PMC:1183529 / 12938-18219 JSONTXT

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and Diaphragm Development in the Fog2 Mutant Mouse\nExperiments were conducted to evaluate the role of Fog2 in pulmonary and diaphragmatic development. The pulmonary phenotype is characterized by diffuse hypoplasia and specific loss of the accessory lobe and a portion of the right middle lobe.\nIt is well established that abnormalities in either diaphragmatic development or fetal breathing can result in a secondary pulmonary hypoplasia, although loss of normal structure has never been documented in this context [17,18]. Fog2 is expressed diffusely in the pulmonary mesenchyme during the period of branching morphogenesis, while later expression is restricted to the smooth muscles of airways and pulmonary vessels (Figure 3). This, and the observation that lungs appeared small on transverse sections evaluated prior to diaphragmatic muscularization or function, suggests that the pulmonary hypoplasia occurs independently of the diaphragmatic defect. To test this hypothesis, lungs were dissected from Fog2 −/− mice and littermate controls before the onset of fetal diaphragmatic motion. Fog2 −/− mice were used for this experiment, as we wanted to avoid potential phenotypic variance from the lil hypomorphic mutation. Lungs dissected at E12 from Fog2 −/− embryos were smaller in size and lacked the development of an accessory lobe. In 11 viable Fog2− /− lung culture explants, there was never development of an accessory lobe, and the weights of mutant lungs cultured for 5 d were significantly lower than those of littermate controls (Figure 4). These data demonstrate that the pulmonary hypoplasia in Fog2 mutant mice is a primary defect.\nFigure 3 Fog2 Is Expressed in the Developing Lung and Diaphragm\nFog2 is expressed in the diffuse pulmonary mesenchyme at E13.5 (A) (arrow shows mesenchyme) and is restricted to the bronchial and vascular smooth muscle (sm) at E16.5 (B). Fog2 is expressed diffusely in the developing diaphragm (Dia) both prior to (E11.5) (C) and after muscularization (E13.5) (D).\nFigure 4 Fog2 Is Necessary for Primary Lung Development\nFog2 null lungs removed prior to diaphragmatic muscularization and grown in vitro show no accessory lobe development. Accessory lobe is labeled with black arrow in control littermate lungs. Branching in the unaffected lobes appeared to be delayed by 6–12 h, but all mutants developed an intricate branching pattern in the unaffected lobes after culture for 5 d.\nBecause the Fog2 phenotype is striking for specific lobar loss, the spatial pattern of Fog2 expression was evaluated in normal embryos during the period of early lobar establishment to determine whether Fog2 expression is specifically different at these lobar buds. Expression was evaluated in mice carrying a lacZ gene incorporated into the Fog2 locus (S. Tevosian, unpublished data). In nine mice examined at E11.5, all lungs showed a specific enhancement of Fog2 expression in the mesenchyme surrounding the accessory bud and the right middle lobe bud, which are the lobes that do not develop normally in Fog2 mutant mice (Figure 5). By E12.5, expression was diffuse in the pulmonary mesenchyme, as was seen previously with in situ hybridization on tissue sections.\nFigure 5 Fog2 Expression in Embryonic Non-Mutant Lungs\nIn embryonic non-mutant lungs, Fog2 is most highly expressed at the tips of the accessory (single arrows) and right middle lobes (double arrows) at E11.25 (A) and E11.75 (B), while expression is diffuse by E12.75 (C). Diaphragms from Fog2 lil mice show an intact membrane with a defect in muscular patterning (see Figure 1B). The membranous portion of the diaphragm is populated by a migratory population of muscle precursor cells, much like the limbs [19,20]. Mice with defects in genes known to be important for the control of this process have intact but amuscular diaphragms [17,21,22]. Hepatocyte growth factor/Scatter factor (HGF) is one potential candidate responsible for the guidance of muscle precursors to the membranous diaphragm. It has been shown that HGF is expressed along this anatomic pathway [23], and mice with absence of the HGF receptor c-Met fail to form muscularized pleuroperitoneal folds (PPFs), and thus have amuscular diaphragms [24,25]. Fog2 is expressed diffusely in the early amuscular diaphragm at E11.5 as well as in the later muscularized diaphragm (see Figure 3C and 3D). Pax3 and MyoD, transcription factors required for appropriate migration and determination of myogenic precursors, were detected in the PPFs of Fog2 lil mice (data not shown). However, in situ expression analysis demonstrated that the expression of HGF in the region where this structure meets the membranous diaphragm was markedly reduced in Fog2 mutant mice (Figure 6). We hypothesize that Fog2 is required (either directly or indirectly) for the induction of HGF in the developing diaphragm, and dysregulation of HGF patterning along the path of muscle precursor cell migration between the PPF and the diaphragm accounts for the abnormal phenotype in these mice.