing species, KUpPFFER* concludes that no method will apply in all cases, that all methods are of importance, and that when the several methods are used conjointly, little difficulty is experienced. KupFFER then turns to the methods of K6LREUTER, based upon the sterility of hybrids, as a method which has not been used to the extent its merits warrant. Sterility of the hybrids being presumably due to defective germ-cells, he depends for his measure of sterility upon the condition of the pollen, basing his method upon the fact pointed out a few years ago by JENcIc5 that viable pollen swells immediately upon the introduction of water, while the sterile pollen remains - Shrunken, and that this capacity of the normal pollen to swell is retained for many years in herbarium materials (more than 50 years in Viola, fide KUPFFER). Although the author eee that considerable sterility of the pollen has been observed in many “good” species, he has himself never found a pure species in which more than a few (ein Paar) per cent. of the pollen grains remained shrunken, the implication being that the reported instances would bear further consideration. After examining a number of species and their hybrids, especially among the Violaceae, he concludes that when a supposed hybrid shows much less fertility of the pollen than its supposed parents, it is not a mecessary but a sufficient proof (1) that the supposed hybrid is truly a hybrid, and (2) that its parents belong to distinct species, Application of this aehed is then made with interesting results to forms of Potentilla, Viola, Thymus, etc., which have puzzled the systematist—GEORGE H. Suu. Propagation of grain rust—Further comments by Dr. JaKosp Ertxsson® on the question of the origin and distribution of the rust-diseases of plants have recently been presented to the botanical public through separata. The author has not essayed so much to put forth new facts, as to bring together and review those recently published, in so far as they bear upon his mycoplasm theory, giving especial attention to adverse criticisms. 4Kuprrer, K. R., Kélreuters Methode der Art Abgrenzung nebst Beispielen ihrer Anwendung und einigen allgemeinen Betrachtungen iiber legitime and hybride Pflanzenformen. Acta Hort. Bot. Univ. Imp. Jurjevensis 6:1-19. 1905. 5 JENcIc, Untersuchungen des Pollens hybrider Pflanzen. Oesterr. Bot. Zeits. $0:1, 41, 81. 1g00. 6 Errxsson, J. Zur Frage der Sulsictoaing und Verbreitung ie Rostkrank- heiten der Pflanzen. Arkiv for Botanik 53:1-54. 1905. 302 BOTANICAL GAZETTE [APRIL He maintains that after taking into consideration the studies and observa- . tions of MArsHALL Warp and PLlowricut in England; McALpine and Coss in Australia; Bottey, HircHcockx, and CaRrLeTon in North America; BaRr- cLay in India; KLEBAHN, DreTEL, SCHROETER, and MAGNus in Germany; LAGERHEIM in Sweden, and others, the wintering of the uredo-bearing mycelium, or of the uredospores, so as to be a source of infection for the coming season, has not been proven. The evidence, chiefly as brought forward by KLEBAHN, to show that the first appearance of the rust in spring can often be accounted for by uredospores being carried long distances by the wind, is reviewed, and the conclusion reached that this is an assumption based on no direct evidence and highly improbable. The author then enters upon the vital part of the subject and discusses the mycoplasm theory and its recent criticism, especially that which has been most ably presented by KLEBAHN and MarsHALL Warp. After an extended examina- tion of the works of these authors, he finds that his theory has not been affected. He directs attention to a report by BIFFEN of recent experiments in hybrid- izing wheat carried on at Cambridge, England, in which the appearance of rust on the plants can best be explained by assuming that the mycoplasm of certain varieties was transmitted through the pollen to the resulting hybrid. —J. C. ARTHUR. Gynodioecism.—CorrENS? “has presented a second® report on the gyno- dioecism of Satureia hortensis and Silene inflata, giving full confirmation of his earlier conclusion that the pistillate form produces only, or mostly, pistillate offspring when fertilized, as it must be, by the bisporangiate form. If the pis- tillate form is a mutant from the bisporangiate and differs from the latter by the possession of a distinct hereditary unit, as suggested by Burck,® all the seeds produced by a pistillate plant are of hybrid origin, and the observed facts would be best explained as a case of dominance of the newly risen character over the older. In Satureia this dominance (?) is complete, but in Silene the offspring of the pistillate plants were pistillate in only 87-93 per cent., the rest being bi- sporangiate. Although this behavior looks very much like Mendelian inheri- tance, a number of cases are cited in which quite contradictory results have been obtained, so that while the author states it as a law that each sex has a tendency to transmit its own sex form, he does not look upon this as dominance in the Mendelian sense—Grorcre H. SHULL An ear of corn.—The origin of such economic plants as wheat and maize, which have a wide distribution in cultivation but are unknown in the native tis Bot. GAZETTE 39:304. Ap. 19 , C., Weitere cae iiber die Gynodioecie. Ber. Deutsch. Bot. Gesell. aie 452-463. 1905. , W., Die Mutation als Ursache der Kleistogamie. Recueil Trav. bot. pee I-2:95 sqq. 1905. EDS lg _ ee 1906] CURRENT LITERATURE 303 state will doubtless always be an interesting subject for speculation. The most satisfactory hypothesis for the origin of maize, and that which has been until this time rather generally accepted, derives it from the teosinte (Euchlaena). It has been thought that the ear was formed by an abnormal coalescence of the pistillate spikes of that plant. The ease with which maize and teosinte may be crossed gives strong support to the theory that they are nearly related. An altogether different view of the origin of the pistillate spike of maize is presented by MonrGomERy’® and much evidence is given in its support. His ypotheses are that the ear of corn is the homologue of the central spike of the staminate inflorescence; and that the progenitor of maize was a much branched plant, bearing only terminal branched inflorescences of bisporan- giate flowers. The chief support of these hypotheses is derived from abnormal development of pistillate and bisporangiate flowers in the staminate inflores- cence, and vice versa. A number of photographs show these abnormalities and jig. 74 represents a plant, denuded of its leaves, showing that the same num- ber of internodes intervene between the central axis and the ear as are found between the ear and the tassel. Nothing in this new interpretation of the pistil- late spike of maize need lessen the conviction of its near relationship with Eu- chlaena.—GerorcE H. SHULL. The laws of inheritance —CorrENsS*' published a lecture on the laws of inheritance which presents in a very satisfactory manner the recent advances which have been made in this discipline. He would include in hybridization every union between two germ-cells having one or more different character- The laws of dominance and of the purity of the parental gametes are illus- trated from his own experiments on Urtica, Mirabilis, and Zea, and emphasis is given to the fact that these two laws are absolutely unrelated to each other, and that reference to them jointly as Mendel’s Law is ing. Latency is considered at some length, but the present state of knowledge of this subject leaves much to be desired. He makes a proper distinction between latency in the sense of invisibility, and frue latency in which there is actual inac- tivity of a unit that may be changed at times from a passive to an active state. Regarding the relation between MENDEL’s and GALTON’sS laws, he holds with DARBISHIRE,"? that both are correct and the antagonism only apparent, due to the different manipulation of the data. CorRens still maintains that sex is fundamentally unlike the unit-characters which behave in accord with MENDEL’s laws. Touching on xenia and tel- 10 Montcomery, E. G., What is an ear of corn? Popular Sci. Monthly 68: 55-62. jigs. - Jan. 1906. 11 CORRENS, C., Ueber basi et gn 8vo. pp. 43. figs. 4. Berlin: Gebr. Borntraeger. 1905. 2 DARBISHIRE, A. D., On the supposed antagonism of Mendelian to biometric ari of heredity. Mem. and Proc. Manchester Lit. and Philos. Soc. 49. no. 6. 1905- 19 pp- 304 BOTANICAL GAZETTE [APRIL ing he. holds that neither exists in the strict sense, namely that ids may escape m the ‘germ-cells to produce modification in the surrounding maternal tissues, or to be transferred thence into subsequent germ-cells.—GEORGE H. SHULL. Heterostyly in Primula——The inheritance of heterostylism in Primula has been investigated by BATESON and GreGory,' who find that there is general agreement with Mendelian expectation, the short style being dominant over the long style. A second character, a yellow flush in the center of the flower, which was found associated with an “‘equal-styled” condition, also proved to. be Mendelian and capable of being transferred by crossing to the short-styled form. The investigation showed that whenever the yellow flush occurs in a combination in which the long style would be expected, the styles do not develop beyond the level of the anthers, thus forming the “equal-styled” type. Several aberrant results were observed, the most noteworthy being a case in which a single plant indicated a different composition of its germ-cells, according as it was used as the pollen-parent or pistil-parent.—GrorGE H. SHULL. Asparagus rust —Smitu‘+ has published a final account of his investigation of the asparagus rust in California. One of the most important results of his work is the demonstration of the fact that the spores of this rust depend upon dew for the moisture required for germination. The more detailed account of the water relation of this rust was published in this journal.'s This discovery suggested certain practical methods of controlling the rust, such as planting the rows with the wind and preventing weeds and other plants or trees from forming a windbreak close about the asparagus field. In other words, the field should be well ventilated. The bulletin will long continue to be the standard work of reference for information upon the subject—E. Mrap WILcox. Potato scab.—HENDERSON* has recently published the results of his studies of the methods of control of the potato scab. He found that rolling the potato tubers in sulfur did not prevent the scab, and this is in accord with results secured by other investigators. Formalin and corrosive sublimate gave equally good results with the factor of safety in use in favor of the formalin. If treated pota- toes were planted in soil in which “scabby’” potatoes had grown the previous season, the scab appeared in spite of the treatment. This emphasizes the neces- sity of preventing new ground from becoming infected with the disease by plant- ing none but healthy tubers.—E. Mrap WItcox. "3 Bateson, E., and Grecory, R. P., On the inheritance of heterostylism in Primula. Proc. Roy. Soc. London B. 76:581-586. 1905. 14 SMITH, R. E.—Asparagus and asparagus rust in California. Bull. Calif. Exp. Stat. 165:1-99. figs. 1-46. 1905. 1s Smith, R. E., Bot. GazetrEe 38:19-43. figs. I-21. 1904. *6 HENDERSON, L. F., Potato scab. Bull. Idaho Exp. Stat. 52: 1-8. 1906. ee se ee | 1906] CURRENT LITERATURE cha 305 Nuclear division in Ascomycetes.—GUILLIERMOND"’ has continued his studies on nuclear division in the Ascomycetes, which support in all essentials the conclusions of HARPER and contravene those of MAIRE (except as to Gal- actinia), though they are perhaps not irreconcilable with them. However, his descriptions are not so detailed as those of HARPER in his last paper on Phyl- lactinia, especially as it relates to the centers of spindle formation. In this r GUILLIERMOND discusses chiefly the mother-cells of the asci and secre- tion. The species studied comprise Pustularia vesiculosa, Aleuria cerea, Peziza rutilans, P. Catinus, and Galactinia succosa. —B. M. Davis Soil waters.—CAMERON and BELL show" that as a rule the various mineral constituents of the soil solutions exist in sufficient concentration for the growth of crops, and that the magnitude of the concentrations is practically the same for all soils, because, generally speaking, soils contain all the common rock forming minerals, some of each species presenting its surfaces to the solvent action of the soil water; and on account of hydrolysis of the solutes this solvent action is continuous. The paper strongly supports the previous work of the Bureau of Soils which has been so much criticised, often on a@ priori grounds.—C. R. B. Non-infection by rusts.—Erysiphe graminis has a number of biologic forms which are confined to special hosts. Thus conidia from the form on wheat will not infect barley and that on oats will not infect wheat. SALMon’? has recently shown that the reason of the non-infection is not due to inability on the part of the conidia to germinate, but because the haustoria cannot establish relations with the cells of the host plant.—B. M. Davis. Endoparasitic adaptation —SALmoNn’? shows that Erysiphe haar adapts itself readily to an endophytic life. When spores are sown on oats or barley the mycelium ramifies in the intercellular spaces a haustoria are abundantly produced. Conidiophores develop profusely and perfect conidia where they arise on a free surface; and they even break through a weak barrier when they develop in intercellular spaces.—C. Greening of seeds.— Ernst?" finds that during the ripening of the fruit of Eriobotrya japonica the seeds become green, quite independent of light, by reason of the greening of the amyloplasts. The process begins at the plumule of the 17 GUILLIERMOND, A., Remarques sur la karyokinése des Ascomycttes. Ann. Mycol. witli pls. IO-12. 1905. 18 CAMERON, F. K., and Bett, J. M., The mineral constituents of soils. U. S. Dept. Agric., me Soils Bull 30. pp. 19 SALMON, E. S., On the stages = paige’ reached by certain biologic 5 forms of Erysiphe in cases of non-infection. New Phytol. 4:217. 1905. pl. 20 SALMON, E. S., On under cultural conditi tions. 21 Ernst, A., Das mesa: der pei yon Eriohbotrya japonica. a ce agrees ae by Erysiphe graminis DC. hae y. Soc. London B. 198:87~-97. pl. 6. 1905. Beihefte Bot. a Centralbl. r9™: 118-130. pl. 2. 1905. 306 : BOTANICAL GAZETTE [APRIL embryo and progresses from this region to the inner and outer faces of the coty- ledons. Complete greening, however, only follows illumination.—C. R. B The nucleus and secretion.—In the nectar glands on the stipules of the Vicia Faba, according to STOCKARD,?? the nucleus does not give out granular material directly to the cytoplasm, but it transmits a substance which results in the forma- tion of granules. Changes which occur in the cytoplasm during secretion seem to be controlled by the nucleus—CHARLES J. CHAMBERLAIN. Black rot of cabbage.—HarpDING, STEWART, and PrucHa’s find much of the cabbage seed in the market contaminated with Pseudomonas campestris, which may survive and become a source of infection to seedlings. They advise sterilizing seed by soaking for fifteen minutes in HgCl. 1:1000, or in formalin 1:240.— C.K, B. Movement of diatoms, etc.—Jackson suggests? that the evolution of oxygen is the true cause of movements of diatoms, desmids, oscillaria, nostoc, etc. He has been able to imitate the movements by those compressed tablets and bits of - aluminum of proper shapes which evolve gas—C. R. B Anatomy of Claytonia—A study of this genus by THEo. Hot forms one of the Memoirs of the National Academy,?5 where it may be overlooked by botanists. It contains some of the accumulating details which a master hand must some day correlate.— C. R. B. Apothecia of lichens —Gertr. P. Wotrr?® through some studies on the development of the apothecia in a number of lichens argues against LINDAU’s terebrator theory of the function of the trichogynes in lichens——B. M. Davis. Intercellular ducts.—The intercellular spaces in the cotyledons of Legumi- nosae function at the beginning of germination as conducting canals for aleurone which becomes dissolved and diffuses through them.?27—C. R. B. Mustiness.—The peculiar musty odor acquired by damp straw or corn is due, according to Roussev,?® to the oospora form of oe Dassonvillei and not to other of the fungus flora found thereon.—C. R. B 22STOCKARD, CHAS. R., The structure and cytological aegis aaa RY: secretion in the nectar ne of Vicia Faba. Science 21: 204-5. 19 23H ARDING, H. A., STEwart, F. C., PrucHa, M. J., Vitality of be cba black rot germ on cabbage seed. N. Y. Agr. Exp. Sta. Bull. 251: 177-19 05. 24JacKsoN, D. D., Movements of diatoms and ee microscopic perine Jour. Roy. Mic. Soc. 1905: 554-7. 2SHOLM, THEO., ee a morphological and anatomical study. Mem. Nat. Acad. Sci. 10: 27-37. pl. 7 1905. 26WoLFF, GERTR. P. aa zur Entwicklungsgeschichte der ha awe thecien. Flora 95:31. 27JOFFRIN, H., io BUNT des méats intercellulaires pees les cotylédons des Légumineuses au début de la germination. Rev. Gén. Bot. 17 : 421-2. 1905- 28BrocQ-RovssEv, Contributions a l’étude des causes qui provoquent l’odeur de moisi des grains et fourrages. Rev. Gén. Bot. 17: 417-420. 1905. nm Sincere crcanesrticicenns —— ~ f NEWS. PRoFEssor J. C. ARTHUR spent the greater part of January at the New York Botanical Garden in a study of Uredineae. Proressor B. M. Duccar has been spending the winter in research at the Botanical Institute at Montpellier, directed by Professor CH. FLAHAULT. THE Bulletin de l’Académie Internationale de Géographie Botanique an nounces the limitation of leading articles to thirty-two pages. We hope the movement will become general among journals. By proper condensation an author can say all he is entitled to say on one subject in such a space. Dr, Jesse M. GREENMAN spent some six weeks in Yucatan and adjacent Mexico collecting plants for the Field Natural History Museum, of whose her- barium he is assistant curator. He had a violent attack of malarial fever which interfered seriously with his work, but he has returned in good health and with fair collections. LAST SUMMER after lousy the Vienna Congress, Professor GrorGE F. ATKINSON spent some time in the vicinity of Nice, Paris, and especially in the Jura mountains in the vicinity of Pontarlier, studying the fleshy fungi. He collected over 300 species, made photographic studies, and preserved material for morphological investigation. THE VIENNA ConcREss nominated as presidents of the Committee of Organ- ization for the Brussels Congress of 1910 Lfo ERRERA and URAND. On account of the lamented death of Professor ERRERA the Association internation- ale des botanistes has named Senator Count Osw. DE KERCHOVE DE DEUTER GHEM you are ne any trouble with the finish r floors, or are not entirely pleased with eee appearance, it is certain you have not shown, and its strange and used LIQUID GRANITE, the finest floor finish absorbingly interesting ever introduced. It makes a finish so tough that, although the wood will dent under a blow, the finish will not its wonderful resources history recounted, in the Seaboard Magazine. crack or turn white. This is the highest achieve- ment yet attained in a Floor Finish, and is not SENT FREE ON REQUEST likely to be improved on Finished samples a woudl and instructive pamphlet on the care of natural wood floors sent free for the asking. BERRY BROTHERS, Limited, J. W. WHITE, General Industrial Agent Varnish Manufacturers, PORTSMOUTH, VIRGINIA NEW YORK PHILADELPHIA CHICAGO ST. 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MODERN EDUCATIONAL METHODS APPLIED TO RELIGIOUS TRAINING This characterizes the purpose and plan of the Constructive Bible Studies FOR PUPILS OF THE SECONDARY sobhongentit we recommend Studies in the Gospel According to Mark, by Ernest DeWitt Bur The auth has in view a two-fold purpose: first, to help the coil: through a study of the Gospel of Mark, to acquire a knowledge of the life of Jesus, 0 come into s ic acquaintance with him; and, secondly, to help him to form ee habit of coming to all the books of the Bible, with the question “What does it The book contains e igh remo notes, questions requiring written replies, and helpful ssdiaeier eal for oat and teacher FOR TEACHERS OF CHILDREN hap 8 to II years of age, the he ger entitled An /ntroduction to the Bible for Teachers e ag aa by Georgia L. Chamberlin, w prove especially helpful. 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Instructor in Botany in the University of Chicago HIS BOOK contains directions re collecting and preparing see material for microscopic investigation. It is based upon a course in ical micro- nd is i r kingdom from the Algae up to the flowering plant. the staining of karyokinetic figures, and formulas are given for the reagents more attention has been given to collecting material. New c chapters deal with the Venetian turpentine method, microchemical tests, free-hand sections, special methods, and the use of the microscope. These changes and additions have enlarged the volume from 168 to 272 pages 272 pp., ‘Bvo, cloth, net, $2.25; postpaid, $2.39. THE UNIVERSITY OF CHICAGO PRESS natin, ee G. H. STOELTING 60. ® 31-45 W. 