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Shoot and Floral Development in Calla Palustris (Araceae-Calloideae) Author(S): R

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Shoot and Floral Development in Calla Palustris (Araceae-Calloideae) Author(S): R Shoot and Floral Development in Calla palustris (Araceae-Calloideae) Author(s): R. W. Scribailo and P. B. Tomlinson Reviewed work(s): Source: International Journal of Plant Sciences, Vol. 153, No. 1 (Mar., 1992), pp. 1-13 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2995699 . Accessed: 12/04/2012 13:42 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. http://www.jstor.org Int. J. Plant Sci. 153(1):1-13. 1992 ? 1992 by The University of Chicago.All rights reserved. 1058-5893/92/5301-0005$02.00 SHOOTAND FLORAL DEVELOPMENT IN CALLA PALUSTRIS (ARACEAE-CALLOIDEAE) R. W. SCRIBAILO'AND P. B. TOMLINSON2 HarvardForest, HarvardUniversity, Petersham,Massachusetts 01366 Callapalustris shows annualproduction of one or two inflorescenceson each ephemeralseasonal shoot. Shoots are sympodial; the terminal inflorescencesare produced in the summer of the year before the mostly perfect flowers open. Overwinteringterminal buds show limited preformationof foliage leaves on renewalshoots; most leaves areneoformed. The spadixproduces a seriesof prominentflower primordia in up to eight irregularorthostichies. Each primordiumdevelops no perianthbut a series of indistinctly trimerousstamen initials in centripetalorder. The gynoeciumoriginates as a single dorsiventrallyasym- metriccollar-like outgrowth that becomesovate and enclosesthe seriesof basalanatropous ovules, showing no evidence of a pseudomonomerouscondition either in development or vasculature.Floral variation within a spadix is considerable,distal flowersare usuallywholly male, basal flowersshow reducedstamen number,but no strictlyfemale flowersare produced.Vascular traces to the floralappendages are derived from an anastomosingsystem that obscuresany uniformpattern. Morphogenetic considerations describe the vascular system better than any referenceto imaginaryevolutionary antecedents. Introduction (1974). Flowers open in the late spring,the fruits Despiteconsiderable floral diversity in Araceae, ripening in late summer of the same year. virtuallyno developmentalstudy of aroidflowers In the classification of the Araceae developed has beencarried out. Diversityof flowersis usu- by Englerand Krause(Engler 1877; Krause 1908), ally described,in evolutionaryterms, as a mod- Calla is included in the subfamily Calloideae to- ificationof a dimerousor trimeroushermaph- gether with three other north temperate genera, roditicflower, even thoughAraceae with perfect Lysichiton, Orontium, and Symplocarpus.Gra- flowershave themselvesconsiderable organiza- yum (1984, 1990) suggests,primarily on the basis tional diversity (Engler1884). Developmental of pollen morphology,that Calla is more isolated studiesof flowersare essentialto a preciseun- within the family and includes it as a monotypic derstandingof morphologythat can revealcom- subfamily,Calloideae, the remainingthree genera mon floralplans of systematicand phyleticsig- being added to other groups. Barabe and Forget nificance(Leins et al. 1989).In this presentstudy (1987) also suggest a position isolated from Ly- we show that the inflorescenceaxis (spadix)of sichiton, Orontium,and Symplocarpuson the ba- Calla initiateslateral floral primordia that lack sis of cladistic analysis, although they still retain any subtendingbracts. All floralappendages are all four genera in the subfamily Calloideae. initiatedwithout reference to anyconsistent sym- Previous accounts of the shoot organizationof metry,in partbecause stamens are "interpolated" Calla go back to Doll (1843) and Wydler (1856) in anirregular centripetal pattern. The gynoecium and have been summarizedby Braun(1859), En- showsno developmentalevidence for a trimerous gler (1877), Krause (1908), and Dudley (1937) condition.The sequenceof appendagescondi- without the addition of significant new data. tions a vascularsystem without obvious regular- Dudley provided additional information on seed ity. and seedling morphology and pollen develop- Callapalustris is a temperateherbaceous rep- ment and clarified the cytology. More recently resentativeof the primarilytropical family Ara- Barabeand Labrecque(1983) describedthe floral ceae,but with a wide circumborealdistribution. vasculatureand concluded that this provided ev- It persistsby meansof creepinghorizontal axes, idence for a pseudomonomerous condition, the whichare either permanently or temporarilysub- gynoecium