Neural Induction in Xenopus: Requirement for Ectodermal and Endomesodermal Signals via Chordin, Noggin, β-Catenin, and Cerberus

The origin of the signals that induce the differentiation of the central nervous system (CNS) is a long-standing question in vertebrate embryology. Here we show that Xenopus neural induction starts earlier than previously thought, at the blastula stage, and requires the combined activity of two distinct signaling centers. One is the well-known Nieuwkoop center, located in dorsal-vegetal cells, which expresses Nodal-related endomesodermal inducers. The other is a blastula Chordin- and Noggin-expressing (BCNE) center located in dorsal animal cells that contains both prospective neuroectoderm and Spemann organizer precursor cells. Both centers are downstream of the early β-Catenin signal. Molecular analyses demonstrated that the BCNE center was distinct from the Nieuwkoop center, and that the Nieuwkoop center expressed the secreted protein Cerberus (Cer). We found that explanted blastula dorsal animal cap cells that have not yet contacted a mesodermal substratum can, when cultured in saline solution, express definitive neural markers and differentiate histologically into CNS tissue. Transplantation experiments showed that the BCNE region was required for brain formation, even though it lacked CNS-inducing activity when transplanted ventrally. Cell-lineage studies demonstrated that BCNE cells give rise to a large part of the brain and retina and, in more posterior regions of the embryo, to floor plate and notochord. Loss-of-function experiments with antisense morpholino oligos (MO) showed that the CNS that forms in mesoderm-less Xenopus embryos (generated by injection with Cerberus-Short [CerS] mRNA) required Chordin (Chd), Noggin (Nog), and their upstream regulator β-Catenin. When mesoderm involution was prevented in dorsal marginal-zone explants, the anterior neural tissue formed in ectoderm was derived from BCNE cells and had a complete requirement for Chd. By injecting Chd morpholino oligos (Chd-MO) into prospective neuroectoderm and Cerberus morpholino oligos (Cer-MO) into prospective endomesoderm at the 8-cell stage, we showed that both layers cooperate in CNS formation. The results suggest a model for neural induction in Xenopus in which an early blastula β-Catenin signal predisposes the prospective neuroectoderm to neural induction by endomesodermal signals emanating from Spemann's organizer

Optical Spectroscopy of the IRAS 1-Jy Sample of Ultraluminous Infrared Galaxies

This paper discusses the optical spectroscopic properties of the IRAS 1-Jy sample of ultraluminous infrared galaxies (ULIGs). One hundred and eight of the 118 1-Jy ULIGs have been observed at dlambda = 8.3 AA resolution over the wavelength range ~4500 A -- 8900 A. These data are combined with large, previously published sets of optical spectroscopic data of lower luminosity infrared galaxies to look for systematic trends with infrared luminosity over the luminosity range L_ir ~ 10^{10.5}-10^{13} L_sun. As found in previous studies, the fraction of Seyfert galaxies among luminous infrared galaxies increases abruptly above L_ir ~ 10^{12.3} L_sun --- about 50% of the galaxies with L_ir > 10^{12.3} L_sun present Seyfert characteristics. Many of the optical and infrared spectroscopic properties of the Seyfert galaxies are consistent with the presence of a genuine active galactic nucleus (AGN). About 30% of these galaxies are Seyfert 1s with broad-line regions similar to those of optical quasars. The percentage of Seyfert 1 ULIGs increases with infrared luminosity, contrary to the predictions of the standard unification model for Seyfert galaxies. Comparisons of the broad-line luminosities of optical and obscured Seyfert 1 ULIGs with those of optically selected quasars of comparable bolometric luminosity suggest that the dominant energy source in most of these ULIGs is the same as in optical quasars, namely mass accretion onto a supermassive black hole, rather than a starburst. These results are consistent with recently published ISO, ASCA, and VLBI data. (abridged)Comment: Text and 23 figures (45 pages), Tables 1 - 6 (16 pages