Evolution of extreme stomach pH in bilateria inferred from gastric alkalization mechanisms in basal deuterostomes

The stomachs of most vertebrates operate at an acidic pH of 2 generated by the gastric H+/K+-ATPase located in parietal cells. The acidic pH in stomachs of vertebrates is believed to aid digestion and to protect against environmental pathogens. Little attention has been placed on whether acidic gastric pH regulation is a vertebrate character or a deuterostome ancestral trait. Here, we report alkaline conditions up to pH 10.5 in the larval digestive systems of ambulacraria (echinoderm + hemichordate), the closest relative of the chordate. Microelectrode measurements in combination with specific inhibitors for acid-base transporters and ion pumps demonstrated that the gastric alkalization machinery in sea urchin larvae is mainly based on direct H+ secretion from the stomach lumen and involves a conserved set of ion pumps and transporters. Hemichordate larvae additionally utilized HCO3- transport pathways to generate even more alkaline digestive conditions. Molecular analyses in combination with acidification experiments supported these findings and identified genes coding for ion pumps energizing gastric alkalization. Given that insect larval guts were also reported to be alkaline, our discovery raises the hypothesis that the bilaterian ancestor utilized alkaline digestive system while the vertebrate lineage has evolved a strategy to strongly acidify their stomachs.

Geochemistry of basal dolomitic and Fe-Mn sediments from the Tyrrhenian Sea

In the Tyrrhenian Sea (Western Mediterranean), unusual reddish, soft to lithified, dolomitic sediments up to 45 m thick overlie igneous crust at the base of thick Pliocene-Quaternary deep-sea sediment successions in the Marsili (Site 650) and Vavilov (Site 651) basins. These sediments also overlie the Gortani Ridge, a basaltic Seamount near the base of the Sardinian continental margin (Site 655). At both basinal sites (650, 651), the lowest sediments are dolomitic, with manganese oxide (MnO) segregations. Whole-rock X-ray diffraction indicates abundant dolomite and quartz, with subordinate calcite, illite (authigenic), feldspar and minor kaolinite, chlorite, and anhydrite. Chemical analyses show strong enrichment in magnesium oxide (MgO) and MnO relative to shale or deep-sea clay. Mg and Mn correlate positively and exhibit decreasing concentrations up the succession in the Marsili Basin (Site 650). The following scenario is proposed: peridotites were exposed on the seafloor in the Vavilov Basin (Site 651) and then eroded, depositing talc in local fine-grained dolomitic sediments within the igneous basement. After local magmatism ended, the igneous basement at each site subsided rapidly (about 800 m/m.y.) and was blanketed with calcareous and clay-rich oozes. During early diagenesis (from isotopic evidence; McKenzie et al., this volume) tepid fluids, of modified seawater composition, reacted with and dolomitized the overlying deep-sea sediments. At Site 651 additional Mg may have been extracted from asthenosphere peridotite cored at shallow depths (about 100 m). One can hypothesize that fluids rich in Mg and Mn were flushed from the igneous basement, triggered by extensional faulting and local tilting during subsidence of the basement, and that these fluids then dolomitized the base of the overlying sediment succession. Late tectonic movements in the Vavilov Basin (Site 651) fractured already lithified dolomitic sediments and more reducing (? hydrothermal) fluids locally remobilized Fe and Mn and corroded dolomite crystals.

Basal dolomitic sediments ofthe Tyrrhenian Sea

Basal dolomitic sediments were recovered at three drill sites in the Tyrrhenian Sea during Ocean Drilling Program (ODP) Leg 107 (Sites 650, 651, and 655). These sediments overlie the basaltic basement complex and are enriched in iron, and in some instances, also in manganese. The manganese enrichments, together with a very slight enrichment in trace transition elements, strongly suggest that the basal sediments have an affinity to deep-sea metalliferous deposits of hydrothermal origin. At Sites 651 and 655, the dolostones contain variable amounts of authigenic palygorskite, a Mgrich clay mineral. At Site 651, the basal sediments are 40 m thick and contain nonstoichiometric dolomite, sometimes Ca rich, but primarily Mg rich. The occurrence of Mg-rich dolomite with excess Mg up to 4% is unusual for the deep-sea environment; it may be associated with a hydrothermally driven flux of altered sea water through the directly underlying basement complex, which comprises basalt, dolerite, and serpentinized peridotite. Low-temperature alteration of the basement complex could produce solutions enriched in Mg. Oxygen-isotope equilibrium temperatures indicate that all of the studied dolomites formed under low-temperature conditions (i.e., < 70?C). The carbon-isotope compositions, together with the strong isotopic covariance, suggest that the Mg-rich dolomite precipitated more rapidly than the Carich dolomite. We suggest that the low-temperature, hydrothermal convection of Mg-rich solutions through the basal sediments in this back-arc basin environment (1) overcame kinetic problems related to the formation of massive dolostones, and (2) provided a mass-transport mechanism for dolomitization.