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PORTER 326 Dearborn Street, CHICAGO Scientific Instruments ‘9061 pAvd SSa4PPV *g061 ‘x A[nf a10Jaq 10 UO JapUleUaI ay} puv ‘gob ‘1 ArenuLf d10Jaq 10 UO YIFY-aU0 ‘Lo61 ‘1 Ajnf ar0jaq 10 uo Yyyy-at0 ‘Lo61 ‘1 Arenuef s10jJaq 10 uO YIY-aU0 tgo61 ‘1 A[Nf{ 10Jaq IO UO JooI9Y} YIFY-IUC) : SMOT[OJ sv ‘sjuouiAed [enuUe-IUIaS aay UI CS) 0 acme aang JO uIns 94} OSvoIyD jo AytssaAluG ayy, 03 Avd 03 saaise pousisiapun oy} syoo[qo pies 0} uondiosqns A199 pu yore jo puv sasimaid oyi Jo tlonesopisuos ul ‘310ja1agD ‘move ‘spuey Auvul ul pasouoy st aueu srodivzy quapiserg wogm Aq 1aquinu a81v] ay) Jo ynq ‘| U0 MI} BV JO JOU ILS OY) 9q P[NOYS SuIpying [vuOUIIUI SIy} Jey} Potisop pur popudsjuUl st yI ‘anaiaq ge Quy ‘ooofoSz‘1g ynoqe 4soo 0} OSvdIY) Jo AzISIDAIUL) 94} Jo so[suvipenb ay} Ul Suipying Areiqiy & wy 07 [elIOMUT & sv yoaI0 07 asodoid ‘19};9R¥I~YD poqexXd sIy pUe YIOM [NJ -asn sy jo Aroma oy} Yystsyo 07 Surysim ‘s9divpy Aourey wei, Woptserg Jo spuatsy oy) ‘svIaag gz qunfe davagn ss jerr08wmayf LadAVG_E CIMIVVE wuiKEgE 34D ‘ODVOIHS ‘ODVO -IHD 40 ALISHZAINN J3HL ‘SAaLSNYL JO GHVOG FHL 4O AYVWLAYOAS FHL OL LIGQNAS AGNV 39031d SNIMO1104 3AHLNDSDIS GNV LNO T1114 OL GALSANOAY AV IVIYOWAW SIH OL ASIHYOSENS OLHSIM OHM YadYdVH LNAGISaYd 4JO SANFId4 a ie Sata ee —~ VOLUME XLI NUMBER 5 DOTANICA“E GAZETTE MAY, 1906 NEW AND NOTEWORTHY WESTERN PLANTS. III." A. D. E. ELMER. ’ Phacelia acanthominthoides, n. sp.—An annual or biennial, 2 to 5°" high or higher: stems many, profusely branched from the base, erect or reclining, cinereous: leaves at least in the mature plants all cauline, alternate, usually subtending the branches, those from near the base 1o™™ long, pinnately 3 to 5-lobed or toward the apex only pinnatifid; the pubescent petiole almost equaling the blade proper; lobes hispidly strigose on both sides, 1°™ long or less, margins with few much-rounded teeth; upper leaves finer dissected and without petioles: inflorescence ample, in widely branched scir- poid racemes; flowers bluish, upon short pubescent pedicels: the 5 sepals pubescent, 4™™ long, 1.5™™ wide, linear-oblong, very obtuse, united at base, much exceeded by the flower: corolla 8™™ long; petals 5, very short, obovate, lateral nerves extending from the middle basal primary ones: stamens 5, exserted, inserted near the corolla base and alternating with the segments; anthers elliptic, 1™™ long, versatile; filaments glabrous, slender, 8 to 10™™ long, with minute hyaline appendages at base: ovary ellipsoid, pubescent; style per- sistent, 7™™ long, cleft nearly to the base, the united portion sparsely pubescent; stigmas minute, terminal: herbaceous sepals of the mature capsule 8™™ long, 4™™ across the widest part, ovate, acute, sub- coriaceous, with ciliate margins, strongly 1-nerved with prominent reticulations: capsule 2-valved, sessile, 4-seeded: seeds dark brown 2.5™™ long, oblong, triangular, pitted. * The first four new species have been in manuscript more than two years, and the types are in the herbarium of Stanford University. 59 310 BOTANICAL GAZETTE [MAY Type specimen collected in May 1903, by Miss Laura M. Lathrop at Her. nandez, San Benito County, California. This species can be distinguished readily by its reticulately nerved, broadly ovate, and ciliate mature calyx lobes, not unlike the bracts of Acanthomintha ilicijolia Gray. Trichostema rubisepalum, n. sp.—Erect annual, 2 to 3° high: stems chiefly branched from near the base, the branches usually in pairs and ascending, soit pilose and glandular, the lower ones becom- ing reddish: leaves cauline, opposite, entire, subsessile, linear- lanceolate, pilose on both sides and finely glandular, acute or acumi- nate, 2°™ long or longer, about 5™™ wide: inflorescence in axillary short-pedunculate cymes; flowers blue, solitary, on 2 or 3 glandular pubescent pedicels, subtended by linear bracts: calyx united below the middle, bristly pilose and somewhat glandular, about 6™™ long; _ the 5 subequal segments straight, acute, ultimately turning to a pink or light purple: corolla exceeding the calyx, 7™™ long, curved, pilose, throat oblique; its segments also pilose, thin, obscurely bilabiate; upper lip somewhat shorter and 2-segmented; lower one with 3 segments: anthers much exceeding the corolla, in two pairs of unequal lengths; filaments curved, equaling the tube, slender, glabrous, apparently adnate to the entire length of the thin corolla tube; anther cells united toward the apexonly, attached dorsally to the filament, ovoid, the base ultimately much spreading: style glabrous, filiform, recurved, equaling the shorter stamens and insert- ed in the depression of the ovary lobes; ovary short pubescent, deeply 4-lobed: seeds amphitropous. pe specimen collected by Miss Laura M. Lathrop at Hernandez, San Ben- ito County, California, August 1go2. This is closely related to T. laxum Gray, but distinguished by its lang pilose and glandular pubescence, sessile or subsessile leaves, and by its usually pilose corolla. The tips of the sepals soon turn red. Collinsia Hernandezii, n. sp.—Annual, 10 to 20°™ high: stems branched from the base, central ones erect, the outer reclining, soft yellowish pubescent, glandular: leaves cauline, opposite, oblong to oblanceolate, the larger ones 4°™ long, 1.5°™ wide, apex obtusely rounded, gradually tapering at the base to a 1°™ long pubescent petiole, margins entire, short and dirty glandular pubescent on both sides, rather thick, the 3 to 5 obscure nerves parallel; upper leaves ES os eee 1906] ELMER—NEW WESTERN PLANTS 311 becoming bract-like: flowers large, widely scattered along the spicate racemes, half nodding upon short glandular pubescent peduncles, subtended by leaf-like bracts: the 5 distinct sepals glandular pubes- cent, 5™™ long, 1.5™™ wide at base, acuminate: corolla bluish, 2°™ long, strongly bilabiate, saccate at base, with gibbose throat; lower lip obscurely 3-lobed, the middle lobe longer and prominently con- duplicate; upper lip ascending, shorter, and broadly bilobed: fertile stamens 4, equally inserted upon the tube near thé base, longer pair 1.5°™ long, shorter pair 2™™ less, jointed and papillate at base; filaments winged, downwardly recurved, glabrous or the longer ones glandular above the middle; fifth stamen represented by an oblong membranous pouch on the lower portion of the corolla tube; anthers 2-celled, round or reniform, cells united at apex: ovary soft pubes- cent and finely glandular; style usually straight, thick, about equal- a the stamens, sparsely glandular toward the base. specimen collected by Miss Laura M. Lathrop at Hernandez, San Ben- ito ce California, June 1903. Its habit and pubescence is that of C. Greenei Gray, but the leaves and flowers are different. Fritillaria succulenta, n. sp.—Stems glabrous, erect, simple, 2 to 3°" high, from a bulb of fleshy scales: basal leaves ascending, 5 to 10°™ long, 2 to 3°™ wide, in pairs or in whorls of three, elliptic-oblong, obtuse, succulent and covered with a bloom; cauline ones few, erect, © alternate or the lower in pairs, lanceolate, also fleshy and glaucous: flowers solitary or on the larger plants in racemes of three, nodding; peduncle subtended by a leaf-like bract, glabrous, 1 to 2°™ long: perianth campanulate, 2 to 3°™ long, wider than that when spreading; the 6 segments 2 to 3°™ long, obtuse, oblanceolate to obovate, glab- rous beneath, purple, entire, margins at apex yellowish, numerously striate with darker purple and the upper surface pulverulent or obscurely crested: stamens 6, inserted upon the base of the segments, included; anthers 3 to 4™™ long, elliptic-oblong, versatile, extrorse; filaments 8™™ long, glabrous, more or less expanded toward the base: style 1°™ long, glabrous, cleft into three segments half way down, the recurved segments subcompressed and bearing terminal stigmas; ovary smooth, truncate at apex. Type specimen collected in April 1903 by Miss Laura M. Lathrop, at Her- nandez, San Benito County, California. . 312 BOTANICAL GAZETTE [MAY Its leaves are quite thick and fleshy, and are usually glaucous on both sides. Sanicula serpentina, n. sp.—Low spreading biennial or perennial herb, from slender rootstocks, 2 high or less, wholly glabrous and frequently somewhat glaucous, a rich brown color when cured: stems chiefly branched from the base, the central one erect, the marginal ones ascending: leaves mostly from the base, subtending the branches, the radical ones upon membranously flattened 3-nerved petioles 2°™ long; blade proper 2°™ long or longer, ovate in outline, 3°™ across the base, pinnately divided into laciniate lobes which are again divided into slender acuminate usually somewhat recurved and sharply pointed segments: inflorescence branched from near the base, long-pedunculate; involucre of sessile leaf-like bracts; heads 3 to 5, the peduncles of the lateral heads usually much shorter at least when in flower, densely flowered, about 4™™ in diameter; involucels of entire lanceolate bracts slightly shorter than the yellow flowers; marginal flowers sterile, pedicelled, the fewer inner ones sessile and fertile: calyx 5-toothed: petals 1-nerved, quite broad across the middle, the setaceously acuminate apex strongly inflexed and emarginate on its bend: stamens incurved near the apex; anthers broadly elliptic, o.5™™ long: ovary with uncinate prickles; styles 2, slender, recurved, each persistent from the inner face of the stylopodium: fruit not observed. ype specimen no. 4498, collected in April 1903 near Portola, San Mateo County, California. This form is nearest related to S. laciniata H. and A., but the latter is a much more rigid herb, with coarser, broader, spinosely toothed leaf divisions; and with the bracts of the involucels often 3-parted or at least 3-nerved from near the middle. It was discovered on serpentine rocks near Searsville Lake, of the chaparral formation. Trifolium bicephalum, n. sp.—More or less tufted, from an annual root: stems slender, 8 to 18°™ long, decumbent or the outer ones prostrate, rather numerous from the base, rarely branched, sparsely pubescent: leaves both radical and cauline, the basal ones somewhat smaller and more numerous, with slender flexuous pubes- cent petioles 2°™ in length; stipules adnate, 6™™ long, membranous, strongly nerved, subglabrous or finely ciliate along the edges, termi- nated by two setae 2™™ long; leaflets 7™™ long, 4™™ wide, soft pubes- 1906] ELMER—NEW WESTERN PLANTS 313 cent with brownish hairs, obovate or truncate and usually emarginate, entire or obscurely dentate above the middle, with prominent ascend- ing nerves beneath: peduncles pubescent, equaling half the length of the stem or branch, subflexuose above the middle; bicephalous heads terminal, sessile, unequal in size, each subtended by a sub- sessile trifoliate leaf with broad ovate stipules; involucre none: calyx densely pubescent, 4™™ long including the 2™™ sharply acuminate teeth: corolla exceeding the calyx teeth by 1 or 2™™, hyaline and united with a stamineal tube below the middle; upper lip whitish, obovately rounded and surrounding the lateral lobes or wings; lateral lobes oblongish, slightly shorter than the banner, obtuse or acute apical portion nearly white, the middle portion purple, the basal portion hyaline and with an auricle; keel obtuse, shorter than the wings, purplish: anthers very small: ovary glabrous, 2-ovulate; style glabrous, equaling the stamens, terminally recurved, bearing a capitate stigma. Type specimen no. 4812, collected at San Pedro, San Mateo County, Cali- fornia, May 1903. his species comes nearest to the so-called Californian T. Macruei H. and A., but is a much smaller and more prostrate clover, with leaves distinctly obo- vate and emarginate. It forms dense prostrate mats on a high promontory near the sea. Eriophyllum Greenei, n. sp.—A cespitose perennial, from a woody base: stems many from the crown, lanate, branched above the middle: leaves numerous on the sterile stem, the lower ones opposite, the upper ones alternate, petiolate, triangularly ovate in outline, 2 to 3-pinnately divided, lanose on both sides; the segments short, blunt, very narrow, with incurved margins inclosing a dense matrix of woolly hairs; petiole about equaling the blade, as broad as the segments with edges incurved: heads heterogamous, solitary, terminating the leafy branches, ovoid, 1°™ broad; peduncle white tomentose, without bracts; involucre quite rigid, cup-shaped, densely lanate, united at base; bracts in one series, acute, 1o™™ in length: ray flowers light yellow, 15™™ long including the achene, pistillate; tube 2™™ long, pubescent, ligule 8"™ long, 3™™ wide, obovate or oblanceolate, many-nerved, apex obscurely 3-toothed: style arms 1™™ long, narrowly flattened, obtuse: receptacle obscurely pitted, somewhat raised and subconic: disk flowers perfect, very numerous, 314 BOTANICAL GAZETTE [MAY 6°™ long with the achenes: tubular corolla yellow, sparsely pubescent, terminated by 5 obtuse segments: anthers 1.5™™ long, with apex triangularly ovate, bases obscurely auriculate; filaments barely as long, inserted upon the middle of the tube: style arms flattened, bearing small capitate stigmas: achene brownish black, 3™™ long, usually curved and attenuate from the base, subcompressed or 4-angled, edges ciliate: pappus persistent, less than 1™™ in length, of unequal paleae. Type specimen no. 4335, collected in the Mocho Creek Canyon, Alameda County, California, May 1903. Tt is intermediate between E. arachnoideum Greene and E. caespitosum Dougl., but sufficiently distinct from either. Named for Professor E. L. GREENE Navarretia Abramsi, n. sp.—Densely lanose herbs, about 6°” high: stems solitary or several from the base, rigidly erect, chiefly branched from the middle; branches rather stout and straight, ascending, terminated by solitary comparatively large heads: leaves mostly subtending the heads, the larger ones 2°™ long, 1 or 2-lacini- ately divided, soon becoming dry and brittle; the lobes very narrow, becoming glabrous, usually recurved and terminated by a fine sharp point: heads turbinate, the larger ones 1°™. across at top, densely surrounded by a matrix of lanate hairs, 6 to 10-flowered, the sub- tending bracts similar to the leaves but smaller: corolla easily sepa- rating from the base, 8™™ long including the 3™™ long segments, bluish or nearly white, glabrous, hyaline, conspicuously nerved; seg- ments 5, subequally divided, linear spatulate, entire or finely dentate _ at apex: stamens 5, barely exceeding the throat of the corolla, sub- equal in length, filaments threadlike, subequally inserted upon the tube 2™™ below the throat; anthers ovate or elliptic, 1™™ long, apex obtuse, base sagittate: style persistent, glabrous, minutely lobed at apex: calyx of the mature capsule 6™™ long, divided nearly to the base; the sepals straight and erect, linear, hyaline below the middle, held intact by the hairy matrix; the upper part of the sepals glabrous,. foliaceous, and acuminately pointed: capsule triangular, when mature easily falling out from the persistent calyx, straw-colored, smooth and shining, 4™™ long, 1.5™™ in diameter, apex pointed, 3-celled, loculicidally dehiscent: seeds solitary in each cell, subterete, 3™™ long, brown and very hard, with a gelatinous cover which * adily dissolves in water. { ol. 1906] ELMER—NEW WESTERN PLANTS 315 Type specimen no. 4586, collected on Black Mountain, Santa Clara County, California, July 1905. It is a very late summer-flowering annual, chiefly confined to dry gravelly soil immediately bordering thickets of the Californian chamiso (Adenostoma jasci- culatum H. and A.). Named for Mr. L. R. ABRAMS, a former student of botany and classmate at Stanford University. Ribes Stanfordii, n. sp.—A rigidly branched shrub, 1 to 1.5™ high, nearly as broad: bark on the younger branches light brown, becom- ing grayish white with age, thin, separating into shreds; branchlets subtended and protected by 3 spines, very short and rigid; spines about 1°™ long, straight, shining brown, divaricate, distinct, the middle ones usually longer, exceeding the axillary leafy branchlets; branchlets terminated by 1 to 3 small tufts of leaves, subtended by diminutive spines, each tuft provided with a subwhorl of 3 to 5 leaves and terminated by a small inflorescence of 1 to 3 flowers: leaves orbicular, 8™™ long including the finely glandular pubescent 2 to 3"™ long petiole, deeply cleft into 3 segments, soft pubescent on both sides, rather thick, the segments usually terminated by subequal obtusely rounded teeth or lobes, obscurely 3 to 5-nerved; petiole gradually expanded at base into the adnate stipules: flowers 3, upon a short and pubescent peduncle, each separately inserted and sessile, subtended by conspicuously broad pubescent bracts: calyx about the ovary densely pubescent, 3™™ in diameter, its tube 2"™ in diameter, less pubescent, about 2™™ long, the 5 segments exceeding the corolla by 1™™, triangularly obtuse, puberulous on the outer surface, 2™™ long, yellow, rotate or much reflexed: corolla deeper yellow, inserted upon the calyx throat and alternating with the seg- ments, straight, obovate, 2™™ long: anthers 5, inserted upon the calyx throat and opposite the segments, equaling the corolla; fila- ments glabrous, flattened, 1.5™™ long: style erect, subterete, slightly exceeding the stamens: anthers ovate, obtuse at apex, light yellow, truncate or only obscurely lobed at base, 1.5™™ long, 1™™ wide at the base: berry yellow and pubescent at least when young. Type specimen no. 3958, collected on Mt. Pinos near Griffin’s Postoffice, Ventura County, California, July 1902. It was discovered in open pine regions in the vicinity of cliffs and rocky out- croppings at the summit. Not common. Distributed as R. nubigenum Mc- Clatch 316 BOTANICAL GAZETTE [MAY Pedicularis Dudleyi, n. sp.—Perennial herbs, 2 to 3°" high, usually from a branched caudex: stems solitary from each of the scaly crowned caudices, the central ones erect, the outer ones ascend- ing, simple or sometimes branched, not exceeding the basal leaves, lanose especially toward the base, more or less curved; basal bracts brown, lanceolate, entire, glabrous, marcescent: leaves chiefly from the base of the stem, alternate, the uppermost at about the middle of the stem but not exceeding it, the lowest ones longest and some- what decumbent, lanceolate in outline, the larger 25°™ long, 6°™ wide; leaf segments about to pairs, subglabrous or short pubescent on the nerves, membranous; lower pairs distinctly petiolate, the upper pairs not only sessile but broadly united, those along the middle largest; each lobe ovate or oblong in outline, 3°™ long, 2°™ wide, cleft into irregular lobes or merely dentate, the margins unequally serrate or dentate, its teeth sharply pointed: inflorescence spicate, densely flowered, at most 5°™ long and 3°™ in diameter, upon peduncles equaling half the length of the stem, usually erect but frequently somewhat curved; bracts subtending the flowers, foliaceous, serrately toothed, the upper ones equaling the flowers, the lower ones much exceeding them: calyx 1°™ long, unequally 5-cleft, the segments acute and obscurely toothed toward the apex, densely lanose on the exterior: corolla 2°™ long, the narrow tubular part half that in length, glabrous; upper lip conduplicate, slightly notched at apex, pink or whitish, much protruding and arched; lower lip subequally 3-toothed: stamens equal; filaments glabrous; anthers broadly elliptic, attached to the basal dorsal side, the cells connate and rounded at apex, the base not united, acute: style filiform, sub- persistent, thicker and more or less flattened toward the apex, con- spicuously recurved and protruding from the upper lip; ovary glab- rous, dark brown, 2-celled, flattened, acuminately pointed: capsule coriaceous, 12™™ long, 7™™ wide, acuminately terminating in an upwardly curved point: seeds about 4, black when mature, pitted, subterete or obscurely angular. 3 Good flower and fruit of this type specimen, no. 4289, was collected in May and June 1903, at Iverson’s Ranch on the Pescadero Creek, San Mateo County, ornia. Only known from this locality, where it is rare and confined to the deep shade of Sequoia sempervirens. Its proximity to a camping ground endangers its exist- 1906] ELMER—NEW WESTERN PLANTS I 317 ence. This denizen of the Santa Cruz Mountain redwoods is named in honor of Professor W. R. Duprey of Stanford University, who first discovered it. Orthocarpus longispicatus, n. sp.—A profusely branched decum- bent biennial or perennial, forming rather dense mats: stems slender, elongated and distantly branched, often 1™ in length, usually pubes- cent with soft glistening white hairs: leaves alternate, evenly scat- tered, sessile, membranous, puberulent on both sides, or with glisten- ing hairs on the margins and along the 3 obscure nerves, cleft into 2 pairs of strap-like segments, the middle one longest: inflorescence spicate, very long and usually curved; bracts not exceeding the flowers, 5 to 7-laciniately cleft, the obtuse apices light red: calyx 4-cleft, soft pubescent, equaling the corolla, with colored tips: tube of the corolla 2°™ long, externally pubescent, gradually expanded from the constriction above the ovary; upper lip 1™™ longer than the lower one, rather straight, apex obtuse, finely pubescent on the lower surface, margins soft and hyaline; lower lip with 3 obtuse finely pubescent teeth which bear moderate sized sacs: stamens 4, inserted upon the middle of the corolla tube, the lateral pair shorter, the upper pair nearly equaling the galea and enclosed by it; filaments hyaline, linear-flattened, glabrous; the upper anther cell usually somewhat longer and shedding its pollen before the lower one: style persistent, glabrous, much exserted, thickened or expanded toward the base of the capitate flattened or obscurely lobed stigma; ovary oblong, truncate at the apex: capsule 10o™™ long, smooth, loculicidal: seeds numerous, lenticular, with broad reticulate wings. ype specimen no. 4938, collected in July 1903, at Point Reyes, Marin County, California. It was quite abundant among the pickle weed (Salicornia ambigua Michx.), along edges of brackish water. Distinguished by its long decumbent fragile stems and branches, numerous leaves, and elongated densely flowered spikes. Godetia lanata, n. sp.