\nFigure 6 HGF Patterning Is Abnormal in Fog2 Null Mice\nIn situ hybridization of HGF in E12.5 wild-type (A) and Fog2 −/− (B) embryos demonstrates decreased expression in the region where the PPF meets the membranous diaphragm. Li, liver; Lu, lung."}

{"project":"2_test","denotations":[{"id":"16103912-10696067-85559739","span":{"begin":521,"end":523},"obj":"10696067"},{"id":"16103912-9355800-85559740","span":{"begin":524,"end":526},"obj":"9355800"},{"id":"16103912-11063939-85559741","span":{"begin":3716,"end":3718},"obj":"11063939"},{"id":"16103912-10066685-85559742","span":{"begin":3719,"end":3721},"obj":"10066685"},{"id":"16103912-10696067-85559743","span":{"begin":3843,"end":3845},"obj":"10696067"},{"id":"16103912-12388344-85559744","span":{"begin":3846,"end":3848},"obj":"12388344"},{"id":"16103912-10226008-85559745","span":{"begin":3849,"end":3851},"obj":"10226008"},{"id":"16103912-10079225-85559746","span":{"begin":4075,"end":4077},"obj":"10079225"},{"id":"16103912-7651534-85559747","span":{"begin":4221,"end":4223},"obj":"7651534"},{"id":"16103912-8681797-85559748","span":{"begin":4224,"end":4226},"obj":"8681797"},{"id":"T66232","span":{"begin":521,"end":523},"obj":"10696067"},{"id":"T74273","span":{"begin":524,"end":526},"obj":"9355800"},{"id":"T90741","span":{"begin":3716,"end":3718},"obj":"11063939"},{"id":"T7030","span":{"begin":3719,"end":3721},"obj":"10066685"},{"id":"T63606","span":{"begin":3843,"end":3845},"obj":"10696067"},{"id":"T80598","span":{"begin":3846,"end":3848},"obj":"12388344"},{"id":"T49937","span":{"begin":3849,"end":3851},"obj":"10226008"},{"id":"T83298","span":{"begin":4075,"end":4077},"obj":"10079225"},{"id":"T51026","span":{"begin":4221,"end":4223},"obj":"7651534"},{"id":"T15263","span":{"begin":4224,"end":4226},"obj":"8681797"}],"text":"Lung and Diaphragm Development in the Fog2 Mutant Mouse\nExperiments were conducted to evaluate the role of Fog2 in pulmonary and diaphragmatic development. The pulmonary phenotype is characterized by diffuse hypoplasia and specific loss of the accessory lobe and a portion of the right middle lobe.\nIt is well established that abnormalities in either diaphragmatic development or fetal breathing can result in a secondary pulmonary hypoplasia, although loss of normal structure has never been documented in this context [17,18]. Fog2 is expressed diffusely in the pulmonary mesenchyme during the period of branching morphogenesis, while later expression is restricted to the smooth muscles of airways and pulmonary vessels (Figure 3). This, and the observation that lungs appeared small on transverse sections evaluated prior to diaphragmatic muscularization or function, suggests that the pulmonary hypoplasia occurs independently of the diaphragmatic defect. To test this hypothesis, lungs were dissected from Fog2 −/− mice and littermate controls before the onset of fetal diaphragmatic motion. Fog2 −/− mice were used for this experiment, as we wanted to avoid potential phenotypic variance from the lil hypomorphic mutation. Lungs dissected at E12 from Fog2 −/− embryos were smaller in size and lacked the development of an accessory lobe. In 11 viable Fog2− /− lung culture explants, there was never development of an accessory lobe, and the weights of mutant lungs cultured for 5 d were significantly lower than those of littermate controls (Figure 4). These data demonstrate that the pulmonary hypoplasia in Fog2 mutant mice is a primary defect.\nFigure 3 Fog2 Is Expressed in the Developing Lung and Diaphragm\nFog2 is expressed in the diffuse pulmonary mesenchyme at E13.5 (A) (arrow shows mesenchyme) and is restricted to the bronchial and vascular smooth muscle (sm) at E16.5 (B). Fog2 is expressed diffusely in the developing diaphragm (Dia) both prior to (E11.5) (C) and after muscularization (E13.5) (D).\nFigure 4 Fog2 Is Necessary for Primary Lung Development\nFog2 null lungs removed prior to diaphragmatic muscularization and grown in vitro show no accessory lobe development. Accessory lobe is labeled with black arrow in control littermate lungs. Branching in the unaffected lobes appeared to be delayed by 6–12 h, but all mutants developed an intricate branching pattern in the unaffected lobes after culture for 5 d.