PMC:1440874 / 17739-26875 JSONTXT

{"target":"http://pubannotation.org/docs/sourcedb/PMC/sourceid/1440874","sourcedb":"PMC","sourceid":"1440874","source_url":"http://www.ncbi.nlm.nih.gov/pmc/1440874","text":"Early Embryonic Lethality in Atrxnull Male Mice\nTo investigate the role of the Atrx protein during mouse development, we initially established lines of mice bearing the Atrx flox allele. Two independently targeted Atrx flox ES cell clones with normal male karyotype were injected into C57BL/6 blastocysts to produce chimaeric mice, which were then used to obtain germline transmission. Intercrosses between males hemizygous (Atrx flox/Y) and females heterozygous (Atrx WT/flox) for the floxed allele were also carried out to generate homozygous females (Atrx flox/flox). Males hemizygous and females heterozygous or homozygous for the Atrx flox allele were viable, appeared healthy, and bred normally, suggesting that, as expected, the Atrx flox allele was functionally normal. To generate Atrxnull mice by Cre-mediated recombination, the Atrx flox mice were crossed with mice harboring a transgene in which the Cre recombinase is expressed under the control of the regulatory elements of the mouse GATA-1 gene (GATA1-cre) [14]. Widespread expression of the GATA1-cre transgene has been demonstrated during early embryogenesis [14]. We more accurately defined the onset of GATA1-cre expression using a ROSA26 reporter strain, in which a β-galactosidase/neor fusion reporter gene is expressed only after Cre-mediated excision of loxP-flanked transcription and translation termination signals [14]. We found that the GATA1-cre transgene was already active at the 16-cell morula stage of development (0.5 days postcoitus [dpc]) (Figure 4A).\nFigure 4 Timing of Onset of GATA1-Cre Expression and PCR Genotyping of Atrx Alleles\n(A) GATA1-cre +/+ transgenic males were crossed to females of the ROSA26 reporter strain (ROSA26 +/−), and embryos were recovered at 0.5 dpc (~16-cell morula stage) and stained with X-gal. Cre-mediated activation of the ROSA26 β-galactosidase reporter allele was detected in all cells in embryos in which both alleles are coinherited.\n(B) Top gel: PCR genotyping of Atrx alleles in embryos using primers PPS1.15 (exon 17) and Mxnp30 (exon 20) as described in Protocol S1. The sizes of PCR products from the different alleles are indicated. Both the Atrx Δ18 (resulting from recombination event B in Figure 2A) and the Atrx Δ18Δneo allele (resulting from recombination event C in Figure 2A) are null for full-length Atrx protein. The bottom gel shows products from a PCR reaction (primers DG52/DG53) used to sex embryos as described in Protocol S1. A 450-bp PCR product is amplified from a mouse Y chromosome-specific satellite repeat. To generate Atrxnull mice, heterozygous floxed females (Atrx WT/flox) were mated with homozygous GATA1-cre transgenic males (Atrx WT/Y;GATA1-cre +/+ ). No Atrxnull males (Atrx null/Y;GATA1-cre +/− ) were recovered at birth, indicating that the absence of Atrx results in embryonic lethality. This finding was unexpected, since human ATR-X patients clearly survive to adulthood (see Discussion). Embryos were dissected at 7.5, 8.5, and 9.5 dpc and genotyped by PCR analysis of DNA extracted from yolk sac or total embryo (Figure 4B and Protocol S1). Atrxnull males were present at expected mendelian ratios (~25%) at both 7.5 dpc and 8.5 dpc (Table 1). However, by 9.5 dpc, depletion was observed both in the number of Atrxnull males (7%) and in the total number of males recovered (31%). No Atrxnull males were recovered after 9.5 dpc. Thus the absence of Atrx gives rise to embryonic lethality in mice before 9.5 dpc.\nTable 1 Distribution of Atrx Genotypes in Timed Matings To investigate the morphology of Atrxnull embryos prior to death, embryos from the above crosses were initially dissected in their deciduas at 7.5 dpc, and paraffin sections were stained with haematoxylin (Figure 5A) or with an anti-ATRX antibody (Figure 5B–5E). Immunohistochemical staining revealed that Atrx was widely expressed in wild-type 7.5 dpc embryos (Figure 5B). Expression was highest in the embryonic region (Figure 5C) and the chorion (Figure 5D). Detectable but lower levels of expression were observed in the ectoplacental cone (Figure 5D) and surrounding decidual tissue. We also observed very high levels of Atrx expression in trophoblast giant cells (TGCs) surrounding the Reichert's membrane (Figure 5E). Within the large nuclei of these TGCs, the typical nuclear association of Atrx with blocks of pericentromeric heterochromatin [15] was clearly observable. Only background staining was seen in the corresponding Atrxnull embryonic