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Aspects of Floral Development in Philodendron Grandifolium and Philodendron Megalophyllum (Araceae)

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Aspects of Floral Development in Philodendron Grandifolium and Philodendron Megalophyllum (Araceae) Int. J. Plant Sci. 162(1):47–57. 2001. ᭧ 2001 by The University of Chicago. All rights reserved. 1058-5893/2001/16201-0004$03.00 ASPECTS OF FLORAL DEVELOPMENT IN PHILODENDRON GRANDIFOLIUM AND PHILODENDRON MEGALOPHYLLUM (ARACEAE) Denis Barabe´ 1 and Christian Lacroix Institut de Recherche en Biologie Ve´ge´tale, Jardin Botanique de Montre´al, 4101 Sherbrooke Est, Montre´al, Que´bec H1X 2B2, Canada; and Department of Biology, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island C1A 4P3, Canada This study deals specifically with floral organogenesis and the development of the inflorescence of Philo- dendron grandifolium and Philodendron megalophyllum. Pistillate flowers are initiated on the lower portion of the inflorescence, and the staminate flowers are initiated on the distal portion. The presence of extracellular calcium oxalate is observed on the surface of immature stamens of P. megalophyllum. An intermediate zone consisting of sterile male flowers and bisexual flowers with fused or free carpels and staminodes is also present on the inflorescences. This zone is located between the male and female floral zones. In general, the portion of bisexual flowers facing the male zone forms staminodes, and the portion facing the female zone develops an incomplete gynoecium with few carpels. In P. grandifolium, the incomplete separation of some staminodes from the gynoecial portion of the whorl shows that they belong to the same whorl as the carpels. The bisexual flowers of P. grandifolium and P. megalophyllum are believed to be a case of homeosis in which carpels have been replaced by sterile stamens on the same whorl. Keywords: flower, inflorescence, homeosis, development. Introduction on the inflorescence. They are located at the junction between the female portion and the male portion of the inflorescence. The family Araceae comprises 105 genera and more than Developmental studies have shown that carpels and stami- 3300 species (Mayo et al. 1997). Members of this family are nodes are inserted on the same whorl on bisexual flowers (e.g., characterized by two main types of inflorescences: (1) those Mayo 1986; Boubes and Barabe´ 1996; Barabe´ and Lacroix with bisexual flowers, as represented in the genus Anthurium, 1999, 2000). This phenomenon can be explained as the re- and (2) those with unisexual flowers, as represented in the placement of carpels by staminodes, and it represents an ex- genus Philodendron. Inflorescences of the Philodendron type ceptional case of homeosis (e.g., Barabe´ and Lacroix 1999; consist of female flowers located in the lower portion and male Barabe´et al. 2000). flowers in the upper portion. Several species with unisexual In the atypical bisexual flowers of Philodendron, there is a flowers (representing a number of genera) have been investi- lot of variation in the number of carpels replaced by stami- gated by a variety of authors from the perspective of floral nodes among species. For example, in Philodendron solimoe- anatomy and developmental morphology (Engler and Krause sense, an average of 3.2 carpels are replaced by one staminode. 1912; Eckardt 1937; Eyde et al. 1967; Hotta 1971; Barahona In Philodendron fragrantissimum, this number is reduced to Carvajal 1977; Uhlarz 1982, 1986; French 1985a, 1985b, 1.5 and represents a more or less one-to-one shift (Barabe´et 1986a, 1986b; Mayo 1986, 1989; Barabe´ and Forget 1988; al. 2000). The high degree of variability in the mode of ex- Carvell 1989; Buzgo´ 1994; Boubes and Barabe´ 1997). Re- pression of carpel/staminode in homeotic bisexual flowers is cently, Barabe´ and collaborators (Barabe´and Bertrand 1996; not only observed between species belonging to different sub- Barabe´and Lacroix 1999; Barabe´et al. 2000) used the flower genera, such as P. solimoesense (Meconostigma) and P. fra- of the Philodendron type as a model to analyze theoretical grantissimum (Philodendron) but is also observed between spe- aspects of morphogenesis, such as the phenomena of homeosis cies of the same subgenera and even among individuals and morphogenetic gradient. They observed that the nature of belonging to the same species. How can we explain this var- floral organs on different types of flowers depends on the po- iation? To answer this question, comparative developmental sition of the flowers on the inflorescence. In other words, the studies between a greater number of species belonging to the nature of floral organs is independent of the whorl on which same subgenera must be initiated. It is from this perspective it is initiated. that the floral development of Philodendron grandifolium and In the genus Philodendron, bisexual flowers, called “mon- stro¨sen Blu¨ten” by Engler and Krause (1912), are also present Philodendron megalophyllum will allow us to provide new data supporting the morphological nature of bisexual flowers in the genus Philodendron. More precisely, we intend to ad- 1 Author for