being interpreted as being made of merged,or at the surfaceof swamps.Leaves are three fused carpels because of the presence of alwaysemergent. The shootunits are ephemeral, three principal carpellary traces. This interpre- lastinglittle more than a year, but populations tation is contradictedby the observations on flo- persist by both regenerativeand proliferative ral vasculatureby Hotta (1971). The concept of branching,in the terminologyof Tomlinson pseudomonery in aroids apparently originated with Engler(1884) and was extensively elaborat- ed upon by Eckardt(1937). Manuscriptreceived June 1991; revisedmunuscript received September1991. Materialand methods 1 Presentaddress: Purdue University, North CentralCam- Populationsof Callapalustris were studied from pus, 1401 South U.S. 421, Westville, Indiana 46391. Black Gum Swamp, Tom Swamp, and Harvard 2 Authorto whom correspondenceand requestsfor reprints Pond, Petersham, and Spectacle Pond, Athol, should be sent. Massachusetts. Since no differences in patterns I 2 INTERNATIONAL JOURNAL OF PLANT SCIENCES of shoot and floral development were found per year, although the younger spadix in shoots among these collections they have been treated with two spadices may sometimes abort. Phyl- as one. Shoots were sampled at about biweekly lotaxis is distichous throughout,with no change intervals throughout the growing season (May- in the plane of distichy at branching. Although October 1990) and also in the spring of 1991, to no leaves are present in figure3D the distichous a total of about 20 samples of over 150 shoots. phyllotaxis of the shoot is made obvious by the Material was either dissected fresh before fix- arrangementof axillarybuds. In nature,the plane ation or afterfixation in 70%F.A.A. For the study of distichy is horizontalso that plants spreadover of shoot and floral ontogeny fixed spadices were the surface of the substratewith all leaves simi- dehydrated to 95% ethyl alcohol, stained over- larly erected by unequal growth at the base of night in 1% acid fuchsin in 95% alcohol, and each petiole. photographedusing epi-illumination techniques The regenerative renewal shoot is developed (Poslusznyet al. 1980). For scanningelectron mi- by syllepsis (Halle et al. 1978) and is adnate to croscopy(SEM) spadices were dehydrated through the parent axis for about half the associated in- an ethanol series into absolute ethanol, critical- ternode (dotted line in fig. 2). A bicarinatemem- point dried in a SamDri PVT. 3 critical-point branous prophyll is present in the adaxial posi- dryer and coated with gold palladium for 5 min tion (P in figs. 1, 2, 3B). The first foliage leaf on in a Hummer III sputter coater. Spadices were the renewal shoot is inserted on the same side as viewed using an AMR 1000 SEM set at 10 kV at the prophyll, not the opposite side, as would be the Museum of ComparativeZoology of Harvard expected with distichous phyllotaxis (L'o in figs. University. For the study of histology and vas- 1, 2, 3B, 6A). This unusual arrangement also culature at several stages of development, ma- characterizes proliferative vegetative shoots, terial was embedded in paraplast after routine which arise in the axils of each foliage leaf from dehydration in a tertiary butyl alcohol-ethanol dormant buds by prolepsis (PS in fig. 3D). Only series, serially sectioned at 7-10 um in longitu- axillarybuds farthestfrom the shoot tip grow out dinal and transverse planes and stained in saf- and form proliferative shoots with the degree of ranin and alcian green. Serialsections of spadices suppressionof buds increasingmarkedly toward and shoots were analyzedby recordingindividual the shoot tip. The ultimate (Lu) and the unusually sections on a Panasonic GX-4 model AG-1950 placed (Lo) foliage leaf usually lack axillarybuds. multifunction video cassette recorder using a Damage of the leader shoot brings about the re- Javelin Model JE3012 video camera attached to lease of previously dormant buds. a Wild compound microscope. This method of Individual foliage leaves show a regularsym- analysis was complemented by drawings of sec- metry, which influences overall shoot symmetry, tions made with a Wild drawing apparatus.Ad- describedas antidromousby Engler(1877). Each ditional specimens for the study of vasculature leaf has a basal open sheath with overwrapping were preparedby clearingthick sections of older margins, an extended petiole, and a somewhat spadices in 5%alcoholic sodium hydroxide, fol- hastateblade with convolute vernation.The blade lowed by lactic acid, afterwashing in water.These margin
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