The genus Rumex includes hermaphroditic, polygamous, gynodioecious, monoecious, and dioecious species, with the dioecious species being represented by different sex-determining mechanisms and sex-chromosome systems. Therefore, this genus represents an exceptional case study to test several hypotheses concerning the evolution of both mating systems and the genetic control of sex determination in plants. Here, we compare nuclear intergenic transcribed spacers and chloroplast intergenic sequences of 31 species of Rumex. Our phylogenetic analysis supports a systematic classification of the genus, which differs from that currently accepted. In contrast to the current view, this new phylogeny suggests a common origin for all Eurasian and American dioecious species of Rumex, with gynodioecy as an intermediate state on the way to dioecy. Our results support the contention that sex determination based on the balance between the number of X chromosomes and the number of autosomes (X/A balance) has evolved secondarily from male-determining Y mechanisms and that multiple sex-chromosome systems, XX/XY1Y2, were derived twice from an XX/XY system. The resulting phylogeny is consistent with a classification of Rumex species according to their basic chromosome number, implying that the evolution of Rumex species might have followed a process of chromosomal reduction from x = 10 toward x = 7 through intermediate stages (x = 9 and x = 8).
Super high-molecular-weight (SHMW) asphaltene aggregates, hereafter called asphaltene clusters, have been observed for the first-time using gel permeation chromatography with inductively coupled plasma mass spectrometry (GPC-ICP HR MS). Presumably these clusters are composed by nanoaggregates joined by physical interactions such as polar and dispersion forces. These clusters were observed for three asphaltenes samples (Hamaca, Cerro Negro and Boscan) and in their corresponding subfractions, A1 (insoluble in toluene) and A2 (soluble in toluene). Under the present conditions, dilution experiments showed no significant change of these profiles suggesting that these clusters, as well as the other lower molecular weight (LMW) aggregates and molecules are not in equilibrium with each other and behave as independent units. This is supported by the presence of trapped compounds within asphaltenes (TC) which are released after the p-nitrophenol treatment. A temperature cycle was implemented to monitor possible changes of GPC-ICP HR MS profiles with temperature. Here Boscan samples were heated from 25 to 200 °C, left standing by for 24 h and then cooled back to 25 °C before measurement. Whereas we found small changes for Boscan asphaltenes (As-Bo), vast changes in the profiles were detected for A1-Bo and A2-Bo subfractions, and these disclose interchange of material between the cluster and the other components of the solution. These results are discussed in terms of different arrays that may result when asphaltenes are separated in the two subfractions, and nanoaggregates are relocated within the cluster before and after heating. Preliminary molecular dynamics calculation afforded intensity-size distribution coherent with the usual experimental GPC chromatography profiles.
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