—Erect annual, 3 to 6% high, single or branched from near the base, quite rigid; mature stems shining, straw-colored, scaling at base into membranous shreds, with ascend- ing branches from or above the middle; the younger branches yellow- ish tomentose: leaves cauline, lower ones soon falling, alternate and clustered, sessile, very unequal, cinereous on both sides, semicoriace- ous, lanceolate or linear-oblong, equally tapering at both ends, acute, the larger ones 5°™ long, 15™™ wide, midnerve quite promi- 318 BOTANICAL GAZETTE [MAY nent beneath, lateral ones obscure: inflorescence short, spicate or subcapitate, terminating the branches, usually densely flowered, 3 or 4°™ in diameter; buds erect, from the terminal central axis; flowers easily separating from the ovary, subtended by strigose lanceolate acuminate bracts, subsessile: calyx tube obconic, 3 to 4™™ long, lanose pubescent; its 5 equally pubescent segments 12™™ long, acuminate and ultimately reflexed: corolla and stamens inserted upon the rim of the calyx throat; petals straight, thin, pink, broadly obovate in outline, 6™™ long and wide, irregularly or obscurely 3- toothed, the middle tooth acute, usually the larger: stamens 8, in 2 series, those alternating with the petals nearly equaling them, those opposite the petals barely more than 1™™ long; filaments glab- rous, compressed, broadest at the base; anthers introrse, basifixed, those of the upper series twice as long as those of the lower: style glabrous. or with a few long hairs, barely equaling the stamens, bearing an obscurely 4-lobed stigma; ovary densely and persistently lanose pubescent, upon short thick pedicels: capsules subtended by leaf-like bracts longer than themselves, loculicidally dehiscent from the apex, straight, erect, lanose, subsessile, apex truncate, nearly of the same thickness throughout, subterete or only slightly 4-sided, 8-costate, 4-celled, 4-valved: seeds numerous, in single rows, dark brown, subterete or cubical, a little pointed at one end. ype specimen no. 4376, collected in June 1903 at Bardins railroad switch, Monterey County, California. This characteristic species was found quite plentiful on the sandy plain between Monterey and Castroville, and is quite variable in the density of its pubescence branching habit, and size of leaves. Pentachaeta laxa, n. sp.—A lax very much branched annual, 1 to 2 or 3™ high: stems branched from the base, softly but sparsely pilose: leaves in pairs, subtending the branches, sessile, linear, gradu- ally tapering from the base, the larger ones 3°™ long, 2™™ wide, very thin, sparsely pilose on both surfaces: heads terminal, heterogamous, turbinate, 6™™ long, about 9-flowered; the peduncle ascending, 1 to 4°™ long, pilose, somewhat thickened toward the apex; involucral bracts 3 to 5, persistent, flat, acute, oblong, scantily pilose on the exterior, the reticulate nerves quite prominent, equaling the flowers, more or less membranous: receptacle pitted: each of the ray flowers | a ee ae ER ea 1906] ELMER—NEW WESTERN PLANTS 319 subtended by an involucral bract, yellow, caducous, pistillate, tube 1™™ long, bearing a broad 1™™ long notched ligule: style arms exceeding the ligule, recurved, acute: disk flowers perfect, tubular, light yellow and caducous, 2™™ long, the upper half inflated, bearing 4 obtuse teeth: anthers well included, o.5™™ long, rather broad, bases truncate, each with a very prominent apical appendage; fila- ments thread-like, scarcely longer than the anther: style barely exceeding the corolla, its arms subcompressed, recurved, short, obtuse or truncate at apex: achenes subterete, 5™™ long, black when mature, finely rugose, dotted with sessile yellowish brown glands subtended by short setae; pappus of 2 or 3 paleaceous scab- rous awns, usually persistent. ype specimen no. 4437, collected in May 1903, on Cedar Mountain of the Mount Hamilton Range, Alameda County, California. This distinct species inhabits a steep shaded ravine of fertile soil, and forms a tangled mass with its numerous decumbent branches. Not observed elsewhere, and evidently very rare. ~»Nemophila Fremontii, n. sp.—Delicate annual: stems procum- bent or prostrate, branched, subglabrous or sparsely retrorsely pubes- cent, 10 to 30°" in length: radical leaves similar to the lower cauline ones, frequently forming a rosette, 3°™ long including the 1.5°™ long strigose petiole; blade membranous, ovate or oblong in outline, usually oddly pinnate with two pairs of nearly divided lobes or the uppermost merely sinuate, sparsely pubescent on both sides, paler beneath; the lobes nearly as broad as long, rounded, finely mucronate: flowers solitary, upon slender flexuose usually recurved 2°™ Jong peduncles which are clothed with retrorse bristles: calyx persistent, campanulate, 3™™ long, the basal one-third united, equal- ing or exceeding the corolla, pubescent with fine bristle-like hairs; sepals oblong, obtuse or acutish, foliaceous, with a short blunt recurved appendage from each sinus: corolla white, urn-shaped, its lobes becoming reflexed, at most 3™™ long, 5-cleft, the basal appendages quite obsolete; petals hyaline, ovate to oblong or obovate, obtuse, sparsely ciliate on the edges above the middle: anthers 5, alternate with the petals, erect, quite a little shorter than the corolla; filaments inserted half way down on the corolla tube, glabrous, 1™™ in length; anthers 0.3™™ long, comparatively broad, apex obtuse, 320 BOTANICAL GAZETTE [MAY base subcordate: ovary sessile, densely pubescent: style 1™™ long, cleft into 2 recurved arms, terete, glabrous, persistent; stigma ter- minal, capitate: capsule 4™™ in diameter, globular, sparsely ciliate: seeds compressed, carunculate. Type specimen no. 4991, collected in May 1903, on Fremont’s Peak of the Gabilan mountains, San Benito County, California. It was observed only at the very summit of the peak, among the moss-covered rocks, Monardella franciscana, n. sp.—A sprawling suffrutescent peren- nial: lower stems reclining on the ground or in dense herbaceous thickets, woody, one or more meters long; the leaf-bearing upper stems usually clustered, herbaceous, woolly pubescent, 2 to 3° long, erect or decumbent near the base: leaves opposite, mostly fascicled, very variable in size, densely woolly pubescent on both sides especially beneath; the larger upper ones 2 to 3°™ long including the 5™™ long petiole, 2°™ wide near the base, broadly ovate, entire or with a few obscure teeth, the edges recurved; the lower or axillary ones sessile, lanceolate to elliptic-obovate: inflorescence capitate; heads densely flowered, terminal, rarely more than one, 3 to 4°™ in diameter, sub- tended by a whorl of pubescent leaf-like bracts equaling or exceed- ing the flowers; flowers upon stout short pedicels: calyx about 8™™ long, the marginal ones usually curved upward, conspicuously 11 to 15-nerved, silky pubescent except near the base, tubular, equally 5-toothed; the teeth acute, 1.5™™ in length: corolla blue or light pink, funnel-shaped, the longest 2°™, strigose about the middle, glabrous toward the base, bilabiate; upper lip 5™™ long, erect or straight, apex 2-lobed; lower lip divided into 3 linear segments, equaling the upper lip, usually deflexed: stamens 4, fertile, moder- ately unequal, exserted and spreading; filaments slender, glabrous, inserted at the corolla throat or a trifle beneath it; anthers attached to the base, the cells somewhat recurved: style equaling the stamens, glabrous; stigma minute, terminal; ovary glabrous, distinctly 4-lobed. Type specimen no. 4766, collected at San Pedro, San Mateo County, Cali- fornia, July 1903. It was repeatedly observed in dense herbaceous growths in the ravines on the coast from San Francisco to Santa Cruz, and is a distinct seacoast species. a ee ers ‘How to Read You THE PHELPS COMPANY 106 STATE STREET DETROIT, U.S.A. Myers 18" ST &IRVING PLACE, NEW YORK CITY. | festation of Go Christian Belief Interpreted by Christian Experience THE BARROWS LECTURES By CHARLES CUTHBERT HALL, President of the Union Theological Seminary, New York. “| The volume contains a series of lectures delivered in 1902 and 1903 in India, Ceylon, and Japan. 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BARNES, with the assistance of other members of the botanical Mall of the Guivenicy of Chicago. Vol. XLL No. 6 e Issued June 30, 1906 CONTENTS SOME STUDIES REGARDING THE BIOLOGY OF BUDS AND TWIGS IN WINTER (WITH EIGHT FIGURES). Karl M. Wiegand eee THE LIFE HISTORY OF POLYSIPHONIA VIOLACEA. CONTRIBUTIONS FROM THE HULL BoranicaL Laporatory LXXXIII. Shigeo Yamanouchi - 425 THE STRUCTURE AND DEVELOPMENT OF THE BARK IN THE SASSAFRAS (WITH NINE FIGURES). Howard Frederick Weiss - 434 BRIEFER ARTICLES. THE DIstRIBUTION AND Hairs OF SOME Common Oaks. E. J. Hill - - - = us CURRENT LITERATURE. en REVIEWS - - - - - = = a Es my ae a = 448 ANICAL DICTIONARY. MINOR NOTICES ee eters ent nec a ee pereat iron aires anernnepfimmmee ree orton orl NOTES FOR STUDENTS 4. ee tg eS gee Rs Ber ee ey ek tS ey NEWS mm > - - = -~ - - ~ ow. = e -~ - - - - 45 Communications for the Editors should be add d to tl the University of Chicago, Chicago, Ill. 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GAZETTE JUNE, 1906 SOME STUDIES REGARDING THE BIOLOGY OF BUDS AND TWIGS IN WINTER.’ Karu M. WIEGAND. (WITH EIGHT FIGURES) Durinc the winter months in temperate and arctic climates, the meristematic tissues of shrubs and trees assume a more or less completely dormant or resting condition, and become separated from the surrounding atmosphere by tissues of varying thickness and varying degrees of resistance to the passage of water vapor. A detailed study of these structures during the cold period has brought out many interesting facts ordinarily escaping casual observation. In the twigs the cells of the cambium lie close together without intercellular spaces, but the cortical cells usually do not touch at the corners, and consequently in the cortex there is a more or less elaborate system of intercellular spaces. The main structural pro- tective measure seems to be the firm epidermal layer with heavily cutinized outer wall, which is always present at this period. There were no stomates on the twigs in any of the species I exam- ined. Gas diffusion takes place mainly through the lenticels; but perhaps to a slight extent also through the cuticle its:lf. All the living cells contain a large amount of water, 51-55% in most fruit trees, 63% in Forsythia, and the quantity in each species is remark- ably constant, rarely varying more than four to five per cent., and usually even much less. Regarding the time during the ‘ital summer when the bud fundament is first distinguishable, ALBERT? found that, out of r aay from the Department of Botany of Cornell University. No. 105. 2 ALBERT, P., Beitrage zur ee der Knospen einiger Laub- hélzer. Forstlich naturw. Zeitschr. -3:345, 393- 1894- Shoe 374 BOTANICAL GAZETTE [JUNE 15 species of trees bearing scaly buds, the first leaf fundament in one (Betula alba) was present as early as May, in three at the beginning of June, in eight at the beginning of July, in two August 1, and in one not until September. The flowers were always formed later than the leaves. Some of the naked buds he found to start early in the previous season (Elaeagnus, Cornus); others, as for instance Robinia Pseudacacia, did not start until the spring of the year in which they were to unfold. He found that in general the buds were further progressed at the beginning of the winter the farther north the plants were native. BEHRENS? found that in fruit trees the Sewers are first distin- guishable at a later date, as for example, in the cherry during July, and in the pear about August 11, My own observations lead me to believe that in many cases the fundaments are present quite early. The buds of the peach were well formed July 15, and small buds were evident in the leaf-axils of forest trees as early as June 1. This suggests that, in some cases at least, the bud fundament may be present as early as the unfolding of the previous winter’s buds. Those that start quite early have usually reached an advanced stage in development by the time cold weather overtakes them in the fall. The rudimentary flower or shoot for the next season, together with all its organs, is present in the buds of some species, as for instance in the horsechestnut; while in others a varying number of nodes and internodes are thus stored. To inclose so elaborate a structure a certain number of leaves have been modified into scales which closely overlap, or are firmly cemented together at their edges around the young shoot. Such buds are found espe- cially upon treés and shrubs with definite annual growth. The scales are usually composed of several layers of parenchymatous cells with intercellular spaces, moderately firm and slightly cuti- nized epidermis, on the inner side, and a very strong heavily cuti- nized outer epidermis, usually supported by mechanical tissue of varying amount beneath. The parenchymatous cells of all or 3 BEHRENS, J., Entwickelung und Bau der Bliitenknospen unserer Obstbaume und Obststraucher. Gartenflora 4'7:269. 1898. 4 For published descriptions of bud structure see: BEHRENS, J., /. ¢.; FEIST, A., Ueber die Schutzeinrichtungen der Laubknospen dicotyler Laubbaume wahrend 1906] WIEGAND—BUDS AND TWIGS IN WINTER 375 all but the outermost scales are living and contain a large amount of water throughout the winter. The inner scales are frequently almost destitute of epidermal thickenings and are quite green and fresh. Because of the much larger size of the cells in the scales, and much larger vacuoles, there is much more water present in these structures than in the young shoot whose cells are small and nearly filled with protoplasm. This it will be seen is an important con- sideration when the buds freeze during the winter. The abso- lute amount of water in the whole bud is however very nearly the same as that in the young bark, being about 51 to 55% for the fruit buds examined; and, as in the bark, this amount is remarkably constant for the species. The proportion of space occupied by the young shoot varies with the species and nature of the bud. In flower buds this proportion is usually greater than in leaf buds. In many cases only a very small fraction of the total volume is shoot-tissue, all the rest being composed of scales; but in other cases, as for instance the flower buds of pine, almost the whole volume is made up of cones, leaves, and stem; while the scales are very thin, dry, and firmly ‘cemented together. In this case of course nearly all the water is located within the young shoot. The spaces between the various organs and scales usually contain air alone; but in some cases, as for example in apple and horsechestnut, there is also a large amount of wool present in which the organs are seemingly imbedded. In Populus and some other trees the spaces are more or less completely filled with resin. Buds of most indefinite growers differ from those of the majority of definite growing trees in two essential ways: in the slight develop- ment of the fundament, and in the usual absence of scales. The young shoot is most frequently represented merely by a growing ihrer Entwickelung. Nova Act. Leop. Carol. Ak. Naturf. 2: 303-344, 1887, Ref. Bot. Centralb. 36:43. 1888; ScHUMANN, C. R. G., Anatomische Studien iiber die Knos- Penschuppen von Coniferen und dicotylen Holzgewachsen. Biblioth. Botan. 15:32. Cassel, 1889, Ref. Bot. Centralb. 42:275. 1890; Griss, J., saat zur Biologie der Knospen. Jahrb. Wiss. Bot. 23:637. 1892; Lussock, J., On buds and stipules. Jour. Linn. Soc. 30: 463-532. 1895; 33:202-269. 1897; CapuRA, R., Physiologische Anatomie der Knospendecken dicotyler Laubbaiume. Breslau, 1887, pp. 42; Mrkosca, K., Beitrige zur Anatomie und Morphologie der Knospendecken dicotyler Holzge- Wichse, Sitz. Konig. Akad. Wiss. Wien Math. Wiss. Kl. 74':723-755- 1877- 376 BOTANICAL GAZETTE [JUNE point without well-developed lateral organs, and can therefore be protected more economically by being sunk in a pit produced by a ring-like growth of cortex and cork, as is commonly the case. This pit is then closed at the mouth by an ingrowth of the cork itself, as in Gleditschia, or by a feltlike mass of hair, as in Robinia and other species. . In the case of the large buds with the shoot considerably advanced in growth, the bud-scale method seems the only feasible way of ' covering them. Another advantage in this method lies in the tele- scopic expansion of which scaly buds are capable early in the season while unfolding. Growth is thus permitted, but at the same time the protective qualities are not lost. In the maples and_horse- chestnut the tube formed by the enlarged scales often reaches the length of 2 to 8°". By this means buds may open early in the spring and still be protected from excessive transpiration. Scaleless buds usually remain nearly dormant until later in the spring when the weather conditions are not so severe. PHYSICAL PHENOMENA OF BUDS AND TWIGS WHEN NOT FROZEN. The scaleless buds of the indefinite growing trees and shrubs grow very little before or during the winter. In the autumn the very limited growth is soon stopped by the advent of cold weather, and from this time until late spring scarcely any change can be detected. With the scaly buds however it is otherwise. From the inception of the fundament in July or June until cold weather there is a very considerable growth resulting in the buds of various sizes found upon the different species of trees during the winter. Little accurate work has been done towards determining the char- acteristics of this growth, but the results obtained in our laboratory by W. M. Morcan during the fall of 1901 seem to show that in the case of fruit trees the growth is very uniform and gradual up to about November 15. In some cases slight fluctuations occurred which could not be accounted for, and in one or two instances these seemed periodic; but on the whole there appeared neither accel- eration nor retardation until the time mentioned, when the increase in size ceased quite abruptly. From the middle of November until March 1 there was no growth in peach buds, the curve remaining almost exactly horizontal and fluctuating very little. On March 23, 1906] WIEGAND—BUDS AND TWIGS IN WINTER 377 several days of warm weather occurring, the peach buds began to grow rapidly and uniformly until April 23, one month later, when they came into flower. With the apple and apricot the results were very much the same. Growth almost ceased November 15, and from this time until March 1 the increase was apparent but exceedingly slight, amounting to only $ to 1%. Renewal of activ- ity began March 1, and from this time until April 23, seven weeks later, when the apricots flowered, and eight weeks later, when apple buds opened, the growth was very rapid. The curve after growth began was not so gradual as in the peach, but became much accel- erated just before the flowers appeared. Mr. Morcan’s observations were as a matter of fact quite exten- sive, but only the above summary can be given here. At intervals of one week through the fall, winter, and spring, buds were taken from the same tree and as nearly as possible from shoots of the same vigor, a large number were measured, and the average taken as representing the size at that time. It was found impracticable to measure the same bud at different times, owing to the difficulty of manipulating the micrometer out of doors on very cold days, as well as to the fact that the measurements were liable to be taken at different temperatures cach time. A Zeiss cover-glass measurer was found the most convenient instrument for the work. From the tables thus made a great many curves were plotted representing the changes in various fruit trees. The results, however, agreed very well, and in the peach, apricot, and apple were as stated above. From these careful observations, therefore, contrary to the general belief, it seems that fruit buds at least do not grow to any extent in winter. Their swelling period is confined in the north to a few weeks just previous to the opening of the bud. Regarding our forest trees and shrubs no accurate work seems to have been done toward the determination of their curve of growth. From casual observations, ‘I am inclined to believe that a majority will be found to agree with the fruit buds. This seems to be truce of the sugar maple, whose buds are practically as large in November as in early March, also of the ash, oak, etc. On the other hand, the buds of a few plants, as, for instance, Salix discolor, Ulmus fulva, and Ulmus scabra, seem to increase in size early in February. However, actual measurements are necessary to determine these. points. 378 BOTANICAL GAZETTE [JUNE Kuster’ found that during a specially mild winter the buds of maple did show a very slight growth both in the lateral organs and in the young axis. No new organs were started either in maple or other species examined, except rarely in Alnus cordijolia. ALBERT® found that practically all buds became dormant soon after leaf-fall until spring again. The first change in spring was a stretching of the tissues, further development of the parts taking place only later. It is a known fact in physics that the amount of heat absorption varies, among other factors, with the color of the body investigated. In other words, the same body if colored differently will absorb a varying amount of heat from a constant source, depending upon the color. Winter buds and branches are in many cases highly colored, and the question naturally arises as to how this affects the heat absorption of the bud during the winter and spring months. Regarding the extent to which color will affect heat absorp- tion, in addition to the records in works on physics, the experiments of WHITTEN’ are interesting. He found that thermometer bulbs wrapped in muslin of different colors, green, purple, black, and white, or with pieces of muslin of these various colors spread over them, or with the bulbs coated with a wash of similar colors, showed a marked difference in reading when exposed to bright sunlight. The average difference between the black- and white-washed bulbs was 16°, between the white and purple 15°, and between the white and green 13°. At one time a difference of as much as 21° between the white and purple bulbs was found. However, the actual experiments with buds have been rather few and the results are not so definite as one might wish. The most elaborate were those of WHITTEN described in the above-cited report. He selected a row of peach trees containing several vari- eties, and whitewashed them during the winter. During warm days of the unusually changeable winter the unwhitened buds swelled considerably, and during subsequent cold spells most of 5 KisTEr, E., Ueber das Wachsthum der Knospen wihrend des Winters. Beitr. Wiss. Bot. (FUNFSTUCK) 2:401. 