\nBecause the Fog2 phenotype is striking for specific lobar loss, the spatial pattern of Fog2 expression was evaluated in normal embryos during the period of early lobar establishment to determine whether Fog2 expression is specifically different at these lobar buds. Expression was evaluated in mice carrying a lacZ gene incorporated into the Fog2 locus (S. Tevosian, unpublished data). In nine mice examined at E11.5, all lungs showed a specific enhancement of Fog2 expression in the mesenchyme surrounding the accessory bud and the right middle lobe bud, which are the lobes that do not develop normally in Fog2 mutant mice (Figure 5). By E12.5, expression was diffuse in the pulmonary mesenchyme, as was seen previously with in situ hybridization on tissue sections.\nFigure 5 Fog2 Expression in Embryonic Non-Mutant Lungs\nIn embryonic non-mutant lungs, Fog2 is most highly expressed at the tips of the accessory (single arrows) and right middle lobes (double arrows) at E11.25 (A) and E11.75 (B), while expression is diffuse by E12.75 (C). Diaphragms from Fog2 lil mice show an intact membrane with a defect in muscular patterning (see Figure 1B). The membranous portion of the diaphragm is populated by a migratory population of muscle precursor cells, much like the limbs [19,20]. Mice with defects in genes known to be important for the control of this process have intact but amuscular diaphragms [17,21,22]. Hepatocyte growth factor/Scatter factor (HGF) is one potential candidate responsible for the guidance of muscle precursors to the membranous diaphragm. It has been shown that HGF is expressed along this anatomic pathway [23], and mice with absence of the HGF receptor c-Met fail to form muscularized pleuroperitoneal folds (PPFs), and thus have amuscular diaphragms [24,25]. Fog2 is expressed diffusely in the early amuscular diaphragm at E11.5 as well as in the later muscularized diaphragm (see Figure 3C and 3D). Pax3 and MyoD, transcription factors required for appropriate migration and determination of myogenic precursors, were detected in the PPFs of Fog2 lil mice (data not shown). However, in situ expression analysis demonstrated that the expression of HGF in the region where this structure meets the membranous diaphragm was markedly reduced in Fog2 mutant mice (Figure 6). We hypothesize that Fog2 is required (either directly or indirectly) for the induction of HGF in the developing diaphragm, and dysregulation of HGF patterning along the path of muscle precursor cell migration between the PPF and the diaphragm accounts for the abnormal phenotype in these mice.\nFigure 6 HGF Patterning Is Abnormal in Fog2 Null Mice\nIn situ hybridization of HGF in E12.5 wild-type (A) and Fog2 −/− (B) embryos demonstrates decreased expression in the region where the PPF meets the membranous diaphragm. Li, liver; Lu, lung."}

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and Diaphragm Development in the Fog2 Mutant Mouse\nExperiments were conducted to evaluate the role of Fog2 in pulmonary and diaphragmatic development. The pulmonary phenotype is characterized by diffuse hypoplasia and specific loss of the accessory lobe and a portion of the right middle lobe.\nIt is well established that abnormalities in either diaphragmatic development or fetal breathing can result in a secondary pulmonary hypoplasia, although loss of normal structure has never been documented in this context [17,18]. Fog2 is expressed diffusely in the pulmonary mesenchyme during the period of branching morphogenesis, while later expression is restricted to the smooth muscles of airways and pulmonary vessels (Figure 3). This, and the observation that lungs appeared small on transverse sections evaluated prior to diaphragmatic muscularization or function, suggests that the pulmonary hypoplasia occurs independently of the diaphragmatic defect. To test this hypothesis, lungs were dissected from Fog2 −/− mice and littermate controls before the onset of fetal diaphragmatic motion. Fog2 −/− mice were used for this experiment, as we wanted to avoid potential phenotypic variance from the lil hypomorphic mutation. Lungs dissected at E12 from Fog2 −/− embryos were smaller in size and lacked the development of an accessory lobe. In 11 viable Fog2− /− lung culture explants, there was never development of an accessory lobe, and the weights of mutant lungs cultured for 5 d were significantly lower than those of littermate controls (Figure 4). These data demonstrate that the pulmonary hypoplasia in Fog2 mutant mice is a primary defect.