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Size-Dependent Sex Allocation in a Monoecious Species Sagittaria Pygmaea (Alismataceae)

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Size-Dependent Sex Allocation in a Monoecious Species Sagittaria Pygmaea (Alismataceae) Ann. Bot. Fennici 46: 95–100 ISSN 0003-3847 (print) ISSN 1797-2442 (online) Helsinki 30 April 2009 © Finnish Zoological and Botanical Publishing Board 2009 Size-dependent sex allocation in a monoecious species Sagittaria pygmaea (Alismataceae) Fan Liu1,3, Jin-Ming Chen2 & Qing-Feng Wang1,3,* 1) Hubei Key Laboratory of Wetland Evolution and Ecological Restoration, Wuhan, Hubei 430074, P. R. China (*corresponding author’s e-mail: [email protected]) 2) Laboratory of Plant Systematics and Evolutionary Biology, College of Life Sciences, Wuhan University, Wuhan 430072, Hubei, P. R. China 3) Wuhan Botanical Garden, the Chinese Academy of Sciences, Wuhan, Hubei 430074, P. R. China Received 20 Jan. 2008, revised version received 14 Jan 2009, accepted 12 Mar. 2008 Liu, F., Chen, J. M. & Wang, Q. F. 2009: Size-dependent sex allocation in a monoecious species Sagittaria pygmaea (Alismataceae). — Ann. Bot. Fennici 46: 95–100. Sagittaria species have been reported to display remarkable variation in gender expres- sion. Here, we investigated gender variation in Sagittaria pygmaea, the smallest sized monoecious species in the genus. We used the midvein length as an indicator of plant size and production of male and female fl owers as an indicator of gender variation in a single infl orescence. We counted the total number of infl orescences to assess the effect of infl orescence variation on the gender variation pattern. Our results showed that vari- ation in infl orescence number did not affect gender variation. Male fl ower production increased with increasing plant size, but female fl ower production did not. Plants of S. pygmaea might enhance their paternal reproductive success by increasing the number of male fl owers with increasing plant size. Key words: Alismataceae, monoecious, Sagittaria pygmaea, size-dependent sex allo- cation Introduction The gender of fl owering plants varies widely resource status and investment in female and within and among species (Lloyd & Bawa 1984). male components of reproduction among plants This variation is the product of a complex inter- (Gilbert & Bolker 2003, Sultan 2003a, 2003b, play of genetic and environmental factors (Lloyd Sultan 2005). & Bawa 1984, Sarkissian et al. 2001). Genes In general, when environmental condi- could regulate the production of unisexual versus tions play a role in gender variation in cosex- hermaphroditic fl owers and lead to a wide array ual organisms, sex allocation is considered to of sexual systems (Dorken & Barrett 2003, and be associated with plant size (Policansky 1982, references therein). Klinkhamer et al. 1997, Barrett et al. 1999). Environmental factors play an important role According to several theoretical models con- in sex allocation mainly through their effect on cerned with size-dependent sex allocation, in a 96 Liu et al. • ANN. BOT. FENNICI Vol. 46 plant population the larger individuals should S. pygmaea indicated that with increase in plant be more female than the smaller ones, because size the number of female fl owers on an infl ores- they have more resources to afford the greater cence changed little, while that of male fl owers expense and added costs of fruiting both in varied widely among different individuals in a direct expenditure on current fruit and indirect population. The sex allocation patterns observed consequences for future survival (Charnov 1982, in S. pygmaea do not seem to conform to those Lloyd & Bawa 1984, Klinkhamer et al. 1997, reported in other Sagittaria species. The sex Zhang 2006). In addition, the degree of local allocation patterns in S. pygmaea are still poorly mate competition and increased geitonogamy understood. may increase with size and can be detrimental We investigated the sex allocation patterns for male success for large size plants (Lloyd & in S. pygmaea on four natural populations from Bawa 1984, Klinkhamer et al. 1997). Hence, southeast to southwest China. The plant size large plants will often perform better as females usually changes during the growing season and and worse as males, and so should benefi t by the infl orescence number may change with plant being relatively more female than small plants size throughout the fl owering period. Observa- (Sarkissian et al. 2001). Such gender plasticity tion of variation in infl orescence number in the should be particularly common in monoecious fi eld is often hampered by fl uctuating parameters plants, because the production of separate female including environmental factors such as drought and male fl owers enables greater freedom to and fl ood, which may affect infl orescence pro- respond to specifi c environmental circumstances duction. These hindrances can be partly over- (Sarkissian et