1898. 6 ALBERT, P., Beitrage zur Entwickelungsgeschichte der Knospen einiger Laub- hdlzer. Forstl.-naturw. Zeitschr. 3: 345-376, 393-419. 1894. 7 WHITTEN, J. C., Winter protection of the peach. Bull. 38. Missouri Agric. Exp. Station, April 1897. & | | : | 1906] WIEGAND—BUDS AND TWIGS IN WINTER 379 them were killed. The unwhitened buds swelled and grew percep- tibly before any swelling could be detected in those that were whitened. The difference in size March 20 was plainly shown in drawings of the sections of the two classes of buds. Whitened trees came into bloom about one day later than unwhitened trees of the same variety. In 1896-97, owing to a more moderate spring, the differ- ence in time of flowering was still greater. The whitened buds of each variety opened two to six days later than those that were not whitewashed. The differences in the actual time of flowering, however, does not express the difference in time of the swelling of the buds. The whitewashed buds did not begin to swell until almost time for the flowers of normal trees to appear, while the unwhitened ones began to swell three or four weeks earlier, as shown by the drawings above mentioned. These experiments of WHITTEN seem to show that in the peach, at least, the dark-purple color of the buds tends to cause earlier activity in the spring, accompanied by earlier swelling and flower- ing. The only doubt, it seems to me, lies in the effect of the white- wash upon the growing tissue. As mentioned below, some non- porous substances seem to retard respiration perhaps to such an extent that growth also is retarded, but the whitewash would seem porous enough to escape this criticism. In a more recent paper Wuirren® has shown that the temperature within whitened and unwhitened twigs differs by several degrees. In bright sunlight the difference was as much as 15° C., the unwhitened being the warmer. The whitened twigs were nearly of the same tempera- ture as the atmosphere. WuitTENn® has also shown that purple peach twigs transpire considerably more than green ones. This was probably due to the greater temperature and is probably an additional factor in the winter-killing of the peach. 8 WarrreN, J. C., Preventing frost injuries by whitening. Pacific Rural Press 60:276. 1900. 9 WHITTEN, J. C., Das Verhiltnis der Farbe zur Tétung von Pfirsichknospen durch Winterfrost. Inaug. Diss. Halle. 1902. p. 35. See also in this connection Macoun, W. T., Some results of experiments in spraying, etc. (whitewashing ” Tetard bud development.) Ontario Fruit Growers Ass. Rep. 1899: 100, and “ Experi- mental Farms,” Canada, 1899:92. 380 BOTANICAL GAZETTE [JUNE Wishing to determine the effect of natural color and surface of buds upon the absorption of heat, I carried through several experi- ments with horsechestnut buds, which gave some interesting results as follows. On February 8, in bright sunshine, a large horsechestnut bud was obtained and the scales dissected away, care being taken that they were not unnecessarily injured. Two thermometers previously tested as to accuracy were obtained, and over the bulb of one the bud scales were carefully imbricated and firmly held in place by a few turns of black thread. There was enough resin present to cement the scales firmly together and thus form an artificial horse- ° chestnut bud with the thermometer bulb in place of the normal shoot. The instruments were then placed on a table out of doors ’ and in the shade where they were allowed to lie. As soon as the readings were nearly the same,'® the table and instruments were carried to a place in full sunshine, care being taken that the two bulbs projected about 6.5°™ beyond the edge of the table so as not to be affected by direct radiation from the surface of the latter. The readings were taken as follows: ‘TABLE I. Horsechestnut bulb and naked bulb, from shade out of doors to sunlight. (See jig. I.) Maked bein. | Momoeneh to ee ° Difference ge" F. pire 0:00 oO PS (axe €) 33 32 0:10 I mo: 34 33 0:30 I ae 34.5 34 1:10 0.5 0.2 35 35:7 bee fe) 0.7 0.4 36 a5 2:00 I 0.5 36 38 2:30 2 L-6 or 39 3:00 2 EO 37 40 3:30 3 to" 37 42 4:00 5 2.8 38 44 4:30 6 ae 38 45 5:10 7 3.9 38 48 6:00 10 5.5 38 49 7:00 II 6.1 38 50 10:00 rz 6.6 30 5L 14:00 12 6.6 39 52 19:00 13 7.2 3) 53 24:00 14 7.7 39 54 29 :00 Ig 8.3 ‘© They could not be made to read the same because the overcooling point in the bud had been reached. 1906] WIEGAND—BUDS AND TWIGS IN WINTER 381 60 SS Ee 50 —— ri 40 L a a a 380 me oe» « & & &® & & Sk & 8S & SK EF B ———horsechestnut bulb; .<.s.07:- naked bulb. Abscissas represent 5° F.; BIG. he ordinates, 2 minutes. See Table I. No further rise was noted, although the instruments remained in place more than an hour. The experiment was repeated several times during the spring, both with the same bud and with a fresh one, in every case with practically the same result, namely a much faster rise in the bud-bulb, amounting finally to an excess cf 5 to 12° C. over the other bulb. That the above differences were due mainly to the dark brown color of the bud seems probable from another series of experiments in which the naked thermometer-bulb was coated with brown drawing ink. Readings were taken under the same conditions as before. In these cases the difference was but slightly in favor of the horsechestnut bulb, probably because of the less highly pol- ished surface of the ink bulb. It would seem, therefcre, that the point is fairly well demonstrated that in the case of the horsechestnut, at least, the color does make considerable difference in the absorb- ing power of the bud as regards heat. Although no experiments were performed, there seems no reason why the same shculd not be true also of other dark-colored buds. With these results in mind I made a few observations one spring to see if there was any relation between the color of the buds and the time of swelling and opening. It seemed reascnable to expect that the darker the bud, and consequently the more heat absorbed, the earlier the bud would swell and open in the spring; and all the more because this was the exact conclusion reached by WHITTEN with his peach experiments. Unfortunately, accurate record of the time of swelling and of opening were not kept; but still I believe 382 BOTANICAL GAZETTE [JUNE some important general tendencies can be made out from the notes, and therefore the following table is inserted: Name Color of buds Ga Magnolia acummaitay .. fe. osc. use ee yellow-gray or olive late Ailanthus eli having Chg eR ea. bias yellow-gray late Boer Negu nde n.6 naGkbs.- ecards hire a: pale, whitish medium Ju _acsant senha hi pao ta ats be eens se pale, grayish very late Quereus alla. o 53.16 Hees oad wae oes gray very late Salix dite Lc c inte a oraitas. Stara tak ops yellow early Pops Prandidentatd. 455 esses: pale very early Ringe Wulearis. howl asis cows eso pale, yellowish early Betula Bee: ee Or see ee olive late Fagus SnGAiCaUe << os dress se cd euxs olive medium Paes “dilatata ne Cre eae ee yellow-brown medium Salix) COrdatan: 4 tactea erwiss ste itosnattcaaesivs, Ae olive or purple ly Sie TOCA GATIO 6, 5 4.50 Cesena ar ah te een brown medium Sank SeriGed ve coe okie sle, a2 worsiawesae eo). brown-purple early MNES. POUR. se uals oa gs BOF eed wc brown-purple early Acer obec eg wala PRR ata hee bark olive-brown medium its ROIOONA ns oie be Se ee red medium Aesculus Teecectaaet a Us ach otra ah dark brown » early Aber pactariate 6 25 oo ns 8 cae Ys red very early Prunus persica .......-00,.-00ssese> purple very early Other species ee Sohgan and Pyrus.. red, brown, or gray early CYRISORUE COCKER. 0 ooo ce occ deus a red o house olive medium Dilnriis: Hikwase: tcc geseec cure oe heres dark he early NENTS Sen Dita. ch cca c tat ele ees black very early Stee WRN iy vinu ds 64 ex ba eadaes Hae black very early It is not to be expected that the time of opening or even the time of swelling will in all cases be proportional to the color of the buds alone. The protoplasmic characteristics of the particular species or genus undoubtedly play a very important part; but bearing this in mind the following suggestions appear in the above table. Nearly all of the light-colored buds are also late to swell and open. None of the dark red or especially the black buds open late. On the other hand, a few light buds, as for example those of Syringa, Populus, and Salix alba, open quite early. This may be due to more easily aroused protoplasm than is present in most buds. Pos- sibly if these buds were black they would open still earlier and therefore suffer injury from the frost; thus the present lighter color may serve as a means of protection. During the spring of 1901, about March 1, some experiments were started to determine if possible whether other buds might be influenced by color in a way similar to the peach buds white- 1906] WIEGAND—BUDS AND TWIGS IN WINTER 383 _ washed by WauirtTeN. Instead of whitewash, black paint was used to see if they might be made to open earlier. Two kinds of paint were prepared, one made of lamp black mixed with linseed oil, the other of lamp black and xylol. Buds and twigs of Syringa vulgaris, Ailanthus glandulosus, Populus dilatata, and apple were treated to a coat of oil paint; while some others of Syringa, Ailanthus, and apple were coated with xylol paint. The results were as follows: Syringa.—Xylol-painted buds much behind the normal during vernation; they looked unhealthy, one or two being entirely dead. Oil-painted buds never began to swell, all dead. Atlanthus.—The xylol paint made no difference with the killing back of the branches nor with the development of the buds. Oil paint prevented the swelling of the buds; they never opened. A pple.—Xylol-painted buds much behind the others; one completely dead. Oil-painted ones all dead. Populus dilatata.—Oil-painted buds showed much more rapid swelling than normal. When just opening the blackened buds were 6 to 8™™ longer. These results are evidently in the main exactly opposite what we were led to expect. I suspect that the explanation lies in this, that the coating of the surface of the bud with paint prevented res- piration, and thereby inhibited growth just as did varnish used on naked buds as described later in this paper, although it is possible that some toxic property of these substances might have had some- thing to do with the matter. The xylol furnished a much more porous layer than did the oil, and the inhibition was therefore much less. The buds of Populus dilatata are normally almost completely infiltrated with resin in the spaces between the organs and on the surface, and consequently may have some other means of obtaining oxygen for respiration. The coating even with oil paint, therefore, did not injure them. On the contrary, the black color seemed to cause an acceleration in growth. In concluding this part of the subject we may say that in the climate of New York, buds during the winter seem to remain in an almost dormant condition until a short time previous to their opening in the spring. In Missouri swelling of peach buds began much earlier than in New York. Color through its power of absorbing heat seems to have some effect upon the growth of buds in the spring. Early buds are in 384 Pinna ‘BOTANICAL GAZETTE [JUNE most cases dark, and artificial darkening, when unaccompanied by deleterious factors, seems to accelerate the opening. PHYSICAL CONDITIONS IN FROZEN BUDS AND TWIGS. From many inquiries it would seem that very few people are really sure. whether free ice is actually present in buds in winter. Nevertheless, this is one of the most common phenomena connected with the winter condition of trees and shrubs. To put the matter on a firm basis of observation I undertook, during the winter of tgo1, to section buds of various trees during cold pericds and to determine under the microscope the amount of ice present. The method employed was as follows. Early in the morning, at about sunrise, after a fall of temperature to —18° C. or below, a table, microscope, razor, needles, slides, and cover glasses were placed in a shady situation in the open air, where they were allowed to become thoroughly cooled. Free-hand cross sections of the various buds were then made, and mounted on the slide. For a mounting medium cedar-wood oil was found best. A small quantity of this in a vial was allowed to cool with the instruments. One important advantage in the use cf cedar-wood oil over those of a denser nature lay in the fact that it did not congeal at the low temperatures of the experiment. The ice remained unmelted in the preparation and could be observed at leisure; or if the thawing process was under study the slide could be carried to a warm room and placed under another microscope. The melting ice was unable to evapo- rate from the section, and therefore it was easy to determine whether the water was all reabsorbed, and the approximate rate of absorp- tion. The ice was found to occur always in broad prismatic crystals arranged perpendicular to the excreting surface; and usually formed a single continuous layer throughout the mesophyll of the scale or leaf, to accommodate which the cells were often separated to a con- siderable distance (figs. 2, 3, 4). This ice sheet was composed of either one or two layers of the prismatic crystals, depending on the water content of the adjacent surfaces, and was often as thick as the whole normal scale. The cells surrounding the ice, having lost their water content, were in a more or less complete state of collapse, 1906] WIEGAND—BUDS AND TWIGS IN WINTER 385 depending upon the resistance of the walls, and often occupied a space smaller than the ice itself. These cells were uninjured, however, and would resume their normal condition on thawing. In all cases more ice was found in the scale than in the young shoot; never between the scales but always in the mesophyll. The cells of the embryonic shoot were so much smaller and their water content so much less, that frequently it was difficult to detect any ice forma- Fic. 2.—Populus dilatata: cross-section of bud, showing ice in bud-scales and foliage leaves. tion whatever; but ordinarily very minute and numerous masses, at least, were scattered between the cells, and sometimes there were large masses such as appear in the outer organs. In young anthers the ice often filled almost the entire anther cavity, and in it the pollen grains were imbedded in a completely collapsed state. The results of the observations regarding the occurrence of ice in buds may be summarized briefly as follows. The temperature Was ~33;5° C.to = 28°. 386 BOTANICAL GAZETTE [JUNE 1. Tissue packed full of ice in shoot and in mesophyll of scales forming sheets parallel with the surface; rapidly and completely reabsorbed when the sections were thawed in oil. Sponging out of sections very marked.—Populus dilatata (fig. 2) and P. candicans, Prunus serotina and P. virginiana, Betula enta, Acer Negundo, Pyrus Malus and P. communis, Aesculus Hippocasta- num. 2. Containing a large amount of ice, but the water tardily reabsorbed on thawing in oil.—Acer Saccharym, Tilia americana, Ulmus scabra, Crataegus punctata (jig. 3). Fic. 3.—Crataegus punctata: cross-section of bud, showing ice in bud-scales and floral parts. 3. No ice could be found at o°C. Tissue dense, of small cells.—Castanea dentata, Hamamelis virginiana, Fagus americana, Fraxinus americana, Jug- lans cinerea, Corylus rostrata, Quercus alba, Hicoria ovata. 4. Other cases—In Pinus Strobus and P. sylvestris there was a moderate amount of ice in the shoot and in the anther as well as in the inner scales. In Syringa vulgaris there was a very large quantity of ice in the scales and young shoot, especially in the anthers (fig. 4). In Viburnum dentatum and Prunus persica the amount of ice was small, but water was quickly reabsorbed. 1906] WIEGAND—BUDS AND TWIGS IN WINTER 387 Of the twenty-seven plants examined there were only eight that showed no ice in the buds at —18°C. These eight were sectioned later at —26.5° C., with the result that in Castanea, Hicoria, Fraxi- nus, and Juglans numerous minute ice crystals were found between the cells. It would seem, therefore, that ice may be found in all buds if the temperature is sufficiently low. The accompanying illustrations are reproductions of photo- micrographs taken by the writer during periods of low temperature. Fic. 4.—Syringa vulgaris: cross-section through flower bud while frozen; the light spaces filled with ice. When the mercury registered at zero Fahrenheit or below, freehand sections mounted in oil as already described were photographed, the apparatus being set up in the open. The conditions for sec- tioning were so strenuous that very thin sections could not be obtained, and hence the rather poor quality of some of the photographs. The palisade-like ice prisms fill the light areas through the mesophyll of the scales and young leaves. 388 BOTANICAL GAZETTE — [JUNE The question naturally arose as to the cause of the difference in ice content and why ice was absent in the eight species mentioned. Since lowering the temperature from—18° C. to—23.5° C. caused the appearance of ice in some, it would seem therefore to be simply a matter of temperature. But the degree of cold necessary to cause the separation of ice is proportional to the force which holds the water in the tissue. This in turn depends upon the relative proportion of water to cell-wall and protoplasm. We should expect, therefore, to find in those buds which are difficult to freeze a smaller amount of water than in other buds; also smaller cell-structures, since by this latter means the proportion of cell-wall and proto- plasm is increased. When cells become smaller it is usually the water content that most rapidly diminishes, the protoplasm follow- ing at a much lower rate. I have made the following measurements of the cells and water content in seven of the species in which there was much ice, and in seven in which ice did not appear at —18° C. Max. aver. | Min. aver. | Text. of wall | % of water mm. mm. A. Ice abundant in. bud-scales, leaves, nd growing point— Crntecwtis: pronttata. << 670.23 625 oss 0.040 0.012 thin 49-4 RT ENNOS Ste aie ots or een eae Be O51 0.015 : 46.6 Byrn Vileatigg 6.565. ..55 204 254 0.0045 | 0.009 ot 53-2 Pee MN id a deh 0 ware 0.021 0.009 : 45-9 SerNAl, cues ee a 0.021 0.015 ie 47.6 Populus racine LEP Dee wit 8 = sain to 0.025 0.018 2 39-3 meuttile lites es 5 vic 0 Pa Sh 0.018 0.006 se 37-5 B. Ice not sie at — 715" Cc 7 elles Querc Pee PN Sa. 3s th elements 0.015 0.006 thick 22-9 orylus:rostratac: < .0csce 8 oe see: 0.018 0.006 a 29.7 Castanea dertata. .. 28 igs cds ees 0.018 0.015 = 25.4 F Westen. co ree ces | 0.008 0.003 26.8 a aphedia can foe ee ee 0.048 0.015 very thick 31-4 JUG CUMTOR, 055 65 eo eg oe | o12 0.003 thi ck 25:9 Fraxinus americana... ..........-;. 0.021 0.003 | 29.8 Our supposition regarding the smaller size of the cells and smaller water-content in the second group, therefore, seems to be upheld by these results. In the twigs ice is also present in very cold weather, where it may be found in three different localities. TThe largest quantity occurs in the cortex, where the ice crystallizes in prisms arranged 1906] WIEGAND—BUDS AND TWIGS IN WINTER 389 in single or double series according to the law of freezing tissues. The ice is more frequently in the form of a continuous ring, or really a cylinder, extending entirely around the twig, prying apart the cells of the cortex in which it lies. The outer cylinder of cortex in such twigs is completely separated from the inner layers when frozen. In a few species instead of the continuous layer, lens- shaped ice masses are interpolated irregularly throughout the cortex. The cortical cells after the withdrawal of the water are as com- pletely collapsed as were those in the bud scales, but they also usually regain their normal condition on thawing. In the wood ice rarely forms in large quantities. It is usually confined to small masses in the vessels themselves, or, according to some authors," sometimes extends in radial plates in the pith rays. In sectioning twigs, I myself have never seen ice in the wood elsewhere than in the vessels or wood-cells. In the pith the ice, so far as I have been able to observe, always occurs within the cells and therefore in very small masses. At the time when the buds were sectioned, cross-sections of the twigs were made and mounted in the same manner. Ice was found in the cortex of all those in which it was present in the bud, but usually in proportionately larger quantities. It was also found in the following species which showed no ice in the buds: Corylus rostrata, a small amount in large clefts in the cortex; Castanea dentata, some ice in ordinary small spaces of the cortex but not aggregated; Hamamelis virginiana, a ring of ice completely around the stem in young twigs. In Fraxinus, Fagus, and Juglans none could be found, and Quercus was not investigated. Since water on freezing increases in volume, one would at first thought expect the frozen twigs to be larger in diameter than normal. Such, however, is not the case. In every instance a distinct con- traction occurred, which in some cases was very marked.