\nFigure 3 Fog2 Is Expressed in the Developing Lung and Diaphragm\nFog2 is expressed in the diffuse pulmonary mesenchyme at E13.5 (A) (arrow shows mesenchyme) and is restricted to the bronchial and vascular smooth muscle (sm) at E16.5 (B). Fog2 is expressed diffusely in the developing diaphragm (Dia) both prior to (E11.5) (C) and after muscularization (E13.5) (D).\nFigure 4 Fog2 Is Necessary for Primary Lung Development\nFog2 null lungs removed prior to diaphragmatic muscularization and grown in vitro show no accessory lobe development. Accessory lobe is labeled with black arrow in control littermate lungs. Branching in the unaffected lobes appeared to be delayed by 6–12 h, but all mutants developed an intricate branching pattern in the unaffected lobes after culture for 5 d.\nBecause the Fog2 phenotype is striking for specific lobar loss, the spatial pattern of Fog2 expression was evaluated in normal embryos during the period of early lobar establishment to determine whether Fog2 expression is specifically different at these lobar buds. Expression was evaluated in mice carrying a lacZ gene incorporated into the Fog2 locus (S. Tevosian, unpublished data). In nine mice examined at E11.5, all lungs showed a specific enhancement of Fog2 expression in the mesenchyme surrounding the accessory bud and the right middle lobe bud, which are the lobes that do not develop normally in Fog2 mutant mice (Figure 5). By E12.5, expression was diffuse in the pulmonary mesenchyme, as was seen previously with in situ hybridization on tissue sections.\nFigure 5 Fog2 Expression in Embryonic Non-Mutant Lungs\nIn embryonic non-mutant lungs, Fog2 is most highly expressed at the tips of the accessory (single arrows) and right middle lobes (double arrows) at E11.25 (A) and E11.75 (B), while expression is diffuse by E12.75 (C). Diaphragms from Fog2 lil mice show an intact membrane with a defect in muscular patterning (see Figure 1B). The membranous portion of the diaphragm is populated by a migratory population of muscle precursor cells, much like the limbs [19,20]. Mice with defects in genes known to be important for the control of this process have intact but amuscular diaphragms [17,21,22]. Hepatocyte growth factor/Scatter factor (HGF) is one potential candidate responsible for the guidance of muscle precursors to the membranous diaphragm. It has been shown that HGF is expressed along this anatomic pathway [23], and mice with absence of the HGF receptor c-Met fail to form muscularized pleuroperitoneal folds (PPFs), and thus have amuscular diaphragms [24,25]. Fog2 is expressed diffusely in the early amuscular diaphragm at E11.5 as well as in the later muscularized diaphragm (see Figure 3C and 3D). Pax3 and MyoD, transcription factors required for appropriate migration and determination of myogenic precursors, were detected in the PPFs of Fog2 lil mice (data not shown). However, in situ expression analysis demonstrated that the expression of HGF in the region where this structure meets the membranous diaphragm was markedly reduced in Fog2 mutant mice (Figure 6). We hypothesize that Fog2 is required (either directly or indirectly) for the induction of HGF in the developing diaphragm, and dysregulation of HGF patterning along the path of muscle precursor cell migration between the PPF and the diaphragm accounts for the abnormal phenotype in these mice.\nFigure 6 HGF Patterning Is Abnormal in Fog2 Null Mice\nIn situ hybridization of HGF in E12.5 wild-type (A) and Fog2 −/− (B) embryos demonstrates decreased expression in the region where the PPF meets the membranous diaphragm. Li, liver; Lu, lung."}

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and Diaphragm Development in the Fog2 Mutant Mouse\nExperiments were conducted to evaluate the role of Fog2 in pulmonary and diaphragmatic development. The pulmonary phenotype is characterized by diffuse hypoplasia and specific loss of the accessory lobe and a portion of the right middle lobe.