al. 2001, and references therein). come by conducting the experiments on plants Sagittaria is a worldwide genus comprising in cultivation. In our study to establish whether of approximately 30 species (Chen 1989). Most infl orescence number varies with plant size Sagittaria species are basically monoecious throughout the fl owering period in S. pygmaea (Bogin 1955), but some species display remark- we included one cultivated population. In the able variation in gender expression (Barrett et al. study we addressed the following questions: (1) 2000, Huang et al. 2002, and references therein). Is there any evidence of size-dependent sex allo- Sagittaria pygmaea is an emergent or submersed cation in S. pygmaea? (2) What is the adaptive aquatic annual in shallow waters along marshes, signifi cance of this size-dependent sex allocation ponds, stream banks and rice fi elds in southern pattern in the species? and southeastern Asia. The species is 7–15 cm high and has 3–5 sequential infl orescences; each infl orescence typically has only 0–2 female and Material and methods 3–8 male fl owers (Chen 1989). The species is dichogamous and the fl owering periods of the Data collection sequential infl orescences do not overlap. Only one or two fl owers on an infl orescence of S. During July 2005 and July and August 2006, we pygmaea bloom in a day and the stigmas become investigated four different populations of S. pyg- non-receptive within eight hours after fl ower- maea in southeastern and southwestern China ing (F. Liu unpubl. data). Due to the remarkable (Table 1). In 2005, we collected 35 corms from gender plasticity in Sagittaria species, several Population ZJ and grew them in Wuhan Botani- studies have been conducted on the sex alloca- cal Garden. In March 2006 ramets were develop- tion in the genus (Sarkissian et al. 2001, Huang ing and were transplanted one month later into et al. 2002). Most of those studies have focused 4 ¥ 18-arranged 20-cm-deep pots placed 0.5 m on sex allocation patterns in which the female apart. fl ower production varied widely with plant size In both natural and cultivated populations, for while male fl ower production changed little or each plant we counted the number of male and was unaffected (e.g. S. latifolia, Sarkissian et female fl owers on each infl orescence and meas- al. 2001; S. trifolia, Huang et al. 2002); how- ured the plant height and midvein length. For ever, our fi eld observations on populations of cultivated populations, we also calculated the ANN. BOT. FENNICI Vol. 46 • Size-dependent sex allocation in Sagittaria pygmaea 97 Table 1. General information on the four sampled natural populations and one cultivated population of Sagittaria pygmaea in China. Population Location Latitude/longitude Habitat Sample size ZJ Zhijang, Hubei Province 30°20´/111°35´ Marsh 22 WYS Wuyishan, Fujian Province 27°33´/117°51´ Paddy fi eld 26 YS Yanshan, Guangxi Province 25°01´/110°19´ Paddy fi eld 9 PB Pingba, Guizhou Province 26°25´/106°16´ Ditch 21 WH Wuhan, Hubei Province 30°29´/114°19´ Cultivated 32 number of infl orescences and the total number of ZJ: r = 0.4555, F1,21 = 5.24, P = 0.0331; WYS: male and female fl owers. We used the midvein r = 0.5808, F1,25 = 12.22, P = 0.0018; YS: r = length of the longest leaf rather than plant weight 0.4558, F1,8 = 1.66, P = 0.2391; PB: r = 0.6239, (Sarkissian et al. 2001) as a measure of plant F1,20 = 12.11, P = 0.0025). size, because it was easy to measure the midvein The number of fl owers per infl orescence in length on natural populations and the midvein the cultivated population displayed similar sex length strongly correlated with the plant height allocation patterns as those in the natural popula- (r = 0.809, F1,146 = 274.54, P < 0.001) and the tions. The cultivated plants produced a total of plant weight (the above ground dry mass) (r = 77 infl orescences and each infl orescence pro- 0.845, F1,146 = 360.75, P < 0.001). duced a single female fl ower. The number of male fl owers was positively correlated to plant size in all infl orescences (r = 0.3332, F1,76 = 9.36, Statistical analyses P = 0.0031). However, throughout the fl owering period, we did not fi nd any correlation between We fi rst performed an analysis of covariance the midvein length (plant size) and the number (ANCOVA) on female and male fl ower pro- of infl orescences on an individual (Fig. 2, r = duction, with plant size as the covariate and 0.0111, F1,31 = 0.00, P = 0.9512). population as a fi xed effect using GLM proce- dure. Thereafter, we compared the relationship between male and female fl ower production and Discussion plant size using correlation analysis. All the analyses were performed using SAS/STAT (SAS Several studies have investigated the relationship Institute 1998). between gender variation and plant size in Sagit- taria. Sagittaria trifolia and S. latifolia exhibited a positive correlation between the female fl ower Results Each of the S.
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