*? _ ™ MULLER-Tuurcau found ice present in the large vessels of Syringa, Cornus, and in pears, almost completely filling them; and several times he could also dem onstrate it in the wood-cells. The ice was the most distinct, however, in the vessels of the grape. Ueber das Gefrieren und Erfrieren der Pflanzen. II. Landw. Jahrb. ™2 Both Sacas and MU@LLER-TuuRGAU have shown that a similar contraction occurs quite generally when herbaceous tissues freeze. SACHS, J., Krystallbildung 390 BOTANICAL GAZETTE [JUNE To determine the exact amount of contraction the Zeiss cover- glass measurer was used. . Pieces about 10°™ long of one or two year old twigs taken at—18° C. were inserted in the clamps of the machine, a record taken, and then the whole carried to the warm laboratory. The increase in size on thawing could be followed by watching the movement of the indicator as the ice melted, and when at last stationary another reading was taken. Some results are given in the following table: : Frozen Thawed Difference Exp. or contr. Cormis etolodifera < 00.25% 62. 4.3 2.58mm/ 2, 60mm o.o2mm| expanded he oF i oP se acs Eire ares 3.38 3-43 0.05 i "EWR QUGIYDNGUOS Soa ee 2.03 2.10 0.07 si - Feces ok cath gare. tovalvee sid 3.18 3-39 O.12 i Pennies Mlstates 66. Peony os 5: 3-17 3.28 O.11 " ee We is © pe ee gk 4.72 4.80 0.08 i Acer piataniless yee kat 2.82 2.86 0.04 ig Oe es gare hgh as 3.44 3.48 0.04 = Pyrus Malus......... 0.2.0.6... 2.97 3.03 0.06 # WANES Set. yas a arate 3.89 ze 2 0.06 ale MORO Gare nce diy es & 94 5.4 0.10 is pee ne) Sit a ee eS: 5-9 0.14 . Many twigs at — 18° C. or below appear very much wrinkled on the surface as though dried and dead. This is especially true of the polished shoots of Salix cordata. On very cold mornings shoots of this species appear as though dead and dry, the bark being com- pletely covered with fine longitudinal wrinkles. Some of these shoots were brought to the laboratory and allowed to warm, during which process the disappearance of the wrinkles could be watched with ease. In about ten minutes the twigs were entirely smooth and normal. It was from such twigs that the above readings were taken. To show more graphically the expansion during thawing, some twig-sections about 10°™ long were taken from the same willow and the ends while still frozen dipped in melted paraffin. The caps thus produced at the ends of the twig were in every case rup- tured down the side on thawing, leaving in most cases a cleft of considerable size between the two edges. Twigs of plum were bei dem Gefrieren u. Veranderung der Zellhiute bei dem Aufthauen saftiger Pflan- zentheile. Bericht Verhand. Konig. Sachs. Gesell. Wiss. Leipzig, Math.-Phys. Klasse 12:1-50. 1860. Mirrer-Taurcav, H., Ueber das Gefrieren und Erfrieren der Pflanzen. Landw. Talnb. 9:187. 1880. . ————ooo a ee ee 1906] WIEGAND—BUDS AND TWIGS IN WINTER 391 also much wrinkled, those of Negundo and apple showed a slight furrowing, those of black cherry and pear scarcely any at all. The photographs of wrinkled twigs of Salix cordata, reproduced in the accompanying illustrations (fig. 5), were made in the open at a temperature of —20° C. The same twigs were then placed in the laboratory, and after about one hour were photographed again. The slightly wrinkled «appearance in the upper shoot in D Fic. 5.—Salix cordata: A and B, twigs ears in the open at—20° C., showing etic condition due to contraction; C and D, the same twigs thawed in the laboratory; the furrows have disappeared except the minute normal striae on the lower twig. the second photograph was normal for that shoot when thawed, during both winter and summer. It seemed desirable to determine whether this contraction was mainly in the bark or in the wood or in both. At a temperature of —18° C. much wrinkled twigs of Salix cordata were collected, and the following measurements made: 392 BOTANICAL GAZETTE [JUNE With bark on twig the diameter, 7.80™™ expanded to 8.04™™ on thawing; difference o.24™™. With bark removed from a small spot for the clamps of the measuring instrument, the diameter, 5.05™™, expanded to 5.15™™; difference, Lon Therefore more than half of the total expansion was in the bark. Thickness of the bark was 0.5 ™™ on each side; thickness of the wood and pith, 2.05™™ on each side; expansion of the bark, therefore, was 13.5 per cent.; of the wood, only 2.5 per cent. With thicker twigs, containing more hard wood, the expansion would have been still less. Where the bark was whittled away entirely around the end of the twig and for some distance back, the expansion of the wood was not detected; probably because water had passed to the bark to freeze and being removed there was none to cause swelling again when the twig thawed. The explanation of the contraction of twigs on freezing probably lies in the following considerations. When the water is extracted from the walls of the wood-cells, the latter contract to a slight extent just as they do when wood seasons. This accounts for a part of the shrinkage. The rest and greater part occurs in the cortex. Here the intercellular spaces are quite large and numerous, and are normally filled with air. When freezing occurs the ice forms in the spaces and the cells collapse, while the air is mostly driven completely out of the twig. The contraction in the cortex will be approximately equal to the volume of air expelled plus that of the air compressed minus the expansion of the ice while freezing. This: is for contraction in all directions; only a portion of this will be radial, depending upon the structure of the particular species; much the greater part, however, is radial in all twigs. With buds the study is not quite so easy. The record of buds measured at —18° C. and then again after thawing is shown in the adjoining table. From this table it will seem that in all cases, except in Populus and Acer, there was a decided increase in size on freezing and a consequent decrease when thawing out. In the two named cases there was a slight contraction as in the twigs. It is not quite clear why the buds should behave so differently from the twigs. The only explanation I can offer at present is that the contraction of 1906] WIEGAND—BUDS AND TWIGS IN WINTER 393 | Frozen Thawed Difference oO +e ed | eae) er een Cornus stolonifera............ | 2.22mm 2,20mm 9.02mm | contraction a“ SO veg fal NS Chote l= 2860 2.61 0.05 a Tilia platyphyllos.......... 3.63 3.54 0.09 i Pim. Tres dais okie *.i5 3.07 0.08 S Populus dilatata.... «20.66 ..04. | 2.97 ey | ©.00 ° PS eet eT roe fae 2.03 3.04 0,01 expansion Acer platanoides............. 5.06 5.08 0.02 - iy cody emai S ee 4-34 4-34 0.00 s “s pa rete Pe 4.16 4-17 0.01 = ie a | ea ae as 4-92 4.88 0.04 contraction “ee se ae is Bo eRe Pee het ag eee 4.04 4-03 0.01 - eee ee 4.92 4.68 0.04 : Prunus persica............... 2.62 2.59 0.03 ' * Sige Te Te 2.50 2.46 0.04 . ‘a POD alte Sete ies 2.72 2. 66 0.06 . Prunus americana............ 2.17 2.09 0.08 s Meee ree ee 1.78 Ei71 0.07 ie ve fe he ec or ae 2.10 2.06 0.04 “ the wood is eliminated, of course, and that the formation of ice tends to bow out the scales so that they stand less closely together. If the bud scales curve like leaves in freezing this result is to be expected. When the temperature rises sufficiently, the buds and twigs thaw out and regain their normal condition. In the sections under the microscope the reabsorption was so rapid as in most cases to be entirely completed when the ice itself had finished thawing. This results in an active sponging-out movement in the sections as the cells recover from their collapsed condition (fig. 6). On account of the rapidity it is frequently difficult to keep the point of observa- tion in the field of the microscope. Thawing seems not to harm these tissues in the least, no matter how frequently or how abruptly it is done. I have often tried the experiment of transporting twigs abruptly from —18° C. to the warm laboratory at 21° C. and back again several times, thus alternately thawing and freezing them. No matter how many times this was repeated no injury could be detected in the buds, even when subsequently placed in the green- house to grow. Buds and twigs do not thaw at o° C. if the rise in the surrounding temperature is gradual, as it is in atmospheric changes, but at a much lower degree. The thawing like the freezing is proportional to the temperature, and is almost if not quite completed when the 394 BOTANICAL GAZETTE [JUNE freezing point of the tissue is reached. This, in the case of buds; lies at about —3.5° C. to —2.3°C. Hence, if tissues which have been subjected to—18° temperature in the open are to be observed with the maximum ice content, it must be while the temperature is still low. If in the morning the temperature has risen to —7° C. before observations are made, very little more ice will be found than if the cooling to —7° C. had just taken place.'3 6.—Syringa vulgaris: same section as in fig. 3 thawed in the laboratory; note sponging out of tissue and closing of spaces occupied by the ice. 13 GOEPPERT gives a similar experiment. “Twigs with buds of Cornus mas- cula, Prunus Cerasus, and Aesculus Hippocastanum were on January 2, 1871, placed ten hours at a temperature of —16 to —20° C. Then while frozen stiff they were eri into the warm tube of an oven at 25° C. and placed in water for further obser- tion. They waht a later just as others that had not been subjected to this riment.” Some other experiments with herbaceous plants led GOEPPERT to iain that in most cases alternate thawing and freezing, when taking place many times, gradually weakened the tissue. Ueber das tseFeleie Erfrieren der Pflan- zen und Schutzmittel dagegen. Stuttgart, 1883, pe 33. Warttrn believes that rapid thawing gat freezing is very detrimental to the a eT 1906] WIEGAND—BUDS AND TWIGS IN WINTER 305 WINTER FUNCTION OF BUD SCALES. Bud scales are obviously for the purpose of protecting the tender inner shoot from detrimental external influences; but how is this protection accomplished? This is a subject regarding which opinion has varied widely and does still at the present time. I believe we shall find that the most widely accepted views, strangely enough, are not the correct ones, even though the subject appears so simple. We can conceive of such protection taking place along four lines: (1) by keeping out external moisture; (2) by preventing the penetration of cold or sudden changes of temperature; (3) by preventing the escape of internal moisture; (4) by warding off external mechanical injury. It seems best to discuss each of these in turn, and in this way determine the extent to which cach one is operative. External moisture. There is a widespread belief that bud-scales function by keeping out the wet. The subject, however, is a difficult one to determine experimentally, and I can find no reference in literature to such experiments having been performed. Let us first determine the possible ways in which water might be supposed to injure the embry- onic tissues. First, the cells might absorb too much water and thus become more sensitive to frost. It seems quite reasonable to believe that a cold spell following such an event might «nd the life of the bud completely. Again, through gradual absorption of the air by the water the latter might replace the air in the intercellular spaces, thus preventng free respiration. Or again, if a thawing bud were surrounded by water, the latter, instead of.air, would be drawn in to fill the vacant intercellular spaces, the final result being the same as in the last case. Lastly, one might expect that the freezing of free water between the embryonic foliar and floral parts might cause mechanical injury. purple buds and twigs of peach. Green twigs and especially whitened ones warm up less each day and this color would therefore be protective. I believe it may quite likely be true that delicate buds might suffer by such violent treatment either from stimulated activity or increased transpiration, even though hardy trees are apparently indifferent. Das Verhiltniss der Farbe zur Tétung von Pfirsichknospen durch Winterfrost. Inaug. Diss. Halle. 1902. 396 BOTANICAL GAZETTE [JUNE Taking up these in turn, if the cells were so unprotected as to be capable of absorbing water in this way, they would be expected to lose a large part again when dry conditions returned, and thus quickly following frosts alone could do harm. ‘There is also con- siderable doubt whether sufficient water would be absorbed by the cells to cause any perceptible difference in sensitiveness. Water at winter temperatures absorbs air very little, and especially after having fallen in the form of raindrops it may be considered as nearly saturated. The air in the leaves would probably be absorbed very little, if at all, although compression of the air due to capillarity might allow some water to enter. If the thawing tissue has its spaces filled with water instead of air, this will not necessarily cause harm. In experiments on leaves it was found that only the ivy leaf was unable to recover when the spaces were filled with water. Many leaves allow the water to evaporate and then become normal. Mechanical injury is not probable since the air spaces of the tissue would be elastic enough to overcome the compression of the expand- ing ice between the organs, and after the tissue froze slight pressure from the outside on the compressed cells would do no more harm than the pressure of the ice masses within ordinary tissuc. However, the greatest objection to this theory, it seems to me, lies in the fact that protection against moisture might be obtained in a much simpler manner. The embryonic tissue might be densely clothed with strigose hairs, or densely glaucous, either of which would cause the rain drops to roll off without wetting, at the same time allowing gas-interchange to continue; or a coating of resin would effectually prevent all danger of water absorption. All of these devices are more simple than the elaborate system of bud- scales found on many trees. On the other hand, the wool produced on many buds would tend directly to retard the drying of the bud surface. The result of an experiment may here be given. During the winter of 1902, about January 24, several buds of Acer platanoides, deprived of their scales but still remaining on the tree, were each inserted in a rubber pipette-bulb previously filled with water. The neck of the bulb was then fastened firmly around the twig by means of twine. The experiment was allowed to continue about one ee ee Oe et ee ee se hee eae a Serge A) 1906] WIEGAND—BUDS AND TWIGS IN WINTER 397 week, during which time temperatures of —23.5° C. had alternated with those of 4.3° C., so that the buds were alternately frozen and thawed. After removal of the rubber, the tissue appeared as fresh and sound as ever; the twigs were then cut and placed with their ends in water in the greenhouse, where the treated bud remained fresh as long as did others whose scales were freshly removed as check experiments.'4 There exists, it seems to me, insufficient evidence to sustain the theory that the exclusion of external moisture has played an impcr- tant part in the evolution of scaly buds. Heat conduction. The popular belief is widespread that bud-scales serve to keep out the cold, and indeed such an explanation appears in some of our leading textbooks and in various other works. A moment’s consideration will convince us that this cannot be true. No plant tissue yet known is a perfect non-conductor of heat, or, indeed, less than a fairly poor conductor, and scale tissue is no exception; while the very thin nature of the scaly covering on some buds, as those of Salix, would absolutely preclude their offering more than a moderate amount of resistance to the escape of heat. To keep out the cold during an entire cold spell in winter would require, even in much thicker tissue, an almost absolute non-conductivity, and that is possessed by few if any substances in nature, much less by the bud- scales. This erroneous impression has arisen probably through comparing the action of bud-scales with that of clothing upon the human body, forgetting the fact that in the body there is a constant source of heat without which clothing could not keep it warm for more than a few minutes.'S ™ Kny found that with the bud-scales and cortex intact average twigs will not take up as much water through these organs as they give out in dry air pie ae a similar time. He neglected, however, to experiment with naked buds. Ue die Aufnahme tropfbar-fliissigen Wassers durch winterlichentlaubte Zweige von Holzgewachsen. Ber. Deutsch. Bot. Gesell. 13:361. 1895. 8 It may be suggested that such a constant source of heat does actually exist in a tree, at least so far as the buds are concerned, and that this is provided by the heat accompanying respiration. However, reference to any textbook in plant physi- ology will show that the amount of heat evolved in this way is but slight in the very best examples, which are all herbs, and is mainly evident during the period of most 398 BOTANICAL GAZETTE [JUNE Such substances can only retard, not prevent, the escape of heat. As a final argument we may return to the fact that observation shows that buds are always filled with ice during cold periods, which of course could not occur if they were kept warm. It is a more difficult matter to demonstrate whether the non- conductivity of the bud-scales is of importance to the bud in any other way. Recently Griiss'® has quite firmly upheld the theory that one of their chief functions is to modify the temperatures reach- ing the interior of the bud. We can conceive of several ways in which such protective service might occur. First, poor conduc- tivity might prevent injury from too rapid thawing. Second, bud- scales might prevent extreme fall of temperature by preventing excessive radiation. Third, they might prevent too frequent rapid thawing and freezing due to fluctuating sunlight, and thus prevent excessive water evaporation. Before answering any of these questions let us try to understand a little more fully the actual relation of bud-scales to heat. This problem resolves itself into two parts, namely, a consideration of the conductivity simply, and a consideration of the relation to normal atmospheric heat changes in the open. On the question of conductivity the following experiments seem to throw some light: Two thermometers, previously tested as to their readings, were selected, and the bulb of one was covered with the imbricated scales of a fresh horsechestnut bud, as in the previous experiment to determine the effect of color, thus forming an arti- ficial bud with the thermometer bulb in place of the young shoot. The other bulb was left naked. The experiments were all conducted within the building where the conditions were more constant and presented fewer uncontrollable factors than outside. The room rapid growth. During the dormant winter period it must be very slight in all trees. An ordinary thermometer probably could not measure it. It may also be suggested that since the large size and mass of the trunk would retard heat changes, being warmer than the air meee the temperature is falling, and cooler when the latter is this, by conduction along the gp hae modify the temperatures in the shoots and buds. UIRES has shown (M Bot. Stud. 1:453) that the average empertr in a box elder tree was in ace 1.3°C. higher than the air, in Fe ry the same as the air, and in March 1° lower. The differences between internal ‘esd i deena temperatures during the day was in all cases only a few degrees. he idea that the branches can conduct such slight modifications so long a distance without loss is so evidently unreasonable as to require no more discussion here. 16 Griiss, J., Beitrage zur Biologie der Knospen. Jahrb. Wiss. Bot. 23:651. 1892. AL see ee 1906] WIEGAND—BUDS AND TWIGS IN WINTER 399 selected had a temperature ranging from 3.7° C. to 4.3° C. during the several days on which readings were taken. The two ther- mometers were brought to the same reading in a warmer place, either in another room or over a water bath, then quickly taken out and the readings recorded for every few seconds until they again registered at the same degree in the cold atmosphere of the room. Two classes of readings were taken, one from a temperature only a few degrees above that of the cold room and the other from one far higher. The readings in each class, taken with the same bud, corresponded remarkably. A specimen reading from each set is here given. TABLE II. Horsechestnut bulb and naked bulb transferred abruptly from a temperature of 19.5° C. to one of 3.5° C. (See fig. 7.) Naked bulb a ane | ° Difference | Time difference 67°F. 67°F | osec.| 0 F.(0°C.) © sec. 66 67 § + eg 20 65 67 10 o: (ae) 20 64 67 18 gts 37 ; 63 66 | 25 3 rs) 45 (3 min.) 62 65 | 32 cee ke, _ 6x 65 | 38 4 ap a 72 60 65 45 5 23) 60 Weegee gees 5 63. . 3.0 : §7 Ps : 70 3 ? ba) 100 (1% min.) 62 80 6 3.3) 105 55 62 go 7 i) 125 54 62 100 9 30) 145 52 61 110 9 Ke) a 9° 130 .O ° 3 ine i .o) 200 (34 min.) 47 6 185 9 so.) 255 46 6 200 10 ee) 285 45 5 215 10 a na 4 2 10 a 7 E pe : Be 9 .0 ) 420 (7 min.) 42 I 325 9 re) ed 41 4.4 r 40 8 pe Blea) 765 (12 min.) 49 47 442 z.: 43-0 4} 765 40 46 485 Go t3.4,) 765 3) 45 555 6 (3-3) 960 3) 44 620 5 oe a, 960 ; 38 43 705 et) 1235 (204 min.) 38 42 845 S48 3 38 4? T160 2 1.0 ) 38 3? 