The complete chloroplast genome sequencing analysis revealed an unusual IRs reduction in three species of subfamily Zygophylloideae.

Tetraena mongolica, Zygophyllum xanthoxylon, and Z. fabago are three typical dryland plants with important ecological values in subfamily Zygophylloideae of Zygophyllaceae. Studies on the chloroplast genomes of them are favorable for understanding the diversity and phylogeny of Zygophyllaceae. Here, we sequenced and assembled the whole chloroplast genomes of T. mongolica, Z. xanthoxylon, and Z. fabago, and performed comparative genomic and phylogenetic analysis. The total size, structure, gene content and orders of these three chloroplast genomes were similar, and the three chloroplast genomes exhibited a typical quadripartite structure with a large single-copy region (LSC; 79,696-80,291 bp), a small single-copy region (SSC; 16,462-17,162 bp), and two inverted repeats (IRs; 4,288-4,413 bp). A total of 107 unique genes were identified from the three chloroplast genomes, including 70 protein-coding genes, 33 tRNAs, and 4 rRNAs. Compared with other angiosperms, the three chloroplast genomes were significantly reduced in overall length due to an unusual 16-24 kb shrinkage of IR regions and loss of the 11 genes which encoded subunits of NADH dehydrogenase. Genome-wide comparisons revealed similarities and variations between the three species and others. Phylogenetic analysis based on the three chloroplast genomes supported the opinion that Zygophyllaceae belonged to Zygophyllales in Fabids, and Z. xanthoxylon and Z. fabago belonged to Zygophyllum. The genome-wide comparisons revealed the similarity and variations between the chloroplast genomes of the three Zygophylloideae species and other plant species. This study provides a valuable molecular biology evidence for further studies of phylogenetic status of Zygophyllaceae.

Niche conservatism and elevated diversification shape species diversity in drylands: evidence from Zygophyllaceae.

The integrated contributions of climate and macroevolutionary processes to global patterns of species diversity are still controversial in spite of a long history of studies. The niche conservatism hypothesis and the net diversification rate hypothesis have gained wide attention in recent literature. Many studies have tested these two hypotheses for woody species in humid forests; however, the determinants of species diversity patterns for arid-adapted plants remain largely ignored. Here, using a molecular phylogeny and the global distributions of Zygophyllaceae, a typical arid-adapted plant family, we assessed the effects of contemporary climate and net diversification rates on species diversity patterns in drylands. We found the variables representing water availability to be the best predictors for Zygophyllaceae diversity. Specifically, Zygophyllaceae species diversity significantly decreased with the increase in water availability, probably owing to phylogenetic conservatism of water-related niches. The net diversification rates of Zygophyllaceae accelerated sharply in the recent 10 Myr, coinciding roughly with the period of global aridification. The species diversity of Zygophyllaceae significantly increased with the increase in mean net diversification rates per geographical unit, especially in the Old World, supporting the net diversification rate hypothesis. Our study provides a case exploring climatic and evolutionary mechanisms of dryland species diversity patterns, and suggests that the conservatism in water-related niches and elevated net diversification rates in drylands may have jointly determined the global patterns of dryland species diversity.

Insights into the historical assembly of global dryland floras: the diversification of Zygophyllaceae.

BACKGROUND: Drylands cover nearly 41% of Earth's land surface and face a high risk of degradation worldwide. However, the actual timeframe during which dryland floras rose on a global scale remains unknown. Zygophyllaceae, an important characteristic component of dryland floras worldwide, offers an ideal model group to investigate the diversification of dryland floras. Here, we used an integration of the phylogenetic, molecular dating, biogeographic, and diversification methods to investigate the timing and patterns of lineage accumulation for Zygophyllaceae overall and regionally. We then incorporated the data from other dominant components of dryland floras in different continents to investigate the historical construction of dryland floras on a global scale. RESULTS: We provide the most comprehensive phylogenetic tree for Zygophyllaceae so far based on four plastid and nuclear markers. Detailed analyses indicate that Zygophyllaceae colonized Africa, Asia, Australia, and the New World at different periods, sometimes multiple times, but Zygophyllaceae lineages in the four regions all experienced a rapid accumulation beginning at the mid-late Miocene (~ 15-10 Ma). Other eleven essential elements of dryland floras become differentiated at the same time. CONCLUSIONS: Our results suggest that the rise of global dryland floras is near-synchronous and began at the mid-late Miocene, possibly resulting from the mid-Miocene global cooling and regional orogenetic and climate changes. The mid-late Miocene is an essential period for the assembly and evolution of global dryland floras.

In vitro cytotoxic activity of five commercial samples of Tribulus terrestris Linn in Espírito Santo (Brazil)

ABSTRACT The objective was to investigate the total saponin and protodioscin concentrations and the cytotoxicity in vitro, of five samples of the plant Tribulus terrestris, commercially available in the metropolitan region of Vitória - Espirito Santo, Brazil, and to compare them with the aqueous extract of the plant. The chromatographic profile and quantification of protodioscin in commercial samples and plant extract were evaluated by LC-MS/MS. The percentage of total saponins were determined by the colorimetric method. Extracts and protodioscin cytotoxicity were analyzed by the MTT assay in three cell lineages: fibroblasts (L929), ovarian cancer (Ovcar3) and murine hepatoma (Hepa1c1c7). All extracts displayed high levels of total saponins (207.2 to 780.3 mg g-1 of dry extract). The chromatographic profile revealed a wide diversity of compounds, and the saponin protodioscin was detected in only two extracts. One extract displayed high cytotoxicity, with IC50 values of 157.0, 38.2 and 7.4 µg mL-1 for the Ovcar3, Hepa1c1c7 and L929 cell lines, respectively. The other extracts displayed cytotoxic effects only at concentrations equal to or greater than 125.0 µg mL-1. Surprisingly, the most cytotoxic extract displayed the highest protodioscin concentration. Therefore, it is suggested that these products be marketed with caution, and followed-up by a certified healthcare professional.