correspondence; telephone 514-872-1436; fax 514- dress the following questions in the context of the phenomenon 872-3765; e-mail [email protected]. of homeosis: Are there different types of bisexual flowers in Manuscript received June 2000; revised manuscript received August 2000. terms of substitution ratios (i.e., number of carpels replaced 47 48 INTERNATIONAL JOURNAL OF PLANT SCIENCES by number of staminodes or vice versa)? Do carpel and stam- a Cambridge S604 scanning electron microscope (SEM) with inode primordia belonging to the same whorl have a different digital imaging capabilities (SEMICAPS). mode of development than do staminodes and carpels be- longing to sterile male flowers and female flowers, respectively? Is the relationship between the nature of floral organs that will Results be formed and the position of the flower on the inflorescence in P. grandifolium and P. megalophyllum similar to that of other documented cases? Morphology of the Mature Flowers Although there are a few anatomical studies of selected spe- The length of the mature spadices of both species ranges cies of the subgenus Philodendron (Barahona Carvajal 1977; from 10 to 15 cm. Both staminate and pistillate flowers have Mayo 1986, 1989), the mode of development of the flower in no perianth. The staminate flowers occupy the upper portion this subgenus, represented in this study by P. grandifolium and of the inflorescence and take up ca. 65% of the total length P. megalophyllum, remains poorly understood. Consequently, of the inflorescence, whereas the female flowers are located on new morphological and ontogenetic characters related to the the lower portion and occupy ca. 25% of the total length. In development of flowers are needed to complement anatomical the median portion of the inflorescence, there is an interme- studies of subgenus Philodendron species (see, e.g., Mayo diate zone (ca. 10% of the total length) consisting of bisexual 1986, 1989) and to add to our present knowledge of floral flowers and sterile males flowers. types within this group. In both species, modified stomata (water pores) in the sense Developmental studies often reveal particular features that of Vogel (1977) are found on the surface of the apical portion are not visible on fully developed organs. This is particularly of the stamens (fig. 1A,1B) and staminodes on male and sterile the case for secretory structures such as hydathodes, nectaries, male flowers, respectively. The epidermis of stamens and stam- and water pores in the sense of Vogel (1977) and Endress inodes consists of pegged and rigged cells (fig. 1B). (1996). The presence of stomata on stamens and staminodes The bisexual flowers generally consist of carpels and stam- has been observed in different species of Philodendron, such inodes inserted on the same whorl. The portion of the bisexual as Philodendron acutatum (Boubes and Barabe´ 1996) and P. flower facing the male zone consists of staminodes, and the solimoesense (Barabe´and Lacroix 1999). However, there were portion facing the female zone consists of an incomplete gy- no indications concerning the accumulation of calcium oxalate noecium (figs. 1C,5F–5H). On mature flowers, like that rep- crystals on anthers in Philodendron. With this study, we intend resented in figure 1C, it is difficult to determine whether the to present evidence for the presence of extracellular calcium staminode(s) and carpels are inserted on the same whorl. How- oxalate crystals in P. megalophyllum. ever, during the early stages of development, this phenomenon The general aims of this study are (1) to compare the de- is visible (see section on floral development of bisexual flow- velopment of flowers of P. grandifolium and P. megalophyllum ers). The presence of mature atypical organs having both stam- to that of other members of the same subgenus in order to test inodal and pistillate characters (figs. 1C,4H) is a more or less our current assumptions on the phenomenon of homeosis in common feature in the intermediate zone of the inflorescence. the Araceae and (2) to further document the liberation of cal- These organs are characterized by a discontinuous stigmatic cium oxalate crystals by floral organs. surface and staminodal-like tissue. The mature female flowers have a prominent stigmatic sur- Material and Methods face (fig. 1D,1E; fig. 4F,4G). A large quantity of secretions are consistently observed between the papillae on the stigmatic Philodendron grandifolium Brongn. ex Regel and Philoden- surface (fig. 1F). dron megalophyllum Schott belong to the subgenus Philoden- dron, section Philodendron, subsection Philodendron (Croat 1997). Specimens used for this study were collected by D. Development of the Inflorescence Barabe´in French Guiana (Petit-Saut dam) in November 1997 and July 1998 (voucher specimens were deposited at MT: Bar- The inflorescence primordium of Philodendron grandifol- abe´62, Barabe´67). Specimens growing at the Montreal Bo- ium is more or less cylindrical in shape during early stages of tanical Garden were also collected in April 2000 (registration initiation (fig. 2A). The different types of flowers are initiated numbers 2382-92 [P.
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