\nIt is well established that abnormalities in either diaphragmatic development or fetal breathing can result in a secondary pulmonary hypoplasia, although loss of normal structure has never been documented in this context [17,18]. Fog2 is expressed diffusely in the pulmonary mesenchyme during the period of branching morphogenesis, while later expression is restricted to the smooth muscles of airways and pulmonary vessels (Figure 3). This, and the observation that lungs appeared small on transverse sections evaluated prior to diaphragmatic muscularization or function, suggests that the pulmonary hypoplasia occurs independently of the diaphragmatic defect. To test this hypothesis, lungs were dissected from Fog2 −/− mice and littermate controls before the onset of fetal diaphragmatic motion. Fog2 −/− mice were used for this experiment, as we wanted to avoid potential phenotypic variance from the lil hypomorphic mutation. Lungs dissected at E12 from Fog2 −/− embryos were smaller in size and lacked the development of an accessory lobe. In 11 viable Fog2− /− lung culture explants, there was never development of an accessory lobe, and the weights of mutant lungs cultured for 5 d were significantly lower than those of littermate controls (Figure 4). These data demonstrate that the pulmonary hypoplasia in Fog2 mutant mice is a primary defect.\nFigure 3 Fog2 Is Expressed in the Developing Lung and Diaphragm\nFog2 is expressed in the diffuse pulmonary mesenchyme at E13.5 (A) (arrow shows mesenchyme) and is restricted to the bronchial and vascular smooth muscle (sm) at E16.5 (B). Fog2 is expressed diffusely in the developing diaphragm (Dia) both prior to (E11.5) (C) and after muscularization (E13.5) (D).\nFigure 4 Fog2 Is Necessary for Primary Lung Development\nFog2 null lungs removed prior to diaphragmatic muscularization and grown in vitro show no accessory lobe development. Accessory lobe is labeled with black arrow in control littermate lungs. Branching in the unaffected lobes appeared to be delayed by 6–12 h, but all mutants developed an intricate branching pattern in the unaffected lobes after culture for 5 d.\nBecause the Fog2 phenotype is striking for specific lobar loss, the spatial pattern of Fog2 expression was evaluated in normal embryos during the period of early lobar establishment to determine whether Fog2 expression is specifically different at these lobar buds. Expression was evaluated in mice carrying a lacZ gene incorporated into the Fog2 locus (S. Tevosian, unpublished data). In nine mice examined at E11.5, all lungs showed a specific enhancement of Fog2 expression in the mesenchyme surrounding the accessory bud and the right middle lobe bud, which are the lobes that do not develop normally in Fog2 mutant mice (Figure 5). By E12.5, expression was diffuse in the pulmonary mesenchyme, as was seen previously with in situ hybridization on tissue sections.\nFigure 5 Fog2 Expression in Embryonic Non-Mutant Lungs\nIn embryonic non-mutant lungs, Fog2 is most highly expressed at the tips of the accessory (single arrows) and right middle lobes (double arrows) at E11.25 (A) and E11.75 (B), while expression is diffuse by E12.75 (C). Diaphragms from Fog2 lil mice show an intact membrane with a defect in muscular patterning (see Figure 1B). The membranous portion of the diaphragm is populated by a migratory population of muscle precursor cells, much like the limbs [19,20]. Mice with defects in genes known to be important for the control of this process have intact but amuscular diaphragms [17,21,22]. Hepatocyte growth factor/Scatter factor (HGF) is one potential candidate responsible for the guidance of muscle precursors to the membranous diaphragm. It has been shown that HGF is expressed along this anatomic pathway [23], and mice with absence of the HGF receptor c-Met fail to form muscularized pleuroperitoneal folds (PPFs), and thus have amuscular diaphragms [24,25]. Fog2 is expressed diffusely in the early amuscular diaphragm at E11.5 as well as in the later muscularized diaphragm (see Figure 3C and 3D). Pax3 and MyoD, transcription factors required for appropriate migration and determination of myogenic precursors, were detected in the PPFs of Fog2 lil mice (data not shown). However, in situ expression analysis demonstrated that the expression of HGF in the region where this structure meets the membranous diaphragm was markedly reduced in Fog2 mutant mice (Figure 6). We hypothesize that Fog2 is required (either directly or indirectly) for the induction of HGF in the developing diaphragm, and dysregulation of HGF patterning along the path of muscle precursor cell migration between the PPF and the diaphragm accounts for the abnormal phenotype in these mice.\nFigure 6 HGF Patterning Is Abnormal in Fog2 Null Mice\nIn situ hybridization of HGF in E12.5 wild-type (A) and Fog2 −/− (B) embryos demonstrates decreased expression in the region where the PPF meets the membranous diaphragm. Li, liver; Lu, lung."}