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Embryological Development of Lachnanthes Caroliniana (Haemodoraceae) Author(S): Michael G

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Embryological Development of Lachnanthes Caroliniana (Haemodoraceae) Author(S): Michael G Embryological Development of Lachnanthes caroliniana (Haemodoraceae) Author(s): Michael G. Simpson Source: American Journal of Botany, Vol. 75, No. 9 (Sep., 1988), pp. 1394-1408 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/2444463 . Accessed: 05/10/2011 17:31 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]. Botanical Society of America is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany. http://www.jstor.org Amer. J. Bot. 75(9): 1394-1408. 1988. EMBRYOLOGICAL DEVELOPMENT OF LACHNANTHES CAROLINIANA (HAEMODORACEAE)1 MICHAEL G. SIMPSON Department of Biology, San Diego, State University, San Diego, California 92182 AB3STRACT Embryological development of Lachnanthes caroliniana was studied utilizing standard an- atomical techniques and SEM. Lachnanthes has a monocotyledonous anther wall development (endothecial cells with spiral secondary wall thickenings), successive microsporogenesis, and amoeboid (periplasmodial) tapetal development. Mature pollen grains are 2-nucleate with a proximal, fusiform generative cell. Ovules are initiated as 5-7 cylindrical primordia from a common placental base. Basal ovular swellings collectively contribute to the enlarged, peltate placenta. Mature ovules are pleurotropous, anatropous, bitegmic, and crassinucellate; the nu- cellus consists of a chalazal hypostase, radially elongate lateral cells, and a prominent micropylar nucellar cap. Megasporogenesis is successive, forming a linear tetrad of megaspores. Mega- gametogenesis is monosporic; the female gametophyte is of the Polygonum-type with relatively large, pyriform antipodals. Endosperm formation is helobial, resulting in the establishment of a ring of four thick-walled basal endosperm cells (the chalazal chamber) and numerous free nuclei (in the micropylar chamber). The mature cellular endosperm is filled with starch grains and has a chalazal cavity and a thick-walled peripheral layer. The discoid, peltately attached seeds have marginal wings derived by anticlinal divisions and buckling of the outer integument alone. Inner and middle cuticular layers are present in the seed coat. Lachnanthes is similar to all other investigated members of the Haemodoraceae in major embryological features. The significance of embryological evidence with regard to interfamilial classification is discussed. Future studies of ovule and seed development may prove valuable in phylogenetic studies in assessing the homology of placental, ovule, and seed morphology and anatomy. EMBRYOLOGICAL STUDIES have provided char- northern South America, Central America and acters of significant value in angiosperm sys- Mexico, Cuba, and eastern North America. The tematics (see Maheshwari, 1950, 1964; Davis, family is divided into two purported mono- 1966; Johri, 1984). Yet our knowledge of em- phyletic tribes (sensu Simpson, in press b): tribe bryological processes in numerous higher plant Haemodoreae (8 genera), with 3 (rarely 1) sta- taxa consists of an extremely small data base. mens per flower and monosulcate, verrucate Many plant taxa of doubtful systematic affin- (foveolate) pollen grains; and tribe Conostyl- ities remain embryologically unknown, partic- ideae (6 genera), with 6 stamens per flower and ularly among the monocotyledons (Dahlgren porate, rugulate pollen grains. and Clifford, 1982). Embryological studies may Embryological studies have been conducted aid in our understanding of developmental previously on only four species (in four of four- processes and taxonomic relationships; they teen family genera): Anigozanthos flavidus may also, in conjunction with phylogenetic (Stenar, 1927) of the tribe Conostylideae; and analyses, provide a means for hypothesizing Dilatris pilansii (De Vos, 1956), Wachendorfia specific past evolutionary events and give in- paniculata (Dellert, 1933; De Vos, 1956), and sight into the possible adaptive significance of Xiphidium coeruleum (= X. album; Stenar, those events. 