Redalyc.Placentation Patterns and Seed Number in Fruits of South American Solanum Subgen. Leptostemonum (Solanaceae) Species
Total Page:16
File Type:pdf, Size:1020Kb
Darwiniana ISSN: 0011-6793 [email protected] Instituto de Botánica Darwinion Argentina Chiarini, Franco E.; Barboza, Gloria E. Placentation patterns and seed number in fruits of South American Solanum subgen. Leptostemonum (Solanaceae) species Darwiniana, vol. 45, núm. 2, diciembre, 2007, pp. 163-174 Instituto de Botánica Darwinion Buenos Aires, Argentina Available in: http://www.redalyc.org/articulo.oa?id=66945202 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative DARWINIANA 45(2): 163-174. 2007 ISSN 0011-6793 PLACENTATION PATTERNS AND SEED NUMBER IN FRUITS OF SOUTH AMERICAN SOLANUM SUBGEN. LEPTOSTEMONUM (SOLANACEAE) SPECIES Franco E. Chiarini & Gloria E. Barboza Instituto Multidisciplinario de Biología Vegetal (CONICET-UNC), casilla de correo 495, 5000 Córdoba, Argentina; [email protected] (author for correspondence). Abstract. Chiarini, F. E. & G. E. Barboza. 2007. Placentation patterns and seed number in fruits of South American Solanum subgen. Leptostemonum (Solanaceae) species. Darwiniana 45(2): 163-174. Thirty-seven South American species representing seven sections of Solanum subgen. Leptoste- monum were analyzed. Andromonoecious as well as hermaphrodite species were considered. The length and width of their fruits was measured, and the number of seeds per fruit was counted. Medial cross sections of fresh, ripe fruits were observed with a stereoscopic microscope and illustrated. Six placentation patterns and three types of seeds were described. A relationship among seed number, fruit size, fruit color and sexual system was detected by means of statistical analysis. These results suggest that andromonoecy affects fruit size and placentation patterns, in order to contain a higher number of seeds per fruit. Keywords. Andromonoecy, Leptostemonum, placentation, seed number, Solanum, South America. Resumen. Chiarini, F. E. & G. E. Barboza. 2007. Patrones de placentación y número de semillas en frutos de espe- cies sudamericanas de Solanum subgen. Leptostemonum (Solanaceae). Darwiniana 45(2): 163-174. Se analizaron 37 especies sudamericanas pertenecientes a siete secciones de Solanum subgen. Lep- tostemonum. Fueron consideradas tanto especies andromonoicas como hermafroditas. Se midió el ancho y el largo de sus frutos y se contó el número de semillas por fruto. Se estudiaron bajo lupa y se ilustraron cortes por la zona media de frutos frescos maduros. Se describieron seis patrones de pla- centación y tres tipos de semillas. Mediante análisis estadístico, se detectó una relación entre el número de semillas, el tamaño del fruto, el color del fruto y el sistema sexual. Estos resultados sug- ieren que la andromonoecia afecta al tamaño del fruto y su patrón de placentación, para contener un mayor número de semillas. Palabras clave. Andromonoecia, Leptostemonum, número de semillas, placentación, Solanum, Sud- américa. INTRODUCTION “Lulo”), and also weeds (S. viarum Dunal, “Trop- ical soda apple”; S. elaeagnifolium Cav., “Silver- Solanum L. is one of the largest genera among leaf nightshade”; S. sisymbriifolium Lam., “Sticky Angiosperms, including about 1100-1400 species nightshade” or “Wild tomato”; S. rostratum Dunal, (Nee, 1999; Hunziker, 2001; Bohs, 2005) that “Buffalo bur”, “Bull thistle” or “Texas thistle”). grow in all kinds of habitats. The largest subgenus Several species of Solanum subgen. Leptoste- within Solanum is Leptostemonum (Dunal) Bitter, monum have been studied due to their andromo- with ca. 450 species (Nee, 1999). This subgenus, noecy or functional dioecy (Wakhloo, 1975a, b; with nine sections in South America, includes edi- Anderson, 1979; Dulberger et al., 1981; Coleman ble species of economic importance (S. melongena & Coleman, 1982; Solomon, 1987; Anderson & L., “Eggplant”; S. sessiliflorum Dunal, “Cocona” Symon, 1989; Diggle, 1991, 1993). The andromo- or “Cubiu”; S. quitoense Lam., “Naranjilla” or noecy has been linked with fruit type, and a corre- Original recibido el 10 de agosto de 2007; aceptado el 2 de noviembre de 2007 163 DARWINIANA 45(2) 163-174. 2007 lation between fruit size and the degree of Samples of at least 10 fruits per species (of dif- andromonoecy has been proposed for some ferent individuals, when possible) were taken. The species (Whalen & Costich, 1986). Recently, in a length and width of each fruit were measured with study that embraced five species of Solanum sect. a Vernier calliper, and the number of seeds was Acanthophora and eight of sect. Lasiocarpa, Miller counted. Averages of these measurements are & Diggle (2007) demonstrated that fruit size is detailed in Table 1. Medial cross sections of fresh, positively correlated with the proportion of stami- ripe fruits were observed in stereoscopic micro- nate flowers produced within inflorescences. scope preparations. Placentation patterns are based These authors