1515 t (es) 38 33 1942 ° 0.0 ) 400 BOTANICAL GAZETTE [JUNE aN \ sie oe a Se ean — See eeeeetee tebe sess horsechestnut bulb; ........... naked bulb. Abscissas represent FIG. 7; 5° F.; ordinates, 100 seconds. See Table II. The first column of figures represents the readings in degrees from the thermometer with the naked bulb; the second column the same from the bud-covered instrument; the third column shows the time in seconds from the beginning of the experiment; the fourth column the difference in degrees at each reading; and the fifth column is the ‘“time-difference,” so-called, which represents the number of seconds elapsing after a reading on the naked bulb before the same temperature was reached on the horsechestnut bulb, in other words, the number of seconds by which the bud- scales retarded the fall of temperature in the enclosed bulb. While not attempting to deduce the physical laws governing the fall of temperature in each case, we may note from the tables and curves several points which bear upon our problem. It will be seen that theoretically the time required for the temperature to fall in either case is infinitely long, the curve becoming nearly horizontal towards the end of each experiment. But for all practical pur- poses, and as closely as my instruments would measure, the fall was completed in about thirty minutes in each case. The greater part of it, in fact, was completed in ten minutes. As regards time, in Table II the very much more rapid radiation of heat more than balanced the effect of the greater quantity of heat to be radiated. As we should expect, the retarding effect of the bud-scales in degrees, shown in the fourth column, was much greater in case of the greater extremes of temperature, and was greatest when the papas enh yer pn m2) 1906] WIEGAND—BUDS AND TWIGS IN WINTER 4o1 / TABLE III. The same thermometers and bud preparation transferred abruptly from a temperature f 51° C. (over a water bath) to one of 2.7° C. (See fig. 8 Naked bulb | Horsechestnut Time °Difference Time difference 124° F 124°F. o sec riot © sec. sai 123 5 a ks 15 116 122 10 6 (4.2 * hee 120 15 1o (5.5 27 106 118 20 re ne 28 100 116 25 16 (8.8 42 97 114 30 my 184 45 (2 min.) 94 113 35 19 6 (10.5 50 oI 122 40 eo” (15,3 Be 88 109 4s ar (11.6 60 84 106 48 22 12.2 97.7 82 105 sy ag (12.7 80 80 103 65 230 «(12.7) 80 T7 Iol 65 24 144 go (14 min.) 74 99 69 25 (13.9 III 73 97 75 25 (13-9 115 7° 96 80 20> (1320 120 94 85 25 (14.4 125 67 93 go 26 (13.9 135 65 gr 95 26 (14.4 145 63 88 105 25 330 155 j 61 87 115 26 14.4 160 ' 60 86 120 26 14.4) 165 58 83 130 #e-\- 08.6 180 58 81 140 23. {a2.9) 180 54 76 160 22 12.2 200 (3} min.) 52 75 170 23 42.4) 230 50 74 180 24 13.3 250 49 72 Igo 23 12.7) 265 47 69 210 22 12.2) 280 46 68 220 22 12.2) 330 45 65 240 20 ae A 335 44 63 260 19 10.5 240 . 42 61 275 19 (10.5) 435 (74 min.) 41 59 305 18 10.0) 465 41 57 320 16 8.8) 465 = 56 335 15 8.3). 465 ae 55 355 15 (8.3 545 (9 min.) 4o 80 13 7.2 545 39 aa 12 (6.6 580 39 49 455 10 5-5 580 : 38 47 4 g (5.0 ggo (164 min.) 38 46 55° 8 (4.4) 5 37 44 600 7 O20) 1300 (214 min.) 37 43 O..4s3 37 42 710 ee 9 37 41 770 aes fae | 37 40 Mee ee 6. 37 39 I Pais Ge A 37 38 1480 Beg ae 37 37 1900 yore toe ©, Ra al a AO eh ore Rr ane 80 BOTANICAL GAZEFTE [JUNE fall was most rapid. Of much more importance to our problem is the retarding effect in point of time, shown in the fifth column. This increased very rapidly towards the close of the readings, but was for our purpose practically the same in both cases. It was greater in proportion to the slowness of heat penctration, and was also somewhat greater at first in Table I than in Table II. The greatest retardation capable of measurement with the thermometers used was about twenty minutes, while for most of the experiment it was only from one to nine minutes. It was found that decreasing the thickness of the scaly covering decreased this time difference very markedly; while the presence of air Oe i between the scales tended to make it greater. The mass of the thermometer bulb, or of a shoot in a normal bud, and the extent of the ai radiating surface, are important factors in deter- a mining the length of time required for such a structure to cool. While the mass of the mercury N in this case is much greater than that of ae \ a _- NJ a2 c. 3 S688 © Sa 2s 8 2 SS s ce = horsechestnut bulb; _........... naked bulb. Abscissas represent Fic. 8. 5° F.; ordinates, 100 seconds. See Table III. the shoot, its specific heat being only one-thirtieth that of water would render the two not very dissimilar, so far as the present problem is concerned. In apparent volume they do not differ greatly, so that the radiation surfaces of the two would be nearly the same. I believe we are justified in saying that a normal horsechestnut bud would not behave in any essential way differently from the artificial one here used; and that the time for it to cool off would — 1906] WIEGAND—BUDS AND TWIGS IN WINTER fe) 403 be for all practical purposes not over about thirty minutes, no mat- ter whether it was cooled very much or only a little, providing it was plunged directly into the cooler temperature. We may also say, I belicve, that smaller buds with thinner scales and smaller shoots will show a time period correspondingly less than thirty minutes, and a time difference which will approach more nearly zero. In the case of the willow buds with only one thin bud-scale, the time period and time difference must be very small indeed. A number of readings were taken in which the thermometers were warmed up instead of cooled, and it was found, as expected, that the above generalizations applied in this case also. Providing that atmospheric changes out of doors are abrupt, I fail to see how the temperature at the center of a bud of medium size can be retarded more than five or ten minutes over practically all of the range of fall. A small bud would probably be retarded only about one to five minutes. Of course the retarding would be greater than this through the last degree and fraction of a degree, but this slight change, it seems to me, would be of little moment to the present question. Buds in nature, however, are under slightly different conditions. Instead of being transported from one temperature to another, the temperature itself changes. We should therefore conduct some experiments in which the air itself is varied. This change is either very gradual, as when a thaw approaches, or more abrupt, as when the sun shines from behind a cloud upon the bulb, which is the only way in which abrupt changes are produced in nature. In either case they are much less violent than were our laboratory experiments. During warming by the sun, radiation from surrounding objects may play an important part and introduce still another factor. We should thcrefore conduct some experiments in which the air itself is warmed. The experiments with the horsechestnut bud already described in the discussion of the function of color are to the point here. They show in a surprising way that instead of retarding the rise in temperature within the bud, under these very natural conditions the bud-scales actually scem to hasten it. These experiments were with direct sunlight. It seemed possible 404 BOTANICAL GAZETTE [JUNE that the readings might be different if radiated instead of direct heat was employed, especially since there is a considerable difference in the nature of such heat, as shown by the well-known fact that direct heat from the sun passes easily through glass into the green- house, but when radiated passes out with much greater difficulty, thereby warming the house. A number of readings taken with naked and_ horsechestnut bulbs transferred from the shade to the surface of a black book in direct sunlight with the bulb raised 3-4™™, or with the bulbs pro- jecting several inches over the edge of the book which itself was raised several feet from the ground, or with the bulbs raised 7.5—10°™ above the surface of the book, showed no appreciable difference that could be referred to a difference in kind of radiated heat. There was some difference in the readings, of course, but this could ke traced directly to the fact that there was more intense heat where the heat of radiation was also present. In case of more intense heat the extra absorbing power of the bud-scales was at first more obscured by the slightly greater retardation of heat-penetration due to the greater difference in outside and inside temperatures, as we should expect from the deductions from Tables III and IV. This was partially shown by the difference in locality of the crossing of the two curves plotted from each reading. Looking at the matter from still another standpoint, we may consider how much time is required for a bud to thaw. As shown by the cover glass measurer, the wrinkled willow twigs thawed and became perfectly normal in thirty minutes at the temperature of the laboratory. Undoubtedly the ice had disappeared in about half the time. Large buds of horsechestnut will lose all their ice in about twenty-five minutes under similar conditions, and buds of Negundo in about fifteen minutes. The small buds of the black cherry require only about ten minutes for thawing. The time required in the laboratory for the various buds, therefore, is ten to thirty minutes. The question is whether when the temperature changes are slow the buds thaw proportionately more slowly. The answer must be that they will, slightly, just as a cake of ice will thaw more slowly when the temperature rises gradually than when the rise is abrupt. This difference is proportional to the size of the 1906] WIEGAND—BUDS AND TWIGS IN WINTER 405 ice cake, since it depends largely upon the non-conductivity of the ice and the greater quantity of heat required to convert ice into water. This heat is more slowly available when the change is gradual. Although no experiments were made under these condi- tions, it is to be expected, I think, that with long slow rise in atmos- pheric temperature, the retarding effect would almost if not quite disappear. Frozen peach buds, placed in the air at—5.5° C., which gradually rose in 2 to 2.5 hours to a temperature of —1.0° C., were completely thawed, apparently as soon at the temperature reached about —2.3° C., thus following the general rule for frozen tissue. We are now in position to consider the questions outlined on a previous page regarding the various ways in which the bud-scales May be supposed to act beneficially by modifying the temperature. It was first suggested that they might retard the thawing out and thereby be of benefit to the bud. From the tables already given and the observations regarding them, it becomes at once apparent that the temperature modification which scales are capable of producing are, in the cases of moderate sized buds, of very little Moment—not more than two or three minutes during most of the time, and then only when the change from one temperature to another is abrupt. When the transition is gradual, the retarding effect will be very slight indeed, and is frequently wholly offset by the absorbing power of the darker color. I cannot see how under any atmospheric condition the modifications can become great enough to be noticeable unless careful measurements are taken. The idea that a slow thawing is beneficial to plants has come about from analogy with the frosting of human tissue and from the con- sideration of the treatment which gardeners successfully give frosted plants. But the gardener’s treatment consists in keeping the plant cool and dark for hours or even days after the freezing; while recent investigators have shown that slow or rapid thawing (7. e. conversion of the ice into water) in themselves bear no relation Whatever to the extent of the injury. The gardener’s treatment is essentially an after-treatment—while the plants are recovering from the shock. I have already cited the fact that buds of many trees, at least, may be thawed in an oven and then frozen alter- nately many times and still come out in the greenhouse apparently 406 BOTANICAL GAZETTE [JUNE as fresh as others not so treated.'7. The answer to this first question then, is, that bud-scales do not function by preventing rapid thawing of winter buds; neither does bark so function towards the twigs. It has been suggested that bud-scales protect the bud by pre- venting rapid radiation from the delicate tissue during the cold nights, and thereby preventing a harmfully low fall of temperature. MU ier-TuHurGAu,'® by placing one thermometer on some cotton under a o.5°!" cloth screen fastened 1°" above the ground, and another thermometer outside, was able on a clear night to get 4° C. difference due to radiation. Griiss'® states that differences in temperature due to radiation may be one or two degrees on cool nights just before sunrise, and as great a difference as 6° C. has been observed by other investigators. A difference of 4-6° C. would frequently be of importance to tender exotic buds in winter, but it is scarcely to be supposed that so slight a difference would be of much moment to the great majority of perfectly hardy species which withstand all of the fluctuations of our vigorous American climate without injury. Indeed these species seem capable of existing below any atmospheric temperature that has yet occurred in this country, as freezing mixture experiments have shown. Besides, the structure of buds does not lead one to expect a radiation screen as efficient as those specially constructed. Strictly speaking, the ques- tion here is not one of radiation of heat, since the scales are all more or less in contact, but of conduction, and as such has already been treated. HENsLow’? has shown that it scems desirable for plants in tem- perate regions to protect their delicate bud-structures from loss of water when the bud is opening. Such loss he says is favored by radiation and heat absorption. The above objection will apply here also for the first part of this last statement, and the latter part is treated elsewhere in this paper. 17 Motiscu, H., Untersuchungen iiber das Erfrieren der Pflanzen. Jena. 1897 18 MULLER-THuRGAU, H., Ueber das Gefrieren und Erfrieren der Pflanzen- Landw. Jahrb. 15:563. 1886. 9 Gruss, Beitrige zur Biologie der Knospe. Pringsh. Jahrb. 23:651. 1891-92. 20 HENSLOW, G., On vernation and the method of meee of foliage as protective against radiation. Jour. Linn. Soc. Bot. 21:624. 1886. lt 1906] WIEGAND—BUDS AND TWIGS IN WINTER 407 In December 1901 some experiments were conducted to show whether twigs and buds while continuing frozen lost as much water by evaporation as when alternately thawed and frozen several times during the same period. It was found that they did not quite, and hence the question whether bud-scales may function by pre- venting too frequent thawing and freezing. Several buds of Pinus Laricio and horsechestnut, also several twigs 15°™ long of Syringa vulgaris and apple, were sealed at the cut end with Venice turpen- tine, weighed, and quickly placed on a tray in the open air. They were divided into two equal lots, and one of these was brought into the warmer laboratory for a few moments ten times, thus insur- ing ten alternate thawings and freezings. During the experiment, which lasted three days, the temperature ranged from —18° C. to —7° C. in the open. The results were as follows: Horsechestnut buds continued frozen lost 0.4% of their water. «alternately thawed and frozen lost 0.6% of their water. i he oe — continued frozen lost 3.4% of their water. alternately thawed and frozen lost 5.0% of their water. ea twigs continued frozen lost 1.3% of their water. *¢ alternately thawed and frozen lost 2.4% of their water. i a — continued frozen lost 1.6% of their water. alternately thawed and frozen lost 2.4% of their water. In every case there was a greater loss of water from the buds which were alternately thawed and frozen. The difference was very marked, and in each case amounted to about one-third of the total loss. Considering the total quantity of water present, this was really a very slight increase in loss, however, being 0.25% in horsechestnut, 1.1% in lilac twigs, 0.8% in apple twigs, and 1.2% in pine buds; and with me it is a serious question whether, in all of these cases, so slight a difference would not be quickly equal- ized during spells of thawing by conduction from the older wood, if the twigs and buds were connected with the trunk in the normal manner. Again, the thawings in nature would probably be fewer, and it has not been shown that bud-scales prevent such thawings. It seems to me that here again a beneficial functioning of the bud- scales is very doubtful. But the most vital argument against all these cases lies in the fact that experiments have shown that dark buds tend actually to 408 BOTANICAL GAZETTE [JUNE increase the heat absorption. Therefore, these considerations could scarcely have been instrumental in bringing about the existence of such structures. The idea that bud-scales may protect the bud by warding off the hot rays of the sun applies mainly to the tropics. It seems to have been first advanced by TrEvuB,?" who cites several cases, where in plants exposed to the hot tropical sun delicate young tissues were inclosed in enlarged stipular organs or else well-shaded by overlapping leaves or by other special structural provisions. On the same subject, in 1891 another paper was published by Porrer.?? According to this investigator many trees in the tropics protect their young leaves and shoots from direct sunlight by means of stipules. These organs were removed from a number of buds and in every case the leaves from these when mature were deformed and abnormal. The sunlight seemed to produce injury by causing more water to be evaporated than could be replaced. For this reason Artocarpus, the most pronounced type of this class, unlike most trees, produced leaves throughout the dry season, probably because of the stipular protection. Instead of by stipules some tropical plants obtain similar protection by various methods of leaf-folding, shading by older leaves, and coating with gum. Is there not inaccuracy here in his interpretation? Rather than by actually preventing the entrance of heat from the sun, which it seems such structures could do only to a slight extent, is it not more proba- ble that they function simply by preventing the escape of extra moisture vaporized by the intense heat ? The relation of bud-scales to the young shoot when the bud is opening is discussed under internal moisture relations. Suffice it to say that the results reached seem to indicate that even in this case the scales do not function beneficially by modifying the heat. It has sometimes been thought that the layers of hair and wool found in many buds, as for example in the horsechestnut, are for the purpose of modifying the heat conditions inside. To obtain 21 TrEuB, M., Iets over knopbedekking in de tropen. Hand. van. het eerste Nederlandsch Natuur- en Geneeskundig Congres. Amsterdam. 1887, p. 139- Ref. Bot. Centralb. 35:328. 1888. 22 PoTTER, M. C., Observations on the protection of buds in the tropics. Jour. Linn. Soc. Bot. 28: 343. 1891. Oa eal 1906] WIEGAND—BUDS ‘AND TWIGS IN WINTER 429 evidence upon this point I performed the following experiment. The two thermometers used in the previous experiments were selected, and the bulb of one was coated with black cloth; that of the other was wrapped in a layer of cotton about twice the thickness of the wool in the horsechestnut bud, and was then coated with black cloth. The surface of both bulbs was therefore black. TABLE IV. Bulb covered with black cloth, and bulb covered with cotton and black cloth; trans- erred from a temperature of 56° C. to a room of 9° C. Cloth bulb Cotton bulb Time ° Difference Time difference 134-8. 134° F. o sec. o° F. (0° C.) © sec. 130 131 5 I (0.5) 7 128 131 to r Seda SS 10 125 128 15 3 (1.: 15 123 128 20 5 (2:7) 15 222 127 25 5 (2: 15 119 125 30 6 3.3) 20 i17 123 35 6 2-3) 20 114 T21 45 7 3-9) 20 ach a T19 50 8 4-4 25 109 117 55 8 4-4 26 106 Hie 9 (5.0) 30 104 112 70 8 4.4 30 100 110 80 Io oy) 37 98 107 85 9 -0 40 96 105 9 9 iO 40 93 103 105 Io 5 40 QI IOI 113 Io us 42 99 120 to fe 45 87 97 130 10 5 45 86 96 135 Io as 43 oy 93 145 9 (5-0) 59 82 ot 155 9 (5.0 55 80 89 165 9 (5.0) 55 78 88 173 10 8 _ 62 77 i 86 178 9 0 7? 76 85 185 9 -O 75 74 84 195 be) “5 75 73 83 205 Io 5 80 72 82 210 10 “5 85 71 81 215 10 5 95 7° 79 225 9 te) 95 69 78 235 9 .O 100 68 77 250 9 re) 105 67 76 60 9 .0 110 66 74 270 ee) 110 65 73 285 8 (4.4) 120 64 72 295 8 4-4 130 63 71 310 8 4-4 135 63 70 320 7 (3-9 135 62 69 335 7. (9 140 410 BOTANICAL GAZETTE [JUNE TABLE IV.—Continued. Cloth bulb Cotton bulb Time ° Difference Time difference 61°F. 68°F. 355 sec. or (3,90) 155 Bec. 61 68 370 6 aS 155 61 66 380 5 Ce 155 60 65 405 , aia 5 145 9 64 425 5 2a7 160 58 63 445 5 2.7 185 7 65.5 475 5-5 (2.7 210 56 61 510 5 2.7) 270 6 60 550 4 2.1 310 5 59 585 4 2.1) 545 4 8 630 4 2.1) 545 4 57 685 3 1.5) 545 3 7 745 4 2.1) 1083 3 6 780 a £25 1083 3 5 895 2 eae. 1083 2 55 IOI5 a (i.5 1300 2 54 1175 2 (1.0 1300 I aie 2155 a a | 480 49 -! 