tissues (Figure 5B–5D), while expression in the surrounding decidual tissue (of maternal origin) was normal and served as an antibody staining control (unpublished data). Morphologically, 7.5 dpc Atrxnull embryos were dramatically reduced in size and appeared developmentally retarded relative to stage-matched wild-type embryos (Figure 5A and 5B). However, despite their reduced size, the general morphology and organisation of embryonic structures in Atrxnull conceptuses appeared grossly normal. The amnion and chorion were clearly present and the amniotic, exocoelomic, and ectoplacental cavities were distinguishable, as were all three embryonic germ layers (Figure 5A–5C). At 8.5 dpc, embryos were dissected free of deciduas, and observed in whole mount. Individual conceptuses were genotyped by PCR using DNA isolated from yolk sac as described in Protocol S1. Consistent with observations at 7.5 dpc, the general morphology of the embryo proper of Atrxnull conceptuses also appeared grossly normal at 8.5 dpc. The head fold had clearly formed, and expression of the early mesoderm marker brachyury (T) [16] was detected in the primitive streak and emerging notochord by whole-mount in situ hybridisation (WMISH) (Figure 5F), indicating that Atrxnull embryos had gastrulated.\nFigure 5 Morphology of Atrxnull Embryos at 7.5 dpc and 8.5 dpc\nParaffin sections of wild-type or Atrxnull 7.5 dpc embryos (dissected in their deciduas) were stained with haematoxylin (A) or with an anti-ATRX antibody (H-300, Figure 1) (B–E). Photomicrographs C–E show higher magnification images (200×) of the stained sections shown in (B) (40×). Scale bars represent 200 μm (40× magnification) or 40 μm (200× magnification). a, amnion; ac, amniotic cavity; c, chorion; e, epiblast; ec, ectoplacental cavity; ecc, exocoelomic cavity; ep, ectoplacental cone; ne, neural ectoderm; rm, Reichert's membrane; tgc, trophoblast giant cell.\n(F) Detection of brachyury (T) expression in Atrxnull 8.5 dpc embryo (head fold stage) by WMISH. The genotype was determined by PCR (as shown in Protocol S1) using DNA extracted from yolk sac. hf, head fold; n, emerging notochord; ps, primitive streak. To investigate whether the reduced size of the Atrxnull embryos was due to an increase in apoptosis, we analysed sections of paraffin-embedded 7.5 dpc embryos by TdT-mediated dUTP nick end labeling (TUNEL) assay (Figure 6A). Very few apoptotic cells were detected in wild-type 7.5 dpc embryos. In Atrxnull embryos, a slight increase in the apoptotic population was evident. However, consistent with our observation of a grossly normal apoptotic index in Atrxnull ES cells (Figure S2), the apoptotic response observed in Atrxnull embryos was also not uniform, but was restricted to a low number of scattered TUNEL-positive cells. Since this small apoptotic response is unlikely to account fully for the dramatic size deficit observed in Atrxnull embryos, a possible proliferation defect was also investigated by immunohistochemical staining of 7.5 dpc embryo sections for the mitosis marker phosphorylated (Ser10) histone H3 [13]. Relative to the very high mitotic index observed in wild-type embryos, the proportion of mitotic cells observed in Atrxnull embryos at 7.5 dpc was dramatically reduced (Figure 6B). Taken together, these results suggest that the size deficit observed in Atrxnull embryos prior to lethality largely reflects a proliferative defect, with a minor but indirect contribution from increased apoptosis. Although a growth defect was also observed in Atrxnull ES cells (Figure 3A), in contrast to the Atrxnull embryos, the mitotic index of the ES cell population (as measured with the same antibody) was not depleted (Figure S1B). These observations suggest that the mitotic defect observed in embryos is unlikely to be a direct, cell-autonomous effect of the absence of Atrx, and is more likely to be a secondary effect resulting from the failure to develop a normal trophoblast (see below).\nFigure 6 Analysis of Apoptosis and Mitosis in Atrxnull Embryos\n(A) Paraffin sections of wild-type or Atrxnull 7.5 dpc embryos (dissected in their deciduas) were analysed by TUNEL assay and apoptotic cells labelled with fluorescein-dUTP. Sections were counterstained with DAPI.\n(B) Paraffin sections of wild-type or Atrxnull 7.5 dpc embryos were stained with an antibody against the mitosis marker phosphorylated (Ser10) histone H3. Sections were counterstained with haematoxylin. For both (A) and (B), the presence or absence of Atrx in each embryo was determined by staining adjacent sections with the anti-ATRX antibody (H-300) as in Figure 5 (unpublished 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