Cdc48p from Saccharomyces cerevisiae and its highly conserved mammalian homologue VCP (valosin-containing protein) are ATPases with essential functions in cell division and homotypic fusion of endoplasmic reticulum vesicles. Both are mainly attached to the endoplasmic reticulum, but relocalize in a cell cycle-dependent manner: Cdc48p enters the nucleus during late G1; VCP aggregates at the centrosome during mitosis. The nuclear import signal sequence of Cdc48p was localized near the amino terminus and its function demonstrated by mutagenesis. The nuclear import is regulated by a cell cycledependent phosphorylation of a tyrosine residue near the carboxy terminus. Two-hybrid studies indicate that the phosphorylation results in a conformational change of the protein, exposing the nuclear import signal sequence previously masked by a stretch of acidic residues.
Despite intensive morphological, chemical and cladistic studies on Caryophyllidae, the circumscription of this subclass and the interfamilial relationships are still under discussion. Using comparative sequencing of the chloroplast matK gene, hypotheses of relationships between the carnivorous Droseraceae, Nepenthaceae and Dioncophyllaceae and ten other families of the Caryophyllidae s.l. were tested and compared with previously published cladograms based on rbcL, 18S rDNA and ORF2280 sequences. Parsimony analyses indicate two well‐differentiated clades. One strongly supported clade comprises the carnivorous families Droseraceae and Nepenthaceae, along with its close relatives Dioncophyllaceae and Ancistrocladaceae. The second clade is restricted to the Polygonaceae, Plumbaginaceae, Tamaricaceae and Frankeniaceae. The Simmondsiaceae are more closely related to Caryophyllales and are at the base of the remaining taxa. Results of this analysis suggest that carnivory within Caryophyllidae s.l. has a monophyletic origin and, with the exception of Triphyophyllum, this syndrome was lost in the taxa of Dioncophyllaceae and Ancistrocladaceae. The exclusion of Drosophyllum from Droseraceae suggests no close relationship with this family. Finally, the data support a sister group relationship between the Plumbaginaceae and Polygonaceae and the Frankeniaceae and Tamaricaceae. An extensive survey of the rpl2 intron via PCR amplification indicates that the intron is absent from chloroplast genomes of Droseraceae and all taxa of Caryophyllales, but is present in Drosophyllum. Consequently, there is evidence for a multiple loss of the intron and strong support that Drosophyllum has affinities outside the Droseraceae. Our sequence data corroborate many aspects of recent cladistic analyses based predominantly on rbcL sequences. This study shows that matK sequences are useful for'phylogenetic inference among closely related members of Caryophyllidae.
Molecular systematics demonstrate that carnivorous plants have evolved at least ten times independently, in five orders, 12 families, and 19 genera of angiosperms. Carnivory has arisen once in Nepenthales (a segregate of Caryophyllales), once in Oxalidales, twice in Ericales, and three times each in Lamiales and Poales. Estimated crown ages of these ten lineages range from 1.9 to 81 million years (Mya), with the youngest three lineages (1.9 – 2.6 Mya) being all single genera of Poales, and all involving one or two carnivorous species in an otherwise noncarnivorous group. We now understand the evolution of carnivorous plants based on knowing when and (often) where they diverged from specific noncarnivorous ancestors; inferring which traits were gained, which were retained, and which of the latter may have been crucial preadaptations for carnivory; and identifying the evolutionary drivers of carnivory by evaluating the ecological differences between carnivorous plants and their noncarnivorous relatives.
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