Surface, Frank M.

The Ohio Journal of Science

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The Ohio ZS£aturalist,PUBLISHED BYThe Biological Club of the Ohio State University.Volume VI. NOVEMBER, 1905. No. 1.TABLE OF CONTENTSSURFACE—Contribution to the Life History of Sangninaria Canadensis 37!)SCHAFFNER—The Classification of Plants, II '. 886IIINE—New Species of North American Chrysops 391RIDDLE—Brush Lake Protozoa 394FISCHER—An Abnormal Cone of Pinus Laricio 396GLEASON—Notes from the Ohio State Herbarium. IV 897News and Notes 39!)SURFACE—Meeting of the Biological Club 400CONTRIBUTION TO THE LIFE HISTORY OF SANGUINARIACANADENSIS.*FRANK M. SURFACE. ' " ^Sanguinaria is a monotypic genus of the Eastern UnitedStates belonging to the family Papaveraceae. Apparently novery great amount of morphological work has been done on themembers of this family. Shaw (3) in a recent paper gives someobservations on certain Papaveraceae among which is Sanguin-aria. Some of the conclusions in the following paper do not agreewith his. This may possibly be due to a difference in climaticconditions between the two stations where material was col-lected, although Shaw does not state definitely, the locality from,which his material came.The following study of the development of Sanguinariacanadensis L. was begun in April, 1904. Material was collectedat intervals of about one week until the middle of June whenthe seeds were mature and had fallen to the ground. In the fallmaterial was collected in October, November and December,while in the Spring of 1905 young capsules were taken at inter-vals of about two weeks from March 1st to the middle of April.The material was killed and fixed in chromo-acetic acid,passed through the alcohols and imbedded in paraffin. Sectionswere cut on a rotary microtome 10-12 microns thick. In theyounger stages the entire ovulary was cut, either transversely orlongitudinally but in the older stages the individual ovules weresectioned. The orientation of the ovule was not difficult owing' • * Contributions from 'the Botanical Laboratory of the OhioState University, XXI.380 The Ohio Naturalist. [Vol. VI, No. 1,to the distinct crest developed along one side. In the oldeststages it was found necessary to remove the hard covering of theseed before cutting. Several stains were employed amongwhich were anilin-safranin followed by genetian violet, Dela-field's haematoxylih and Haidenhein's iron-alum-haematoxylin.The latter well known stain proved to be the most successful inthe stages of the development of the megaspores and embryo-sac, while Delafield's stain gave the best results with themicrospores.The flower of Sanguinaria begins its development early in thesummer and the ovules and stamens are considerably advancedby the end of September. Material taken on Oct. 6th showed thesingle, hypodermal, archesporial cell distinctly differentiated(Fig. 2). The ovules at this time were just beginning to curvebut no traces of the integuments were discernable (Fig.l). Atthis time the microsporocytes were just separating (Fig. 19) anda few of the tapetal cells already had two nuclei. There arethree intermediate layers between the epidermis and. tapetum(Fig. 19). Material taken on Nov. 9th showed but little changein the development of the ovules, except that they had elongatedand the curving was much more marked. The integuments werejust beginning to make their appearance but the archesporialcell was still undivided. At this time the microsporocytes hadundergone some division and some tetrads were observed. Thetapetal cells contained two nuclei (Fig. 20).By Dec. 9th the archesporial cell had divided twice givingrise to the megasporocyte with two parietal cells above (Fig. 3).The curving of the ovule was much more marked than in theprevious stage and the beginnings of both integuments could bereadily seen in central sections. The division of the micro-sporocytes must have proceeded rather rapidly for at this timethe microspores were present with large resting nuclei and thetapetum was dissolving (Fig. 21). In this stage the microsporespass the winter.No material was taken from December until the first ofMarch when it seemed that the