Phylogeny and biogeography of Sagittaria (Alismataceae) revisited: evidence for cryptic diversity and colonization out of South America.

Sagittaria is a genus of ca. 40 species in the aquatic plant family Alismataceae with a nearly global distribution, and a center of diversity in the New World. Two thirds of the known species are native to the Americas, while only a few species are distributed in Africa, Asia and Europe. A previous biogeographic analysis of the genus suggested an African origin for the genus with subsequent dispersal to North America and then to East Asia. Here we expanded the taxon sampling with a focus on the New World taxa and applied species delimitation and biogeographic analyses to revise the knowledge of the phylogeny and evolution of the genus. We obtained largely similar topologies from the chloroplast DNA and nuclear DNA (ITS) data sets. The 74 accessions sampled for our analyses were delimited into 29 species and several cryptic taxa were revealed in widely distributed species. Biogeographic analysis supported basal diversification in South America and subsequent colonization to North America and Asia.

Effects of population size on synchronous display of female and male flowers and reproductive output in two monoecious Sagittaria species.

BACKGROUND: Flowering synchrony and floral sex ratio have the potential to influence the mating opportunities and reproductive success through female function. Here, we examine the variances in synchronous display of female and male function, ratio of male to female flowers per day and subsequently reproductive output in small populations of two monoecious plants, Sagittaria trifolia and Sagittaria graminea. METHODOLOGY/PRINCIPAL FINDING: We created plant populations of size 2, 4, 10 and 20 and recorded the daily number of blooming male and female flowers per plant to determine daily floral display, flowering synchrony index and ratio of male to female flowers per day. We also harvested the fruits, counted the seeds and calculated the number of fruits and seeds per flower to measure reproductive success through female function. There is less overlap in flowering time of female and male function in smaller populations than in larger populations. Most importantly, we found that male-biased floral sex ratio and imbalanced display period of female and male function for individual plant can lead to a population-size-dependent ratio of male to female flowers per day. Increasing ratio of male to female flowers per day was generally associated with a greater percentage of fruit production. CONCLUSIONS/SIGNIFICANCE: Our results highlight the importance of flowering synchrony of female and male function and population-size-dependent ratio of male to female flowers per day for female reproductive success. This finding improves our understanding of a mechanism that reduces reproductive success in small populations.

Phylogeography of a marsh herb Sagittaria trifolia (Alismataceae) in China inferred from cpDNA atpB-rbcL intergenic spacers.

Sagittaria trifolia L. is a perennial, erect herb that is confined to ponds, rice fields, ditches, and freshwater wetlands. Using chloroplast DNA (cpDNA) atpB-rbcL intergenic spacer sequences, we studied the phylogeographic pattern and demographic history of S. trifolia with 108 samples from 42 populations representing the entire geographic range in China. Twenty-seven haplotypes were characterized and two of them were widely distributed in the populations. In the minimum-spanning network, all tip haplotypes were unique to a particular population, while the interior nodes represented widespread haplotypes. Nested clade analysis (NCA) of cpDNA haplotypes indicated that long distance dispersal characterized the post-glacial recolonization of S. trifolia in China. No specific refugia areas were suggested because genetic differentiation was low among the sampled regions and among populations within regions although a large number of the haplotypes were unique to a single population. The present data support that the unique haplotypes in individual population most likely represent recent mutational derivatives after long distance dispersal rather than the relics in refugia. These results for S. trifolia represent the first phylogeographic analysis of a widespread marsh herb in China and support the importance of long distance dispersal events in the post-glacial migrations of plants.

Expression of alpha-expansin and xyloglucan endotransglucosylase/hydrolase genes associated with shoot elongation enhanced by anoxia, ethylene and carbon dioxide in arrowhead (Sagittaria pygmaea Miq.) tubers.

BACKGROUND AND AIMS: Shoot elongation of arrowhead tubers (Sagittaria pygmaea Miq.) is stimulated by anoxia, ethylene and CO2. The aim of this study was to characterize anoxic elongation by comparison with elongation stimulated by ethylene and CO2. METHODS: The effects of the inhibitors aminoethoxyvinylglycine (AVG) as an ethylene biosynthesis inhibitor, 1-methylcyclopropene (1-MCP) as a potent inhibitor of ethylene action, and pyrazol, an inhibitor of alcohol dehydrogenase, on shoot elongation were examined. Moreover, the effects of these gaseous factors on expression of genes possibly involved in modification of cell wall architecture were examined by polymerase chain reaction (PCR) methods. KEY RESULTS AND CONCLUSIONS: In air, promotion by 5% CO2 and 5 microL L-1 ethylene of shoot elongation occurred. At 1% O2, ethylene also stimulated shoot elongation but CO2 did not. Pyrazol inhibited shoot elongation in hypoxia but not in normoxia, suggesting that alcohol fermentation contributes to elongation enhanced by hypoxia. AVG and 1-MCP partially prevented shoot elongation both in normoxia and in hypoxia, but they did not have significant effects in anoxia, suggesting that endogenous ethylene acts as a stimulator of shoot elongation in normoxia and in hypoxia but not in anoxia. Ethylene is not involved in anoxia-enhanced elongation. We cloned four cDNAs (SpEXPA1, 2, 3 and 4) encoding alpha-expansin (EXPA) and five cDNAs (SpXTH1, 2, 3, 4 and 5) encoding xyloglucan endotransglucosylase/hydrolase (XTH) from shoots of arrowhead tubers. The transcript levels of SpEXPA1 and 2 were increased by anoxia and those of SpEXPA2 were increased by 5% CO2. Ethylene slightly elevated the level of SpEXPA4 transcripts. Anoxia enhanced the transcript levels of SpXTH1 and 4; neither ethylene nor CO2 had any effect. CO2 enhanced transcript levels of SpXTH3 and depressed those of SpXTH5. Ethylene decreased transcript levels of SpXTH5. These results suggest that four SpEXPA genes and five SpXTH genes are differently responsive to anoxia, CO2 and ethylene. Enhancement of SpEXPA1 and 2, and SpXTH1 and 4 transcript levels suggests that these gene products are involved in anoxic shoot elongation through modification of cell wall architecture.