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Floral Organogenesis in Five Genera of the Marantaceae and in Canna (Cannaceae)

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Floral Organogenesis in Five Genera of the Marantaceae and in Canna (Cannaceae) Floral Organogenesis in Five Genera of the Marantaceae and in Canna (Cannaceae) Bruce K. Kirchoff American Journal of Botany, Volume 70, Issue 4 (Apr., 1983), 508-523. Kirchoff, B. K.1983. Floral organogenesis in five genera of the Marantaceae and in Canna (Cannaceae). American Journal of Botany 70: 508-523. Made available courtesy of the Botanical Society of America: http://www.jstor.org/stable/i341826 Your use of the JSTOR archive indicates your acceptance of JSTOR' s Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR' s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSToR archive only for your personal, non-commercial use. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. American Journal of Botany is published by Botanical Society of America. Please contact the publisher for further permissions regarding the use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/botsam.html. American Journal of Botany ©1983 Botanical Society of America JSTOR and the JSTOR logo are trademarks of JSTOR, and are Registered in the U.S. Patent and Trademark Office. For more information on JSTOR contact [email protected]. ©2001 JSTOR http://www.jstor.org/ Tue Oct 9 12:27:39 2001 Amer. J. BOt. 70(4): 508-523. 1983. FLORAL ORGANOGENESIS IN FIVE GENERA OF THE MARANTACEAE AND IN CANNA (CANNACEAE) 1 BRUCE K. KIRCHOFF2 Department of Botany, Louisiana State University, Baton Rouge, Louisiana 70803 ABSTRACT The paired flowers of all species of the Marantaceae studied, except Monotagma plurispicatum, are produced through the division of an apical meristem with a tunica-corpus structure. The solitary flowers of M. plurispicatum develop from a similar meristem which does not bifurcate. The paired flowers of Canna indica are produced in the axil of a florescence bract through the formation of a bract and an axillary flower on the side of the primordium which gives rise to the largest flower of the pair. The sequence of organ initiation for both families is: calyx, corolla and inner androecial whorl, outer androecial whorl, gynoecium. The sequence of sepal formation is opposite in the two families. In the Cannaceae it leads directly into the spiral ereated by the formation of the other organs, while in the Marantaceae the sequence of sepal formation follows a spiral opposite to that of the other floral organs. The members of the corolla and inner androecial whorl separate from common primordia. In general these common primordia sepa- rate into a petal and an inner androecial member through the initiation of two growth centers, at the same level, in the dorsal and ventral flanks of the primordium. In Ischnosiphon elegans and Pleiostachya pruinosa the stamen is initiated at a lower position than the petal in the ventral flank of the common primordium. A similar pattern of initiation is described for the callose staminode in Marantochloa purpurea and Canna indica. This pattern is interpreted as a variation on the more generalized pattern of inner androecial formation found in the other genera. FLoRAL oRGANoGENESIS and floral growth in flower; Maranta spp., Calathea spp. — house five genera of the Marantaceae, and one species plants). Genera of these families were selected of Canna (Cannaceae) will be explored in a for study for several reasons: l) floral devel- series of two papers of which this is the first. The species under study include: Calathea leo- opment was not well known; 2) Andersson's pardinia, Calathea lancifolia, Calathea vinosa, (1977) revision of the genus Ischnosiphon in- Ischnosiphon elegans, Marantochloa purpurea, cludes an assessment of the relationships of the Monotagma plurispicatum, Pleiostachya prui- neo-tropical genera which allowed the selec- nosa (all Marantaceae), and Canna indica tion of a natural subset of the family for study; (Cannaceae). This paper presents a description and 3) the position of the Marantaceae in the and comparison of the patterns of organogen- order Zingiberales allowed the identification of esis found in these species. the Cannaceae as its sister group (that group The Marantaceae and Cannaceae are well which shares a common ancestor with the Mar- circumscribed pan-tropical groups of plants antaceae and with no other third group). some of whose members are of economic im- The choice of genera within the Marantaceae portance (Maranta arundinacea— arrowroot was based on Andersson's (1977) revision of Ischnosiphon. Selection of species within each ' Received for publication 11 February 1982; revision of the genera was based on availability of flow- accepted 2 August 1982. ering material. Three species of were This paper is based on a portion of a dissertation sub- Calathea mitted in partial fulfillment of the requirements for the chosen to represent