weather conditions were favor-able for the continued growth of the plant. Sections of materialfixed March 1st showed that there had been but little activityduring the winter months. One of the parietal cells had againdivided so as to form a row of three cells above the large mega-sporocyte (Fig. 4). In some instances transverse division hadtaken place forming a considerable tissue (Fig. 5). The integu-ments had increased quite a little in size and now reached morethan half way around the developing ovule. The curving of theovule was complete at this time. The microspores were inpractically the same stage in which they were found in Decem-ber. The nucleus was undivided but the vacuole in the centerNov., 1905.] Life History Sanguinaria canadensis. 381of the microspore had enlarged so that the nucleus was forcedto one side of the cell. Material taken on March 13th showedbut little change from that taken on the previous date. Themegasporocyte had enlarged quite a little and seemed to be pre-paring to divide (Fig. 4), and a few spindles were observed(Fig. 6). The microspores showed no change.Material fixed on March 22nd showed that the megasporocytehad divided into four megaspores the lowest being the largefunctional one (Fig. 7). The arrangement of the megasporesvaried considerably. In some cases the transverse walls werenearly parallel forming a row of cells but frequently they werevery irregular (Figs. 7, 8). These divisions are interesting inthat they show a great similarity to the divisions of the micro-sporocyte and often result in a nearly typical tetrad (Fig. S).Above the megaspores the rows of parietal cells could be dis-tinctly seen (Fig. 7). These varied in number but it is evidentthat considerable division has occurred in the three originalparietal cells. Vesque (1) states that Papaver orientale has noparietal cells. The nuclei of the microspores were divided atthis time (Fig. 21). Pollen grains taken at later dates did notseem to show any further division. It is probable that the gen-erative cell does not divide until after the tube has been formed.Strasburger (2) however, reports that in Papaver the generativenucleus divides in the pollen grain.From this time the development of the functional megasporeis rather rapid. Material taken on March 28th showed the com-plete eight celled embryo-sac and often the two polar nuclei hadalready fused to form the definitive nucleus (Fig. 9). Theantipotals are rather large at this stage and each has a singlenucleus. Material taken on April 10th showed the oosphereand definitive nucleus still undivided but the antipodals werevery large and each contained two nuclei (Fig. 10). This stagecorresponded with material taken the year before at about thesame date. The egg then seems to be fertilized about the firstweek in April. Many of the sections show remains of the pollentube which is very prominent (Figs. 10, 11). The synergidsseem to disappear early. Remains of one or more of them couldusually be seen at this stage, lying to one side of the oosporeand staining very dark (Figs. 10, 11). The early divisions ofthe embryo seem to occur very slowly for it was not until May12th that the first division was observed (Fig. 13). This makesa remarkably long period of rest for the fertilized egg. Inmaterial killed May 16th the three celled embryo was found(Fig. 14). The divisions of the definitive nucleus had begun inearly April and by the latter part of that month the endospermhad 20 to 30 nuclei.382 The Ohio Naturalist. [Vol. VI, No. 1,The development of the embryo from this stage on was nottraced very closely but the divisions must occur rather rapidlyfor on-May 23rd the embryo had reached about the 12-celledstage (Fig. 15). On May 29th a large spherical embryo waspresent (Fig. 16). The suspensor had lengthened considerablyand vSections showed a double row of cells (Fig. 16). At this timethe antipodals were degenerating (Fig. 17). They lie in a broaddepression at the base of the sac which is greatly enlarged andfilled with endosperm. The behavior of the antipodals recalls thecondition of these cells in certain Ranunculaceae. On June 6ththe ovules had matured and were falling to the