1938) of the tribe Haemodoreae. As noted by The Haemodoraceae (Bloodwort family), as De Vos (1956, 1961), these four genera have recently circumscribed and described by Simp- several similar embryological features: an son (in press a, b), are a monophyletic monocot amoeboid (plasmodial) tapetum, successive family consisting of 14 genera and approxi- microsporogenesis, two integuments, crassi- mately 70 species with distributions in eastern nucellate ovules, a Polygonum-type female ga- Australia, New Guinea, southern Africa, metophyte, and helobial endosperm devel- opment (observed only in Dilatris and I Received for publication 1 September 1987; revision Wachendorfia). These embryological data have accepted 6 January 1988. not contradicted the monophylesis of the fam- This study was supported in part by National Science ily, supported more definitively from studies Foundation Grant DEB-8 109909. I thank J. M. Herr, Jr., and an anonymous reviewer for comments made on the of chemistry (Cooke and Edwards, 1981) and manuscript. pollen ultrastructure (Simpson, 1983). [Three 1394 September 1988] SIMPSON-EMBRYOLOGY OF LACHNANTHES 1395 TABLE 1. Embryologicalcharacters of the Haemodoraceaeand relatives Taxon Tapetal type Microspore division Nucellus type Documentation Haemodoraceae Lachnanthes Amoeboid Successive Crassinucellate Present study Anigozanthos Amoeboid Successive Crassinucellate Stenar, 1927 Dilatris Amoeboid Successive Crassinucellate De Vos, 1956 Wachendorfia Amoeboid Successive Crassinucellate Dellert, 1933; De Vos, 1956 Xiphidium Amoeboid Successive Crassinucellate Stenar, 1938 Bromeliaceae Glandular Successive Crassinucellate Dahlgren and Clifford, 1982 Cyanastraceae Cyanastrum Glandular Simultaneous Crassinucellate Fries, 1919; Nietsch, 1941 Hypoxidaceae Hypoxis Glandular Successive Tenuinucellate De Vos, 1948 Pauridia Glandular Successive Tenuinucellate De Vos, 1949 Lanariaceae Lanaria Glandular Simultaneous Crassinucellate De Vos, 1961, 1963 Melanthiaceae (in part) Lophiola Glandular Successive Crassinucellate Simpson, 1981 Philydraceae Helmholtzia Glandular Successive Crassinucellate Hamann, 1966 Orthothylax Glandular Successive Crassinucellate Hamann, 1966 Philydrella Glandular Successive Crassinucellate Hamann, 1966 Philydrum Glandular Successive Crassinucellate Hamann, 1966 Pontederiaceae Eichhornia Amoeboid Successive Crassinucellate Banerji and Gangulee, 1937; Schurhoff, 1922 Monochoria Amoeboid Successive Crassinucellate Banerji and Haldar, 1942 Sparganiaceae Sparganium Amoeboid Successive Crassinucellate Dahlgren and Clifford, 1982 Taccaceae Schizocapsa Glandular Simultaneous Crassinucellate HAkansson, 1921 Tecophilaeaceae Cyanella Glandular Simultaneous Crassinucellate De Vos, 1950 Odontostomum Glandular Simultaneous Crassinucellate Cave, 1952 Typhaceae Typha Amoeboid Successive Crassinucellate Dahlgren and Clifford, 1982 Velloziaceae Vellozia Glandular Successive Tenuinucellate Schnarf, 1931; Stenar, 1925 (Pseudocrassi- nucellate) genera-Lanaria, Lophiola, and Pauridia- Lachnanthes S. Elliott, the subject of this which have been placed in the Haemodoraceae study, is a monotypic genus in the tribe Hae- in many recent classifications, differ from all modoreae consisting of the species L. carolin- investigated Haemodoraceae (sensu Simpson, iana (Lam.) Dandy [synonomous with L. car- in press b) in one or more major embryological oliana (Lam.) Dandy and L. tinctoria Elliott; features (Table 1). These embryological differ- see Robertson, 1976]. The genus is the only ences, along with differences in anatomy- North American member of the Haemodor- Schulze, 1893; Ambrose, 1980,1985; Simpson aceae north of Mexico and occurs in Nova and Dickison, 1981-and palynology-Erdt- Scotia, the coastal plain of eastern and south- man, 1966; Simpson, 1983; Zavada, Xu, and eastern U.S., and Cuba. Lachnanthes was first Edwards, 1983-argue firmly for the place- placed in the Haemodoraceae by Lindley (1830) ment of these three genera in other, more dis- and has remained in the tribe Haemodoreae tantly related families (see Simpson, in press (= Euhaemodoreae) since the definitive treat- b).] ment of the family by Bentham and Hooker 1396 AMERICAN JOURNAL OF BOTANY [Vol. 75 pr en ppc spC pint o ml t~~~P 1~~ en ~ 2~ cp t~~~~~~~~~~~~~~~~~ e n e a ept 13s Sn an 8~~ ~ ~~~~~~~~~~~~~~~~~ nc ------ 8 ~~~10 1 2 1 3 4 516 Fig. 1-16. Anther, ovule, and embryo development. 1. Portion of anthercross section, early developmentalstage. Note outerprotoderm (pr) and subexterior primary parietal cell (ppc),the latterwith some cellshaving divided periclinally into an endotheciallayer (en) and secondaryparietal cell layer(spc). x 260. 2. Laterdevelopmental stage. Note endothecial layer (en) and recent periclinaldivisions of secondaryparietal cell (spc) layer, formingthe primarymiddle layer (pml) and primarytapetal layer (pt). x 260. 3. Laterstage. Note primarymiddle layer (pml) and recent divisions of primary September 1988] SIMPSON-EMBRYOLOGY OF LACHNANTHES 1397 (1883). Although no controversy exits is to the either a Leitz Wetzlar, Zeiss Photomicroscope, familial
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