different sections of this degree of Doctor of Philosophy in the Department of Bot- large genus (Schumann, 1902). Collectors, col- any in the Graduate School of Duke University. I am lection numbers and location of voucher spec- deeply indebted to Dr. R. A. White for his advice and imens are given in Table 1. guidance during the completion of my degree. I am also grateful to Mr. Russell Goddard for his help in the prep- MATERIALS AND METHODS—FlowerIng ma- aration of this manuscript, Sally Baker for preparing Fig. terial was collected both in Costa Rica, during 1, 3 and 4, and to Dr. Shirley Tucker, Dr. U. Posluszny, the summer of 1978, and from plants culti- and an unidentified reviewer for their suggestions to im- vated in the Duke University greenhouses. Dr. prove the manuscript. The SEM work included in this paper was supported by a Duke University Graduate G. Prance was kind enough to supply material School Research Award. of Monotagma plurispicatum from Brazil. Present address: Dept. Botany, Hebrew University of The floral apices for epi-illumination study Jerusalem, 91904 Jerusalem, Israel. and paraffin sections were first fixed in for- malin-acetic acid-alcohol (FAA: 50 ml 95% 508 April, 1983] KIRCHOFF — FLORAL ORGANOGENESIS 509 TABLE 1. Species examined Collection Voucher Species no. location Calathea lancifolia Boom KirchOff 281 DUKE Calathea leopardinia Kres s 78 -1001 DUKE (Bull) Reg. KirchOff 381 Calathea vinosa Kennedy Kress 77-879 DUKE Ischnosiphon elegans Standl. Kress 78-896 DUKE Marantochloa purpurea (Ridl.) M. -Redh. Kress 78-894 DUKE 0 Monotagma plurispicatum (Koernicke) Schum. Prance 26320 NY Fig. 1. Diagrams of an imaginary species of the Mar- Pleiostachya pruinosa antaceae showing the relationships of the floral parts. P, (Reg.) Schum. Kress 78-916 DUKE petals; FT, floral tube; C, sepals; 0, ovary; Sc, callose Canna indica L. Kress 76-541 DUKE staminode; A, anther; Pa, petaloid appendage to anther; ST, style; STI, stigma; Sh, hooded staminode; Si, outer staminode. ethanol, 5 ml glacial acetic acid, 10 ml for- malin, 35 ml H 2 0) and dehydrated to 95% in the form of the androecial members. Ac- ethanol in an ethanol series. They were then cording to Pai (1965) the androecium of both stained in 0.5% acid fuchsin (Sattler, 1968) for families is composed of two trimerous whorls. 24 hr and destained in 98% ethanol for l-2 The inner whorl contains the functional anther, wk. The apices were transferred to 100% eth- which is reduced to two loculi with an asso- anol and photographed under epi-illumination ciated petaloid appendage (Fig. l), and two on a Leitz Orthomat microscope equipped petaloid staminodes. In the Cannaceae these with an Ultropak Illuminator and dipping petaloid members are the labellum and the cones (Posluszny, Scott and Sattler, 1980). inner staminode. In the Marantaceae they are When all the buds from a species had been the hooded staminode, which encloses the style photographed they were transferred to tertiary and stigma before pollination, and a callose butyl alcohol and embedded in "Tissue-prep," staminode (Fig. l) on which the tripped style a paraffin embedding medium. Sections were rests after pollination (Fig. 2). cut at 6-9 kim on an American Optical 820 The outer androecial whorl is trimerous in Microtome. Buds were occasionally left in construction but is seldom represented by three 100% ethanol for as long as 3 months with no mature structures (for exceptions, see Costerus, ill effects in sectioning. The sections were 1916). The members of this whorl are always stained in tannic acid-ferric chloride, safranin petaloid and are referred to as the outer stam- and fast green (Berlyn and Miksche, 1976) and inodes. For most of the species included in this mounted in "Permount." Photographs were study (Canna indica, Calathea vinosa, C. leo- taken using a Leitz Orthomat microscope. pardinia, C lancifolia, Monotagma plurispi- Apices for scanning electron microscopy catum, Pleiostachya pruinosa, Ischnosiphon (SEM) were fixed in glutaraldehyde and de- elegans) only one member of this whorl is fully hydrated to 100% freon in an ethyl alcohol- developed (Fig. l). Marantochloa purpurea, freon series. Critical point drying was carried however, has two. out in a Bomar SPC-900/EX Critical Point The ovary in both families is inferior and Dryer. The apices were mounted on stubs, trilocular. In the Cannaceae each locule con- sputter coated with gold-palladium in a Film- tains two series of anatropous ovules while the Vac Inc. EMS-41 Mini-Coater, and observed Marantaceae possess, at most, one anatropous and photographed at 15 kv on a Philips 501 to campylotropous ovule per locule. For many Scanning Electron Microscope. genera of the Marantaceae (including Ischno- siphon, Pleiostachya, and Monotagma) ovules RESULTS— Organography— Floral structure are not produced in two of the three loculi, in both the Marantaceae and Cannaceae is sim- leaving only one ovule per flower.
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