The conserved role of Krox-20 in directing Hox gene expression during vertebrate hindbrain segmentation.

Transient segmentation in the hindbrain is a fundamental morphogenetic phenomenon in the vertebrate embryo, and the restricted expression of subsets of Hox genes in the developing rhombomeric units and their derivatives is linked with regional specification. Here we show that patterning of the vertebrate hindbrain involves the direct upregulation of the chicken and pufferfish group 2 paralogous genes, Hoxb-2 and Hoxa-2, in rhombomeres 3 and 5 (r3 and r5) by the zinc finger gene Krox-20. We identified evolutionarily conserved r3/r5 enhancers that contain high affinity Krox-20. binding sites capable of mediating transactivation by Krox-20. In addition to conservation of binding sites critical for Krox-20 activity in the chicken Hoxa-2 and pufferfish Hoxb-2 genes, the r3/r5 enhancers are also characterized by the presence of a number of identical motifs likely to be involved in cooperative interactions with Krox-20 during the process of hindbrain patterning in vertebrates.

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(Table 3) Foraminiferal 87Sr/86Sr ratios and ages of ODP Hole 162-986D sediments

Site 986 was drilled to 965 meters below seafloor (mbsf) on the western Svalbard margin to record the onset of glaciations and to date and document the glacial evolution in the Svalbard-Barents Sea region during the Pliocene-Pleistocene. In this paper, results of sedimentological analyses are discussed in light of seismic stratigraphy and new age determinations. The latter were difficult to obtain in the glacial deposits, and datums are sparse. Through combined paleomagnetic data, biostratigraphy, and Sr isotopes, however, an overall chronology for the main evolutionary steps is suggested. The cored sequence at Site 986 is younger than 2.6 Ma, and the lower 60 m of the section contains no evidence of a major glacial influence. An initial glaciation is interpreted to have occurred at ~2.3 Ma, resulting in increased sand deposition from debris flows at Site 986 and forming a prominent seismic reflector, R7. However, glaciers probably did not reach the shelf break until ~1.6-1.7 Ma (Reflector R6), after which the depositional environment was dominated by diamictic debris flows. A gradual change in source area from the Barents Sea to Svalbard is recorded primarily by changes in carbonate and smectite content, ~355 mbsf (Reflector R5), at an interpolated age of 1.4-1.5 Ma. During the last ~1 m.y., Site 986 has undergone more distal deposition as the main depocenters have shifted laterally. This has resulted in less frequent debris flows and more turbidites and hemipelagic deposits, with a slight fining upward of the cored sediments.