ground. Theintegument is ve?y hard and it was found necessary to removeit before imbedding in paraffin. At this time the embryo wasmuch larger and the two cotelydons were well developed (Fig.18). The suspensor was still present but showed signs ofdegeneration.SUMMARY.The flowers of Sanguinaria begin to develop very early inthe summer previous to the year in which they blossom.The development of the microsporocytes and microspores ismuch more rapid than that of the megaspores for the micro-spores are formed before the winter season begins.The ovule passes the winter in the megasporocyte stage andduring March its development is very rapid while the microsporedoes not renew activity until the last of March and early April.Three parietal cells are formed and" later these divide forminga parietal tissue of considerable size.The division of the megasporocyte frequently results inrather typical tetrads.The generative nucleus does not appear to divide in thepollen grain.The long resting period of the oosphere is especiallyinteresting.BIBLIOGRAPHY.1. VESQUE, J.Nouvelle recherches sur le development du sac embryon-naire des Phanerogames Angiospermes. Ann. Sci. Nat. Bot.VI, 8:261-390, Pis. 12-21. 1879.2. STRASBURGER, E. . •  .. . -Neue Untersuchungen uber den Befruchtungsvorgangbei den Phanerogamen. Jena. ' 1884. jjj3. SHAW, CHARLES H. * 'Notes on sexual generation and the development of: theseed coats in certain of the Papaveraceae.. Bull. Tor B,otClub 31:429-433, PL 15. 1904.Nov., 1905.] Life History Sanguinaria canadensis. 385The above paper was prepared in the Department of Botanyof trie Ohio State University as a minor for the degree of Masterof Arts. The work was done under the direction of AssociateProfessor J. H. Schaffner, to whom the writer wishes to expresshis sincere thanks for the kind assistance and suggestions,without which the paper could not have been completed,DESCRIPTIONS OF PLATES.All drawings were outlined under a camera lucida with thefollowing optical combinations:Bausch and Lomb—1 oc, § obj.—Fig. 1.Bausch and Lomb—1 oc, ^ obj.—Figs. 20, 21.Bausch and Lomb— f oc, | obj.—Figs. 2-8, 13, 1.5-17, 22, 23.Bausch and Lomb—2 oc, ^ obj.— Figs. 18, 19.Leitz—4 oc, 7 obj.—Fig. 10.Leitz—6 oc, 7 ob'j.—Fig. 14.Zeiss—18 oc, Leitz 7 obj.—Figs. 9, 12.Zeiss—18 oc, Leitz 3 obj.—Fig. 11.Fig. 1. Section of young ovulary, showing incipient ovules. Killed10-6-'04.Fig. 2. Archesporial cell. Killed 10-6-'04.Fig. 3 Megasporocyte with two parietal cells. Killed 12-9-'O4.Fig. 4. Megasporocyte with three parietal cells. Killed 3-13-'O5.Fig. 5. Megasporocyte with parietals divided. Killed 3-l- '05.Fig. 6. First division of megasporocyte. Killed 3-13-'O5.Fig. 7. Four megaspores and parietal cells. Killed 3-22-'O5.Fig. 8. Four megaspores showing tetrad arrangement. Killed3-22-'O5.Fig. 9. Eight celled embryo-sac. Killed 3-28-'O5.Fig. 10. Embryo-sac with oospore. Antipodals with two nuclei.Killed" 4-16-'O4.Fig. 11. Outline of ovule with oospore. Killed 4-10-'04.Fig. 12. Oospore with remains of synergids. Killed 4-16-'O4.Fig. 13. Two-celled embryo. Killed 5-12-'04.Fig. 14. Three-celled embryo. Killed 5-16-'O4.Fig. 15. Young embryo showing eight cells in section. Killed5-23-'04.Fig. 16. Spherical embryo with long suspensor. Killed 5-29-'O4.Fig. 17. Degenerating antipodals. Killed 5-29-'O4.Fig. 18. Embryo showing large suspensor and cotyledons at the timethe seed falls. Killed 6-6-'O4.Fig. 19. Section of stamen with microsporocytes and tapetal cells.Killed 10-6-'05.Fig. 20. Section of microsporangium showing microsporocytes justbefore formation of tetrads. Tapetal cells with two nuclei. Killed 11-Q-'O5Fig. 21. Microspores with remains of tapetum. Killed 12-9-'O5.Fig. 22. Single microspore. Killed 12-9-'O5.Fig. 23. Pollen grain. Killed 3-22-'05.384 The Ohio Naturalist. [Vol. VI, No. 1,OHIO NATURALIST. Plate XXVSURFACE on "Sanguinaria.Nov., 1905.] Life History Sanguinaria canadensis. 385OHIO NATURALIST. Plate XXVISURFACE on "Sanguinaria

Contribution to the Life History of Sanguinaria Canadensis

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Contribution to the Life History of Sanguinaria Canadensis

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