To date molecular data have not revealed the exact phylogenetic position of Balanophoraceae in relation to hemiparasitic Santalales. To elucidate the phylogeny of Santalales and the position of Balanophoraceae, three plastid genes (matK, rbcL, accD), three nuclear genes (SSU and LSU rDNA and RPB2) and one mitochondrial gene (matR) from 197 Santalales samples (including 11 Balanophoraceae species) were analyzed with parsimony, maximum likelihood and Bayesian inference methods. Our results demonstrate that Balanophoraceae is composed of two well‐supported clades: a relatively slow‐evolving one including Dactylantus, Hachettea, and Mystropetalon (Mystropetalaceae) and an extremely fast‐evolving one composed of the remaining Balanophoraceae s.str. Support for monophyly of the two clades was low, thus it appears holoparasitism has arisen twice independently in Santalales. These two clades appeared during a time of great change in the order (ca. 100 Ma) when several major evolutionary innovations emerged, e.g., the root hemiparasites of Santalaceae s.l., the first aerial parasites (Misodendraceae), herbaceous root parasites (Schoepfiaceae), root parasitic Loranthaceae (the ancestors of aerial parasitic mistletoes), as well as the holoparasites in Balanophoraceae and Mystropetalaceae.
Citrus huanglongbing (HLB), previously called greening, is a serious citrus disease in Asia, eastern and southern Africa. It is caused by Candidatus Liberibacter asiaticus (Las), a phloem-limited, nonculturable bacterium transmitted by the Asian citrus psyllid ( Diaphorina citri ) in Asia. A PCR-based assay was developed for monitoring Las in vector psyllids using a rapid DNA extraction from psyllid bodies and PCR amplification. The entire procedure for Las detection in psyllids can be completed within 5 h. Using this method, Las can be accurately detected in psyllid adults as well as nymphs in different instar stages. The assay is sensitive enough for Las detection in single-psyllid extract from adult, fifth, fourth and third instars. In a transovarial transmission experiment, Las was not detected in eggs or in offspring produced by Las-carrying psyllid females. In a retention test, the Las-carrying psyllids remained Las-positive for 12 weeks after they were moved to common jasmine orange, a Las-immune plant. From these experimental results it was concluded that Las persists in the Asian citrus psyllid vector, but is not transovarially transmitted by the vector. These data help in understanding epidemiological characteristics of Las and psyllids in citrus HLB.
Plastid genomes (plastomes) vary enormously in size and gene content among the many lineages of nonphotosynthetic plants, but key lineages remain unexplored. We therefore investigated plastome sequence and expression in the holoparasitic and morphologically bizarre Balanophoraceae. The two Balanophora plastomes examined are remarkable, exhibiting features rarely if ever seen before in plastomes or in any other genomes. At 15.5 kb in size and with only 19 genes, they are among the most reduced plastomes known. They have no tRNA genes for protein synthesis, a trait found in only three other plastid lineages, and thus Balanophora plastids must import all tRNAs needed for translation. Balanophora plastomes are exceptionally compact, with numerous overlapping genes, highly reduced spacers, loss of all cis-spliced introns, and shrunken protein genes. With A+T contents of 87.8% and 88.4%, the Balanophora genomes are the most AT-rich genomes known save for a single mitochondrial genome that is merely bloated with AT-rich spacer DNA. Most plastid protein genes in Balanophora consist of ≥90% AT, with several between 95% and 98% AT, resulting in the most biased codon usage in any genome described to date. A potential consequence of its radical compositional evolution is the novel genetic code used by Balanophora plastids, in which TAG has been reassigned from stop to tryptophan. Despite its many exceptional properties, the Balanophora plastome must be functional because all examined genes are transcribed, its only intron is correctly trans-spliced, and its protein genes, although highly divergent, are evolving under various degrees of selective constraint.
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