Evolution of leaf anatomy in arid environments - A case study in southern African Tetraena and Roepera (Zygophyllaceae).

The dry biomes of southern Africa (Desert, Nama Karoo and Succulent Karoo) are home to a rich and diverse xerophytic flora. This flora includes two morphologically diverse clades of Zygophyllaceae, Tetraena and Roepera (Zygophylloideae), which inhabit some of the most arid habitats in the region. Using a plastid phylogeny of Zygophylloideae we assess whether the evolution of putatively adaptive traits (leaf shape, vasculature, mode of water storage and photosynthetic type: C3 versus C4) coincides with the successful colonisation of environments with different drought regimes within southern Africa. Our results show general niche conservatism within arid habitats in Tetraena, but niche shifts from arid to more mesic biomes with longer and/or cooler growing season (Fynbos and Thicket) in Roepera. However, these distinct broad-scale biogeographical patterns are not reflected in leaf anatomy, which seems to vary at more local scales. We observed considerable variability and multiple convergences to similar leaf anatomies in both genera, including shifts between "all cell succulence" leaf types and leaf types with distinct chlorenchyma and hydrenchyma. Our survey of C4 photosynthesis in the Zygophylloideae showed that the C4 pathway is restricted to Tetraena simplex, which also having an annual life history and a widespread distribution, is rather atypical for this group.

Evolution of Asian Interior Arid-Zone Biota: Evidence from the Diversification of Asian Zygophyllum (Zygophyllaceae).

The Asian interior arid zone is the largest desert landform system in the Northern Hemisphere, and has high biodiversity. Little is currently known about the evolutionary history of its biota. In this study, we used Zygophyllum, an important and characteristic component of the Asian interior arid zone, to provide new insights into the evolution of this biota. By greatly enlarged taxon sampling, we present the phylogenetic analysis of Asian Zygophyllum based on two plastid and one nuclear markers. Our phylogenetic analyses indicate that Asian Zygophyllum and Sarcozygium form a clade and Sarcozygium is further embedded within the shrub subclade. An integration of phylogenetic, biogeographic, and molecular dating methods indicates that Zygophyllum successfully colonized the Asian interior from Africa in the early Oligocene, and Asian Zygophyllum became differentiated in the early Miocene and underwent a burst of diversification in the late Miocene associated with the expansion of Asian interior arid lands due to orogenetic and climatic changes. Combining diversification patterns of other important components of the Asian interior arid zone, we propose a multi-stage evolution model for this biota: the late Eocene-early Oligocene origin, the early Miocene expansion, and the middle-late Miocene rapid expansion to the whole Asian interior arid zone. This study also demonstrates that, for Zygophyllum and perhaps other arid-adapted organisms, arid biomes are evolutionary cradles of diversity.

Genetic diversity among Zygophyllum (Zygophyllaceae) populations based on RAPD analysis.

Zygophyllum species are succulent plants that are drought resistant and/or salt tolerant, growing under severe, dry climatic conditions. Despite their importance and abundance in the Mediterranean and Middle East regions, there is little information concerning molecular variations among species of this genus. Genetic diversity was assessed, using RAPD primers, of 12 populations of Z. coccineum, Z. album and Z. aegyptium collected from various locations in Egypt and Saudi Arabia. Yong leaves were used for DNA extraction. Genetic distances were calculated using Nei's method. A dendrogram was constructed based on the similarity data matrix by unweighted pair group method using arithmetic averages cluster analysis. Analysis with RAPD markers revealed genetic variation between and within populations of Zygophyllum. Zygophyllum coccineum showed higher levels of genetic variation and more unique alleles than the other species.

Comparative exine ontogeny in some members of the family Zygophyllaceae sensu lato.

Exine ontogeny is studied in five taxa of the family Zygophyllaceae sensu lato (Peganum harmala L., Zygophyllum album L., Fagonia cretica L., Tribulus terrestris L., and Nitraria retusa [Forsk.] Asch.). In the beginning of the tetrad stage, the plasmalemma is attached to the callose wall, except in T. terrestris, where it describes crests and hollows. The primexine matrix is fibrillar, bilayered in T. terrestris and unilayered in the other taxa. In all species except P. harmala, the procolumellae are heterogeneous with clear zones and they become compact later. In T. terrestris, they are built on the crests. In Z. album and T. terrestris, a primordial nexinic lamella is set up. It is tripartite with a white line seen at some levels; on its external leaflet, the foot layer is observed, and on its internal leaflet, there is the endexine with numerous lamellae. This white line disappears often in the mature exine. In T. terrestris, there is a thick nexine that is coarsely lamellate inside. In the aperture zone, the columellae are lacking, the tectum and the foot layer get thinner; they unite and form the apertural membrane with the external part of the endexine. There is a granulolamellar endexinic zone well developed in P. harmala, whereas it is threelayered and weakly developed in T. terrestris.