2635 2 (126 1440 49 50 3355 EG s§ 48 49 40°75 Se Ne As expected, the retarding effect was apparent in this rather violent experiment, but it was not great. The maximum degree difference of 10° was less than one-half that produced by the bud- scales in Table III, while the time difference through the greater part of the experiment ranged from o to 4 minutes. I think it may be inferred that the wool in the horsechestnut bud retards the pene- tration of heat, when the changes are at all great, by 0.5—3 minutes. At any rate it seems evident to me that the retarding power of the wool in such buds as horsechestnut is insufficient to explain the presence of such a structure. This appears not only from experi- ment but from a general consideration of the thinness of such struc- tures compared with the relatively great temperature differences which they are supposed to offer protection against, and must in order to be effective. Their true function, it seems to me, lies in an entirely different direction, as we shall see somewhat later. In concluding this study of the relation of bud-scales to temper- ature the following summary may be made: : Bud-scales or bark cannot “keep out cold” during the cold spells of winter. ; They seem not to modify the temperature sufficiently to be of 1 hive apy age 1906] WIEGAND—BUDS AND TWIGS IN WINTER 4II beneficial importance in preventing rapid changes, even if such changes are detrimental. Rapid thawing in itself is probably not detrimental to buds and bark. Bud-scales seem of no benefit in keeping out the heat from sudden bursts of sunshine. They do not appreciably prevent the loss of water by preventing alternate thawing and freezing. They do not retard radiation to any important degree. Dark-colored bud-scales indeed, instead of preventing tempera- ture changes actually seem to absorb more heat than if they were lighter colored. “Wool” in buds does not function by modifying temperature changes. Bud-scales do not seem to function in modifying temperature changes when the bud is opening. Bud-scales may protect the delicate tissues in the tropics from heat, but it would seem rather from excessive transpiration due to great heat than from the heat itself. Finally, we may conclude that as a factor in the evolution of buds and bark in cold climates temperature considerations have probably played a very minor part. Internal moisture. Of all the more important factors concerning the function of bud-scales, perhaps that relating to their inhibiting effect upon the loss of internal moisture is the least recognized by people in general. In scientific literature, however, it has received considerable atten- tion. Most authors now consider this one of the principal functions. of the bud-scales and also of the bark. The subject has been dis- cussed briefly by CapuRrA*3 and Groom,’* but also more fully by Gritiss,?5 who performed a number of experiments to demonstrate the point. His results may be summarized as follows. The first function of the scales consists in protecting the inner meristematic 23 CaDURA, R., Physiologische Anatomie der Knospendecken dicotyler Laub- baume. Breslau. pp. 42. 1887. Pe Groom, P., Bud protection in dicotyledons. Trans. Linn. Soc. II. 3:255. 25 Grtss, J., Beitrige zur Biologie der Knospen. Jahrb. Wiss. Bot. 23: 649. 1892. 412 BOTANICAL GAZETTE [JUNE tissue from loss of water. Even in summer and especially in fall, when the sap flow decreases, the tender embryonic interior of the bud must be protected from too great transpiration. Also in winter this function is not interrupted, for then the cold wind can bring into play its desiccating action. To prevent loss of water, cork layers are formed, or in place of these felty hairs may be produced. A third method. consists in the excretion of resin. If, under con- stant temperature, the scales were removed from an oak bud, it soon died, even though there was a moderate amount of moisture present. The inner bud-scales dried out and perished, as well as the embryonic tissues. The young leaves of a beech bud so deprived of scales persisted much longer than did those of the European oak; which he thinks was because the former were hairy while the latter were not. Buds of horsechestnut proceeded to develop in spite of the removal of the scales, probably, he thinks, because of the thick wool among the young parts. Buds of Abies pinsapo, whose pitch had been removed by carbon bisulfid, dried out in a very short time. These experiments were all performed on twigs cut from the trees and placed in water. In 1895 Kny?° published a paper dealing with the transpira- tion and absorption of water by buds and twigs in winter. He cites WIESNER and PACHER as having shown that horsechestnut loses water from twigs in winter, and also Hartic as having shown that many trees do the same. Experiments are given to show that in general not so much water is absorbed by these parts in saturated atmosphere as may be given off at an ordinary degree of saturation. In 1895 some determinations of the amount of water lost by twigs with buds attached were made by the Cornell Experiment Station.?”7_ The experiments lasted three days, beginning April 7. The twigs were sealed at the cut end and kept in an open shed. The percentage of loss ranged from 2 to 10%, with an average of 5-4%- In 1875 WIESNER and PacHER?® found that twigs of horsechest- 26 Kny, L., Ueber die Aufnahme tropfbarfliissigen Wassers durch winterlich- entlaubte Cutis von Holzgewachsen. Ber. Deutsch. Bot. Gesell. 13:361- 1895- 27 BaILEy, L. H., Cornell University Experiment Station Rep. 1896: 4. 28 WIESNER u. PacHER, Ueber die Transpiration entlaubter Zweige und des Stammes der ete ge Bot. Zeitschr. No. 5. p. 9. 1875. RE i ee Se ee are ot ae al ye 1906] WIEGAND—BUDS AND TWIGS IN WINTER 413 nut transpired an appreciable amount in winter at a temperature of 13-17° C. and a slight amount also at —10° C. This was true in older twigs. The leaf scars transpired more than the periderm. The winter buds also lost some water. That there is actual loss of water in winter probably every one knows. My experiments given below show this very definitely, but perhaps few understand that there can be a loss when the tissue is frozen as well as when thawed, though less in extent. Water may evaporate to a large extent from ice crystals themselves, as is shown by the drying of frozen soil, damp clothing, and the frequent disappearance of small quantities of snow at temperature below the freezing point. In buds not all of the water becomes ice, and the remainder is free to evaporate as at a higher temperature. The fall of temperature on the approach of winter is always accompanied by a decrease in the power of root absorption, and it has been shown that, to a certain extent absorption is propor- tional to the temperature. In the case of our native plants, the decrease must be very considerable when the zero air temperatures have chilled the soil to a depth of many feet. A compensating decrease in transpiration must occur or otherwise the cells will suffer from too small water content. This is mainly accomplished by the fall of the leaves, but is greatly aided also by the coverings of the bud and the waterproof bark. But so far bud-scales would not be a necessity, because very little root absorption would probably be sufficient to supply the slight amount of water that could evapo- rate from unprotected buds, compared with that necessary to supply the leaves. Besides, it has been found that considerable water is present in the wood at all times, and in some trees even a larger amount than in the summer. The necessary factor, I suspect, lies in the decreased osmotic activity and vigor of the young tissue itself, During the summer the tendency to transpire is probably, a large share of the time, not so great as in the winter and spring because of the greater saturation of the air; but there are times during the summer when the transpiration is very great indeed. The young tissues do not then dry up very readily, so that little harm usually results. At this time I imagine the growing cells are osmotically very active and more easily draw to themselves 414 BOTANICAL GAZETTE [JUNE a supply of water sufficient to offset that lost in transpiration. In winter, however, the cells are inactive, and on account of the cold the osmotic force is much decreased, so that the cells find it impos- sible to resupply quickly the transpiration water when this function is very great. That loss of water beyond a certain point is detrimental to the cell needs no further demonstration. It has been shown that each cell demands a certain percentage of water, depending mainly upon its activity and water content, in order to maintain its life-prop- erties. If transpiration even for a short time reduces the water in the cells of the bud below the critical percentage, the cells will cease to remain alive. During January 1gor, I cut some twigs of horsechestnut, stripped off the bud-scales from some of the buds, and exposed the whole to an outside temperature of —18° C. to —12.3° C. for 24 hours, after which the twigs were placed with their cut ends in water in the greenhouse for further development. The buds all lived, although those without bud-scales were the first to commence growth. Sub- sequent experiments show that the reason why none died was because the exposure to the dry air was not long enough. On March 1 of the same year, buds of the black cherry, Crataegus punctata, horsechestnut, lilac, apple, and Pinus Laricio while still on the tree were deprived of their scales and each divided into two lots. One lot was left naked, the other was varnished completely with Venice turpentine to prevent loss of water. When the normal buds were opening May 8-10 it was found that both varnished and naked buds were all dead except on Pinus Laricio. On this plant the naked buds were all dead, but the varnished ones were alive, and later all developed into normal shoots. The varnished buds in all cases seemed to be all sound and turgid until warm weather and time for swelling came, when they seemed to decay rapidly, and in no case except the pine did any swelling occur. I suspect that death here was due to the retardation of res- piration owing to the lack of oxygen. The pine is normally closely surrounded by resin without a space inside as in horsechestnut, and possibly some other way is here provided for obtaining Oxy- gen. The pine, therefore, is the only one of the series in whic 1906] WIEGAND—BUDS AND TWIGS IN WINTER 415 the results of varnishing are important to us here. The naked buds of pine in every case began to dry and shrivel up after only a few days’ exposure, and were quite dead long before the time for them toopen. There seems no doubt whatever that the varnish preserved the pine buds by preventing loss of water. Without the varnish the pine was one of the first to succumb. That this thin layer of varnish replaced effectively the thick layer of scales is also good evi- dence toward the idea that the scales do not function by causing temperature modifications. To determine just how much more water is lost from buds without scales the following experiment was performed. Several buds of Pinus Laricio and horsechestnut were separated from the trees by an incision at the base of the bud and the scales were removed from all. One-half were quickly varnished, weighed, and placed in the open air at —18° C., while the other half without varnish were weighed and exposed at once. Care was taken to seal up the cut end in both sets so that no water could escape that would not if the buds had remained on the trees. After three days at a temper- ature of —18° C. to —7° C. the results were as follows: Orig. aa Final weight| Dry weight | Per cent. HO Pinus Laricio—continued frozen: Wath Trid-ecales. oo... caus eck 2.658m | 2.6158™)| 1.358™ 2.7 Without bud-scales.....:......... 1.28 0.96 0.58 45-0 Aesculus hippocastanum—cont. frozen: Ser Btieh ain bet Go oe es Fk 4.03 4.02 1.80 pia Peel aap EAicice alias 18 ic oso oases 2.21 1.94 1.04 33.0 uring experiment by bring- ing into the laboratory: With tead-deales. 35 c5 sis val es 5.30 5.28 2.26 0.65 _ Without bud-scales..............- 2.01 1.66 0.93 32.0 Syringa vulgaris—continued frozen With Dich -atated 5 i405 awe cas i a 1.133 0.52 2.8 Without Sekxeabes Dell Sew ike elars ©.41 0.32 0.18 39.0 I think that nothing could show better than these experiments the very great difference in amount of water transpired by buds protected by scales and those having none. No wonder that the loss of water oversteps the critical point and causes the destruc- tion of the tissues. 416 BOTANICAL GAZETTE (JUNE The buds in the above experiment were separated from the tree upon which they grew, and therefore could not receive water from it to replace that transpired. It would be interesting to know how much water moves into the bud to replace the quantity lost, thus giving a better idea of the actual decrease in percentage within the cells. This has not been done for twigs at temperatures above freezing, but the following figures are available for the frozen buds of pine. I selected six vigorous buds of Pinus Laricio, all on the west side of the tree, deprived them of their bud-scales and allowed them to remain exposed three days. The temperature during this time ranged from —18° C. to —6.7°C., so that the twigs as well as the buds were constantly frozen. Three of the buds were cut off, the cut surface sealed, and placed in a tray at the base of the tree, while the other three remained attached. The results were as follows: baie’ pala Dry weight | Difference | Per cent. ™ wet at end Cut buds— i ©.1058™| 0.0618™| 0.044 41.9 | " 2c Sk aati eg aR PE Tae PP 0.150 0.090 0.060 pies Maas average Ree eed ree os 0.230 0.134 0.096 41.8 Sra Bas coe No Ae Sy eo 255 65139 0.116 45-5 an 0.240 0.121 0.119 49-5 747-5 average Des aie cP ees a es 0.210 0.110 0.100 47.6 It seems, therefore, that there was a rise of about 5% of water into the bud while the tissues were frozen. This is quite possible, since only a portion of the water was converted into ice, the remainder remaining fluid in the walls and protoplasm and still capable of movement. The figures given above for the loss of water from desquamated buds are therefore slightly too large in every case. It may be noticed by computing corresponding figures that the loss of water during this last experiment is slightly less than in the two previous experiments in which desquamated buds of Pinus Laricio were used. This was due to the fact that the last experiment was conducted in a different place, on the other side of the building. ‘I have found that exposure makes a very considerable difference in the loss of water, and readings which are to be compared must ve SE », . — Con a> = = et 1906] WEISS—BARK IN SASSAFRAS 435 from the layer of parenchyma just within the epidermis. He also remarks that the secondary bast fibers, although usually scattered, ‘form distinct strands in certain genera, and that the individual fibers are normally four-sided in cross section with narrow lumina. The present investigation is based on material collected near New Haven, Connecticut, and is confined to the stem and its branches, no reference being made to the bark of the root. The tissues described may be classified as follows: . PRIMARY TISSUES SECONDARY TISSUES Epidermis Tissues derived from the cambium ring Outer cortex The phellogen and its derivatives Primary medullary rays Primary bast PRIMARY TISSUES. Epidermts. The epidermal cells are characterized by a strongly thickened cuticle. Close to the growing point they are isodiametric and thin- walled, but the cuticle begins to make its appearance very early and OADOOSD SSeS POR ESOC e Satya OBE =a cS i> Ba SC <4 s SORTS ORo Ey <> aes Semel scenessios *) ie I Fa hee Pom ; Cre ac hey AN }} hy | iS 2, a Re) (y rai . tf ce Fic. 1.—Cross-section through bark one year old. X70. cam, cambium ring; ¢, epidermis; m, medullary ray; p, parenchyma; phx, primary phloem; s¢s, primar) sclerenchyma; sc,, secondary sclerenchyma; st, stone cells; x, xylem. Practically completes its development during the first year’s growth. At the close of this period it occupies about half the thickness of the epidermis (jig. 1). During the elongation of the stem the epidermal cells retain the power of growth and division. Since their growth is largely in a longitudinal direction, the cell-division is mainly brought about by transverse walls, division by longitudinal walls being much 436 BOTANICAL GAZETTE [JUNE more infrequent. In an epidermis a year old, seen from the surface, the boundaries of the original epidermal cells can usually be distin- » guished. They are somewhat thicker than the secondary transverse oad) < walls, which in turn are thicker than the secondary longitudinal walls (jig. 2). With the formation of es cork the epidermis is of course split longitudinally and soon begins to undergo disorganization. No trace of it is left in a tree 8°™ in diameter. Or The number of stomata produced varies greatly, L000 SEE but seems to be largely dependent upon external conditions. A rapidly growing tree, for example, in 4 a moist locality has many stomata, while a slow- e growing tree in dry soil develops very few. The . YO stomata are depressed and the epidermal cells bound- ing the guard cells are somewhat modified, being longer and narrower than their neighbors (fig. 3)- Fic. 2.—Surface Most of the stomata are transverse to the axis upon view of epidermal which they are borne, a few are oblique, but appar- ae nay Bag ently none of them occupy a longitudinal position. ain ie miei’ ary Lhis is doubtless to be explained by the fact that of the original cell, the stomata are formed late in the development of which has under- the epidermis, the wall separating the guard cells liad i representing one of the secondary transverse divisions of an epidermal cell. In the majority of cases the cells surrounding a stoma contain nie ‘ik anthocyan, so that to the naked eye the stomatal region looks like a minute red speck in the epidermis. This peculiarity _}{ \@/¢ $# affords a ready means for detecting the SS LE stomata. LY aa Epidermal hairs are developed on very ae young twigs before the primary tissues are Fic. 3.—Stoma_ with sur- fully differentiated. They are simple and TU"dins ie nana crcaals unicellular, with thickened walls, and scarcely cage extend below the cuticle (fig. 4). These hairs never persist through the first vegetative period, but dry up and fall away as soon as the cuticle begins to thicken. Their former position is often marked by iG ay 4, x, eS | 1906] WEISS—BARK IN SASSAFRAS 437 small concave depressions in the cuticle. The number of the hairs "varies, and in a general way is inversely proportional to the number of stomata. Thus, in a moist locality few hairs are formed, while in a dry region they are very abundant. In a meso- phytic area some trees bear few hairs, while others under the same conditions bear very many. It would appear from this that the production of hairs was primarily due to individual peculiarities of the tree in question and secondarily to the external conditions under which the tree developed. Outer cortex. The outer cortex comprises everything external to yc, 4-—Epi- the primary sclerenchyma except the epidermis. It is dermal hairs on composed of a ground mass of parenchyma with ia iis scattered stone-cells. No crystal cells occur. With ” the formation of cork the outer cortex gradually becomes disorgan- ized and eventually disappears. . In cross section the parenchyma cells vary from elliptical to rec- tangular in outline, the long diameter running in a tangential direction (jig. 1, p). They vary considerably in size and some of the larger cells have their walls slightly lignified. Most of the cells, however, have thin walls, which may or may not be provided with simple pits. Many of the smaller cells contain starch and this is especially likely to be true of those which border the strands of sclerenchyma. The presence of ethereal oil in the parenchyma can be demonstrated by appropriate tests, but it does not seem to be localized in special cells. In all probability the oil represents an excretory product of the proto- plasm of the parenchyma cells, and this fact would account for its general distribution. The stone cells form a continuous or interrupted layer extending entirely around the stem (fig. 1, st). They sometimes lie next to the epidermis and are sometimes separated from it by one or two layers of parenchyma cells. The stone cells are at first circular in cross section but afterwards become flattened and assume an elliptical outline. In radial section they appear rectangular, being about three times as long as broad. Their walls are strongly thickened by deposits of 438 BOTANICAL GAZETTE [JUNE ligno-cellulose in distinct layers, and these are pierced by numerous simple and branched pits. Primary medullary rays. The primary medullary rays extend from the cambium to the outer cortex, the ray cells merging into the cortical cells without a distinct line of demarcation. The outer portion of the ray is of course directly differentiated from the meristem at the growing point, while the inner portions owe their existence to the activity of the cambium. Some of the cells in the outer portion retain their power of growth and division for several years, the majority of the dividing i any. Hianeteee oa aa vets be Manes lecveltys. eS) (=) ss ig Oa, esa OS ra YS A Bs CY AT El Of { pa NS ae —s oe re = OS | va ie Ps A) war <2 =a 3 S cia, ae" = a we = & >\ ep } Sent Reocmead ies te g ger ey & =a ar he 4) el b} eat * A eo a Se 1 et me ry, Ai i) wa x Fic. 5.—Cross section through old bast. X25. Fic. 6.