concluded that andromonoecy is a on these observations and are presented as sketch- mechanism to regulate allocation to female func- es. Data about the sexual system of the species tion (i.e., fruit production) independently of allo- studied were taken from the literature, and from cation to male function. our own observations in the field (Table 1). Despite these suggestive previous works, Kruskal-Wallis tests were performed in order to detailed studies on fruit structure of subgenus Lep- detect: a) differences in fruit size and seed number tostemonum are rare. Only some scattered species according to epidermis colours, b) differences in have been studied (Miller, 1969; Dave et al., 1979; fruit size among andromonoecious and hermaph- Dottori & Cosa, 1999, 2003), but a comprehensive rodite species, and c) differences in seed number analysis of the subgenus as a natural group is lack- among andromonoecious and hermaphrodite ing. With regard to the placentation patterns, only species. A linear regression was also conducted to general surveys for Solanaceae (Nee, 1986), and find out whether there is a relationship between for the genus Solanum (Symon, 1987) are avail- fruit size and seeds number. Infostat (2001) was able. This is unfortunate, since the contributions of used for the statistical analysis. morphology and anatomy to systematics are wide- ly known, so a study considering any fruit struc- ture would be valuable for Solanum subgen. Lep- RESULTS tostemonum. The present work attempts to pro- vide morpho-anatomical information in order for it The mean size, color, placentation pattern of the to be used systematically, and as a contribution to fruits, and type and mean number of seeds for each understand the relationship between function and species are summarized in Table 1. Most of the taxa structure in South American species of Solanum have spherical fruits (i.e. width = length = diame- subgen. Leptostemonum. Our proposal is that pla- ter), with some exceptions like S. hieronymi, which centation and seed number are influenced by the has depressed spherical fruits, S. tenuispinum or S. sexual system. aridum, with ovoid fruits, and S. mammosum with its anomalous, 3-5 basally lobed fruit. There are only two species with a hairy pericarp MATERIALS AND METHODS in mature fruits. In S. lycocarpum the hairs are stalked-stellate, while in S. robustum they are sim- We analyzed as many species as possible, try- ple, non-glandular and 1-3-celled. At the early ing to study at least one species of each section. stages, S. quitoense and S. asperolanatum have a pu- Thirty seven wild species representing seven sec- bescent pericarp that turns glabrous at maturity. The tions of Solanum subgenus Leptostemonum were remaining species analyzed are always glabrous. sampled (see appendix 1). Mature fruits are of a single color. They can be Several phylogenetic approaches have being red, orange or orange-red (e.g. S. capsicoides, S. carried out by different authors (Bohs, 2005; Levin sisymbriifolium), yellow (most of the species), et al., 2005, 2006) in which informal infrageneric black or brownish (S. mortonii), dark green (S. grouping (clades) have been defined within bolivianum), greenish yellow (S. platense) or Solanum, and probably will be given a formal sta- greenish grey (S. comptum, S. euacanthum). In tus (Nee et al., 2006). Meanwhile, we have hairy-fruited species, the color is given by the arranged the species according to the last conven- hairs. The fruit of S. lycocarpum is greyish due to tional taxonomic treatment (Nee, 1999) (Table 1). its hairs, but the epicarp underneath them is green- 164 F. CHIARINI & G. BARBOZA. Placentation patterns in Solanum subgen. Leptostemonum Table 1. Macroscopic and anatomical fruit features of the 37 species of Solanum subgen. Leptostemonum studied. Diameter is given when fruit is spherical, and length and width are given when fruit has another shape. Abbreviations: A, andromonoecious species; H, hermaphrodite species; NS, mean number of seeds per fruit. Placen- Fruit shape Sexual Species tation NS Seed Type Fruit Color and size (long. System Pattern x lat., cm) Sect. Acanthophora S. aculeatissimum A 3 117.0 bulky yellow 1.84 S. aenictum A 4 180.2 bulky yellow 2.75 S. atropurpureum H 3 99.2 winged red 1.45 S. capsicoides A 4 298.5 winged red 3.14 S. incarceratum H 3 50.0 winged greenish yellow 1.35 S. mammosum A 3 300.0 bulky yellow 7 x 4 S. palinacanthum A 3 250.0 bulky yellow 3.9 S. platense A 3 80.0 winged greenish yellow 2 S. tenuispinum H 3 81.3 winged yellow 1.37 x1.22 S. viarum A 4 369.4 bulky yellow 2.23 Sect. Crinitum hairs greyish, epidermis S. lycocarpum A 5 600.0 bulky 10.00 greenish S. urticans A 5 431.0 bulky yellow 4.067 Sect. Erythrotrichum hairs greyish, epidermis S. robustum A 5 326.0 bulky 1.8 yellow Sect. Herposolanum S. alternatopinnatum A 1 150.0 bulky yellow 4.2 Sect. Lasiocarpa S. quitoense A 5 1444 bulky yellow 4 Sect. Melongena S. aridum H 1 63.7 bulky yellow 1.567 x 1.1 S.