Dunlevy Louisa P E, Burren Katie A, Mills Kevin, Chitty Lyn S, Copp Andrew J, Greene Nicholas D E
Neural Development Unit, Institute of Child Health, University College London, United Kingdom.
Birth Defects Res A Clin Mol Teratol. 2006 Jul;76(7):544-52. doi: 10.1002/bdra.20286.
Closure of the cranial neural tube during embryogenesis is a crucial process in development of the brain. Failure of this event results in the severe neural tube defect (NTD) exencephaly, the developmental forerunner of anencephaly.
The requirement for methylation cycle function in cranial neural tube closure was tested by treatment of cultured mouse embryos with cycloleucine or ethionine, inhibitors of methionine adenosyl transferase. Embryonic phenotypes were investigated by histological analysis, and immunostaining was performed for markers of proliferation and apoptosis. Methylation cycle intermediates s-adenosylmethionine and s-adenosylhomocysteine were also quantitated by tandem mass spectrometry.
Ethionine and cycloleucine treatments significantly reduced the ratio of abundance of s-adenosylmethionine to s-adenosylhomocysteine and are, therefore, predicted to suppress the methylation cycle. Exposure to these inhibitors during the period of cranial neurulation caused a high incidence of exencephaly, in the absence of generalized toxicity, growth retardation, or developmental delay. Reduced neuroepithelial thickness and reduced density of cranial mesenchyme were detected in ethionine-treated but not cycloleucine-treated embryos that developed exencephaly. Reduced mesenchymal density is a potential cause of ethionine-induced exencephaly, although we could not detect a causative alteration in proliferation or apoptosis prior to failure of neural tube closure.
Adequate functioning of the methylation cycle is essential for cranial neural tube closure in the mouse, suggesting that suppression of the methylation cycle could also increase the risk of human NTDs. We hypothesize that inhibition of the methylation cycle causes NTDs due to disruption of crucial reactions involving methylation of DNA, proteins or other biomolecules.
胚胎发育过程中颅神经管的闭合是大脑发育的关键过程。这一过程失败会导致严重的神经管缺陷(NTD)——脑膨出,即无脑畸形的发育前驱症状。
通过用环亮氨酸或乙硫氨酸(蛋氨酸腺苷转移酶抑制剂)处理培养的小鼠胚胎,来测试颅神经管闭合过程中甲基化循环功能的需求。通过组织学分析研究胚胎表型,并对增殖和凋亡标志物进行免疫染色。还通过串联质谱法定量甲基化循环中间体S-腺苷甲硫氨酸和S-腺苷同型半胱氨酸。
乙硫氨酸和环亮氨酸处理显著降低了S-腺苷甲硫氨酸与S-腺苷同型半胱氨酸的丰度比,因此预计会抑制甲基化循环。在颅神经形成期暴露于这些抑制剂会导致脑膨出发病率很高,且不存在全身毒性、生长迟缓或发育延迟。在发生脑膨出的经乙硫氨酸处理而非环亮氨酸处理的胚胎中,检测到神经上皮厚度减小和颅间充质密度降低。间充质密度降低是乙硫氨酸诱导脑膨出的一个潜在原因,尽管在神经管闭合失败之前,我们未检测到增殖或凋亡方面的因果性改变。
甲基化循环的充分功能对小鼠颅神经管闭合至关重要,这表明甲基化循环受抑制也可能增加人类患神经管缺陷的风险。我们推测,甲基化循环受抑制导致神经管缺陷是由于涉及DNA、蛋白质或其他生物分子甲基化的关键反应受到破坏。