—Radial section through old bast. X75. c, cork; cam, cambium ring; e, epidermis; m, medullary ray, p, parenchyma; ph, secondary phloem; phel, phelloderm; sc2, secondary sclerenchyma; St, stone cells; x, xyiem. walls being radial. Thus, in a stem one year old, the strands of primary sclerenchyma are separated by from two to five layers of cells, in a stem two years old by as many as fifteen layers, while in a stem four years old the number may be increased to thirty or more. Since the portions of the rays derived from the cambium do not undergo further divisions, they remain permanently from one to three cells in width. In consequence of these facts the rays gradually assume a T-shape a 1906] WEISS—BARK IN SASSAFRAS 439 in cross section. This form is retained until the outer cortex has become disorganized, after which they appear like narrow bands (ig. 5,m). In radial section the rays are from four to fourteen cells across (fig. 6, m). In most of the ray cells the walls are slightly thickened and pro- vided with numerous simple pits. They usually contain starch and sometimes ethereal oil as well. When the cells are cut off by cork the starch disappears, showing that it is completely utilized; the oil, on the other hand, persists. Some of the ray cells between the strands of primary sclerenchyma become strongly sclerotic, and in some cases cells of this character completely bridge the space from one strand to another (fig. 1). They can be easily distinguished from the scle- renchyma cells, even in cross section, by their larger size and distinct lamination. In longitudinal section they appear short and resemble the stone cells of the outer cortex. Primary bast. The primary sclerenchyma occurs in well-defined bundles, averag- ing about fifty fibers apiece (figs. 1, 8, sc,). Most of these bundles, in a radial direction, measure from three to eight cells across. In most of the fibers the wall is so strongly thickened that the cavity is reduced to a mere slit; in some cases, however, the thickening is less and this is especially likely to be true in the middle of a bundle. Apparently the deposition of ligno-cellulose upon the cell walls is not completed until the second vegetative period. The primary phloem lies just within the primary sclerenchyma, between the latter and the secondary sclerenchyma, and forms a band. from three to five cells across in a radial direction (fig. 1, ph,). The Sieve tubes are more or less completely separated from the scleren- chyma by a layer of phloem parenchyma. The cells of this layer tend to be rectangular in cross section, and their slightly thickened walls have numerous simple pits. The sieve plates separating the segments of the sieve tubes are nearly always somewhat oblique 3 they are supplemented by numerous lateral sieve plates, especially in the radial walls of the tubes. All of the sieve plates in the primary phloem soon become covered by deposits of callus. The companion cells conform to the usual type. 440 BOTANICAL GAZETTE [JUNE SECONDARY TISSUES. Tissues derived from the cambium ring. The tissues of the bark, regularly derived from the cambium ring, include the secondary sclerenchyma and the secondary phloem. In addition to these, scattered groups of stone cells, which should prob- ably be considered a part of the phloem, also make their appearance. Of course the cambium also adds new elements to the primary medul- lary rays and brings about the development of the secondary rays (figs. 1, 5). The development of these various secondary tissues begins during the first vegetative period. The fibers of the secondary bast do not form bundles. Some of them form interrupted layers arranged concentrically in the stem, others are scattered through the secondary phloem. The layers ar? usually but a single cell across and are separated from one another by several layers of phloem. The individual fibers are rectangular in cross section and about thirty times as long as broad; their walls are very strongly thickened (jigs. 1, 5, 6, sc,). When the bast fibers are cut off by cork all regularity in their arrangement disappears. The sieve tubes of the secondary phloem, except those earliest formed, are arranged in interrupted, concentric layers, one or two cells across (fig. 5, ph,). Many of the sieve tubes are in direct con- tact with the medullary rays, but very few of them adjoin the scleren- chyma fibers. The tubes exhibit essentially the same structure ‘as those in the primary phloem. On account of their delicate walls they become practically indistinguishable when cut off by cork. The bulk of the secondary phloem is composed of parenchyma. When first differentiated from the cambium the cells of this tissue are closely packed together, rectangular in outline, and destitute of intercellular spaces. As they become pushed outward, their outlines become more rounded and minute intercellular spaces appear. Their walls are fairly thin but are provided with simple pits. Until they are cut off by cork the parenchyma cells are arranged in layers, which lie among the layers of sclerenchyma and sieve tubes. The groups of stone cells are irregularly scattered in the secondary bast but always abut against a medullary ray (fig. 5, sé). Such a group in cross section is often larger than a bundle of primary scle- renchyma and is composed of larger elements. The stone cells are the 1906] WEISS—BARK IN SASSAFRAS 441 most conspicuous structures found in the inner bark, and are even more striking in appearance than those found in the outer cortex. In longitudinal section (fig. 6, st) they show the same outlines as in cross section (jig. 7) and are therefore isodiametric. Their strongly thickened walls show a very distinct lamination and their contracted cavi- ties are connected by numerous simple and branched pits. Prob- ably on account of poor material, these stone cells were not seen by MOLLER. The phellogen and its derivatives. The derivatives cf the phellogen are the lenticels, the cork, and the phelloderm. The lenticel phellogen is the first to make its appearance; the primary cork phellogen is, at least in part, a direct extension of Fis. 7.—Stone cells from inner bark, the lenticel phellogen; and the suc- ‘S S*HO™ *45° ceeding phellogens arise more or less independently from the deeper layers of the bark. The primary cork phellogen first appears on the south side of an erect stem and normally on the upper surface of a horizontal branch. From these regions it gradually extends laterally and usually forms a complete layer in the course of three or four years. The development, however, follows no definite rule. For example, in one eight-year old stem there was no cork on the north side except in the immediate vicinity of the lenticels, while in another stem of the same age there were five layers of cork on the South side and three on the north. These observations show that a phellogen layer may be active in one part although it has ceased to be functional in another. They also show that there is no definite relationship between the age of the stem or branch and the number of layers of phellogen. The early appearance of cork in the regions exposed to the sun is probably due to the fact that the sassafras is an intolerant species and that the cork protects the deeper tissues from Sun scalding. 442 BOTANICAL GAZETTE [JUNE The primary lenticels are always formed directly beneath the stomata, following in this respect the general rule first enunciated by TRECUL. Some of the lenticels never break through the epidermis but remain in an undeveloped condition. The lenticel phellogen arises from the layer of cells just within the epidermis. The thin- _ walled complementary cells are at first closely packed together. After about twelve layers of these cells are formed the epidermis is ruptured, and the complementary cells as they become exposed sepa- rate from each other and present very irregular and distorted outlines. The mature lenticel agrees with the second of the types described by Devaux‘ and shows no distinct layers of cork among the. comple- plementary cells (jig. &). In some cases, however, a lenticel contains a few scattered stone cells (fig. 8, st). Secondary lenticels are developed from secondary phellogens and make their appearance in the splits of the bark. These lenticels break through the gt Pig.ic: *i [| i OS é OY ERS we (f ef A O eo The credit to Wisconsin is doubtless due to Dr. Lapua, already cited. South of Chicago this oak appears in the southern part of Cook County in the town of Thornton, extending sparingly up Thorn Creek for a short distance, where it grows in company with Q. ellipsoidalis. It is most abundant east of the village of Thornton, making a good part of a wood growing in a soil of sandy peat, patches of sphagnum being common under the trees. Eastward it is found in occasional spots and in similar soils, and in the clayey soils of swamps in Lake and Porter Counties, Indiana. It comes into the dune region of Lake Michigan north of the village of Porter, in a sandy humus soil similar to that near Thornton. Southward from here in Indiana it increases in frequency and abund- ance. In eastern Illinois it reappears south of the Thorn Creek localities after one crosses the range of hills here forming the water-shed of Lake Michigan basin (the Valparasio moraine), and is oe along: the Kankakee River at Momence. Whether Q. ellipsoidalis occurs south of the most northern counties of Indiana there is no evidence at hand to show. Some time spent in examin- ing the flora in the vicinity of North Judson and English Lake in Stark County did not reveal its presence, though the pin or Spanish oak was common along the Kankakee River there. Specimens of oaks sent from Bluffton in the eastern part of the state, a short distance south of Fort Wayne, lacked this: species, but contained Q. palustris and Q. texana Buckley. - ' It is evident from this survey that Q. ellipsoidalis replaces to a large extent in the north of the Middle West the more southerly Q. palustris. But it is usually with a different and drier habitat, and an adaptability to a wider range of conditions. The boundaries of the two overlap in southern Michigan, northern Indiana, northern ‘Illinois, eastern-central Towa, possibly in southern Wisconsin. It may also be of interest to add that the northern bounds of another biennial fruited oak, the shingle oak (Q. imbricaria) correspond quite generally with those of Q. palustris.— -E. J. Hitz, Chicago. 5 Illinois Agricultural Report 1859:596. CURRENT LITEKATURE. BOOK REVIEWS. Botanical dictionary. IN 1900 JACKSON published the first edition of his Glossary of botanic terms, and last fall the second edition appeared.‘ We welcomed the first edition? as being a marked improvement upon any existing dictionary, and criticized but lightly the most obvious shortcomings. The compiler, most competent in many respects, had certain limitations by reason of his unfamiliarity with the content and consequently the terminology of morphology and physiology, and our general criticisms lay along these lines. In judging the second edition one looks to see whether this weakness of the first has been removed, either by the author’s own efforts, or by his associating with himself those who could supply the lacking knowledge. We find that the “revised and enlarged” of the title page means only that typographical and minor errors have been corrected in the plates of the first edition, and that a supplement of 68 pages has replaced the former “Additions during printing.’” ne can overlook much in a first edition that cannot be forgiven in a second. Perhaps there will be a third with a resetting that will allow the necessary improve- ment. In that hope we may point out certain objectionable features that should receive attention. In the first place it would be desirable to relegate to a separate list the many terms which have become obsolete, most of which are adopted from LINDLEY’S Glossary and were antiquated in his day. Technical language changes rapidly and such terms should be put into a museum and labeled as exhibits, if shown at all. We should then escape reading (except we were on antiquarian research bent) that an ovule-tube is “‘a thread-like extension of the amnios, rising beyond the foramen;” and, when we turn in wonder to see what the amnios in plants could have been, learning that it is ‘‘a viscous fluid which surrounds certain ovules at an early stage.”” We do not need often to know that prosphyses were “abortive pistillidia of the muscal alliance,” and the youngster who has occasion to look for the word should learn that both it and its definition are mere sur- vivals from a past century. Second, space could be gained by omitting to define common words which have no technical meaning, such as congeries, enlargement, entangled, evapora- « JACKSON, BENJAMIN Daypon, A glossary of botanic terms, with their deriva- tion and accent. Second edition, revised and enlarged. 12mo. pp. 37!- London: Duckwith & Co. (Philadelphia: J. B. Lippincott Co.) 1905. 2 Bot. GAZETTE 31: 68. 1901. 448 See 1906] CURRENT LITERATURE 449 tion, minute, parallel, sex, tall, wound. All of these and many others are now include Third, more care should be taken to make definitions sufficiently general to include the various uses of the word, rather than so special as to refer only to particular uses. Thus, conjugate appears as an adjective, but not as a verb; conjugating tubes are defined in a special and unusual sense for the Rhodophyceae and not at all for the Conjugatae; for pistil is given (after a wholly erroneous definition in reference to spermatophytes) an obsolete sense which is restricted to the genus Andreaea, when in the same sense it was formerly applied to the archegonia of all mosses; retardation is not mentioned as other than the “‘influ- ence of light on growth in certain structures;” and a fat enzyme is defined merely as an enzyme ‘‘converting olein into oleic acid and glycerin.” Fourth, greater aia is sadly ar A few examples will illustrate this: Galvanotro pic, A istics, etc.;” geotropism, “‘the force of gravity as shown By curvature;” geotaxis, Secveinias in plants caused by gravity;” stamen, “‘a male sporophyll;” pistil, “the female organ of the flower;” stamz- nate, ‘‘applied to flowers which are wholly male;” oogenesis, “the formation of the oosphere, the early stage of the ovule”’ (but oosphere is correctly defined later in the same paragraph!); sap-pressure, “the force exerted on passing upwards through the tissues;” spermatogenesis, ‘the development of the male elements, antherozoids, pollen-grains, and analogous bodies;”’ and so o Fifth (a matter for the publisher), the use of a more flexible paper ee looser binding would contribute much to the handiness of the volume.—C. R. B. MINOR NOTICES. The dynamics of living matter.s—In the spring of 1902 Professor JACQUES LoEB was invited to deliver a series of lectures at Columbia University. In these lectures, eight in number, he presented the gist of his researches upon the dynamics of living matter. This book, forming the eighth volume of the Columbia University Biological Series, is a somewhat more complete survey of the field of experimental biology, says the author, than was possible in the lectures. In ten “‘lectures’’ he discusses the general chemistry and physical constitution of living matter, certain physical manifestations of life, the réle of electrolytes, effects of radiant energy, heliotropism and other tropisms, fer- tilization, heredity, and regeneration. Through the publication of his collected papers in English in the Decennial Publications of the University of Chicago+ Professor Lors’s point of view and the general results of his experimentation have become even more generally 3 Logs, J., The dynamics of living matter. Columbia University Biological Serizs VIII. 8vo. pp- xiit+233. figs. 64. New York: The Columbia University Press. 1906. $3 4 Logs, J., Studies in general physiology, 1905. 450 BOTANICAL GAZETTE [JUNE known than from the originals. The topics named above are naturally those with which the author has chiefly concerned himself, and it cannot be said that the present volume contributes to general physiological literature anything new. The book is rather a new setting of the brilliant work and suggestive ideas of the author, that have previously enriched physiology, and with them is related the results of others in such wise as to round out the presentation. The lectures are readable and instructive, and they are especially commended to the attention of plant physiologists, who are too apt to pass over literature not strictly per- taining to plants.—C. R. B The problems of life——The third part of this book’ was issued last winter, and extends the author’s fundamental hypothesis to the phemonena of fertili- zation and heredity. To him, if one admits the premises, the difficulties of these phenomena fade away like morning mists. The work does not cite defi- nite observations, nor show, except in the most general way, how the known facts can be correlated by this theory; but it presents a clearly reasoned, logical series of deductions, which impresses the reader at once as too simple to be true. Moreover, one is naturally shy of a theory, which, beginning with an assump- tion regarding the molecular structure of protoplasm and the nature of assimi- lation, makes reproduction a necessary and inevitable consequence of these assumptions, while heredity likewise follows as a matter of course from the phenomena of fecundation. We were inclined to welcome the molecular con- ceptions of the first part,° as possibly embodying a fruitful theory, but we can- not follow the author as he widens and heightens his construction upon the acute fundamental assumption. Such inverted pyramids of logic can have no stability.—C. R. B Pfeffer’s Physiology —The third and last volume of this work was pub- lished about the middle of March.? It treats at length of the movements of plants, including the mechanical responses to various stimuli; and briefly of the production of heat, light, and electric tensions, and of the sources and trans- formations of energy. The translation, or rather the interpretation of the original, is of the same satisfactory character as in earlier volumes. As before, the editor has introduced supplementary and critical matter in footnotes; and in an appendix of eight pages he has supplied some important facts not men- tioned in the first two volumes, and a summary of the more recent literature, especially that connected with the present volume. Throughout, his critical 5 GIGLIo-Tos, ERMANNO, Les problémes de la vie. IIJ¢ partie: La fécondation et hérédité. 8vo. pp. vili+189. Cagliari: The author, at the University. 1905. /r. 8- © Cf. Bot. GAZETTE 31:275. 1901. 7 PFEFFER, W., The physiology of plants, a treatise upon the metabolism and and sources of energy in plants. Second fully revised edition; translated and edited by ALFRED J. Ewart. Volume III. Imp. 8vo. pp. viii+4sr. figs. 70. Oxford: The Clarendon Press. 1906. ats. 1906] CURRENT LITERATURE 451 care and acumen have enriched the already valuable work of the author, so that English readers are indebted to him for far more than a translation of pecul- iarly difficult German. To recommend the English form to all libraries and laboratories as a standard work of reference is, at this date, really quite super- fluous.—C. R. B British flowering plants.—Under this title Lord Avepury,® better known as Sir Jonn Lussock, has brought together a mass of desultory notes on various things connected with a great many plants. The author says that this work is “to describe points of interest in the life-history of our British plants; to explain, as far as possible, the reasons for the structure, form, and color; ane to suggest some of the innumerable problems which still remain for solution.’ A glossary and an introductory chapter indicate that the book may be used by those with no botanical training; and perhaps it will be chiefly so used. Each species is taken as the occasion for the presentation of all sorts of facts and fancies and questions in reference to it, as though the author had emptied his note book under that head. There is no distinct organization and no pam ae index; so that the botanist will simply have to ‘‘run on” to things.—J. Spring flora of Ohio.—Under the title ‘Spring Flora,” the botanical staff of Ohio State University has issued a manual for beginners and amateurs.° It is a revised edition of KELLERMAN’S “Spring Flora of Ohio,” and its range as been extended so as to include Ohio and Indiana and the adjacent states. The time range extends from the opening of the season into the first part of June; and such difficult groups as grasses and sedges are not included. There is also a key to the trees and shrubs based on leaf and twig characters.—J. M. C. Flora of Norway.—AxeEL Buiyrtt’s completed Handbook of the Norwegian Flora, including the vascular plants, has been issued under the peak of AHL.'° In reality it has been in preparation since 1861, having begun by the father, continued by the son, whose name is on the title page,

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Patients should be encouraged to take two quarts per day, i they can, and the relief they will obtain will be all the argument necessary after the first day or so. “The Results Satisfy Me of Its Extraordinary Value.” Dr. tas, AGT of New Orleans, Ex- aur of the State Board of Health of Louisiana, n affections of the kidneys and ‘“T have prescribed BUFFALO LITHIA WATER ... nary passages, particularly in Gouty subjects, in Albuminuria, and in irritable euuatied of the Bladder and Urethra in females. The results satisfy me of its extraordinary value in a large class of cases usually pit difficult to treat.’ “I Have Witnessed Decided Beneficial Results from Its Use.” Wm. B. Towles, M. 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It tells how to prevent Sic A Perfect Food. ness. Send fora free co me to Henry B. Platt, 42 Cliff Street, New York, sole . 46 Highest Awards manufacturers of ——— Piatts Chlorides, es dorless Disinfee tant.| Walter Boker @ Co, Ltd. Dorchester, Mass. A coloriess'liquid ; powerful, safe Src micaL Instantly destroys foul odors and disease-breeding m Specially prepared for house- hold use. Sold only in quart onan “by druggists everywhere j Be Fair to Your Skin, and It Will | Be Fair to You--and to Others | A Beautiful Skin can only be secured through Nature’s work. Ghastly, horrid imitat‘ons of Beauty are made by cosmetics, balms, powders, and other injurious compounds. They put a coat over the already clogged pores of the skin, and double the injury. Now that the use of cosmetics is being inveighed against from the very pulpits, the importance of pure soap becomes apparent. The con- stant use of HAND SAPOLIO produces so fresh and rejuvenated a condition of the skin that all incentive to the use of cosmetics is lacking. HAND SAPOLIO is Ss O PURE that it can be freely used on a new-born baby or the skin of the most delicate beauty. that it can be a part of the invalid’s supply with beneficial results. SO SIMPLE ee 1 } a S18) EFFIC ACIOU - sets =. — ~ pram into a state of ‘surgical clean 6 rT 24% Te Dal A RTD Cy bee bent established over 50 YEARS. By our system ol |