Hayhurst G P, Lee Y H, Lambert G, Ward J M, Gonzalez F J
Laboratory of Metabolism, Division of Basic Sciences, National Cancer Institute, National Institutes of Health, Bethesda, Maryland 20892, USA.
Mol Cell Biol. 2001 Feb;21(4):1393-403. doi: 10.1128/MCB.21.4.1393-1403.2001.
The numerous functions of the liver are controlled primarily at the transcriptional level by the concerted actions of a limited number of hepatocyte-enriched transcription factors (hepatocyte nuclear factor 1alpha [HNF1alpha], -1beta, -3alpha, -3beta, -3gamma, -4alpha, and -6 and members of the c/ebp family). Of these, only HNF4alpha (nuclear receptor 2A1) and HNF1alpha appear to be correlated with the differentiated phenotype of cultured hepatoma cells. HNF1alpha-null mice are viable, indicating that this factor is not an absolute requirement for the formation of an active hepatic parenchyma. In contrast, HNF4alpha-null mice die during embryogenesis. Moreover, recent in vitro experiments using tetraploid aggregation suggest that HNF4alpha is indispensable for hepatocyte differentiation. However, the function of HNF4alpha in the maintenance of hepatocyte differentiation and function is less well understood. To address the function of HNF4alpha in the mature hepatocyte, a conditional gene knockout was produced using the Cre-loxP system. Mice lacking hepatic HNF4alpha expression accumulated lipid in the liver and exhibited greatly reduced serum cholesterol and triglyceride levels and increased serum bile acid concentrations. The observed phenotypes may be explained by (i) a selective disruption of very-low-density lipoprotein secretion due to decreased expression of genes encoding apolipoprotein B and microsomal triglyceride transfer protein, (ii) an increase in hepatic cholesterol uptake due to increased expression of the major high-density lipoprotein receptor, scavenger receptor BI, and (iii) a decrease in bile acid uptake to the liver due to down-regulation of the major basolateral bile acid transporters sodium taurocholate cotransporter protein and organic anion transporter protein 1. These data indicate that HNF4alpha is central to the maintenance of hepatocyte differentiation and is a major in vivo regulator of genes involved in the control of lipid homeostasis.
肝脏的众多功能主要在转录水平上由有限数量的肝富集转录因子(肝细胞核因子1α [HNF1α]、-1β、-3α、-3β、-3γ、-4α、-6以及c/ebp家族成员)协同作用来控制。其中,只有HNF4α(核受体2A1)和HNF1α似乎与培养的肝癌细胞的分化表型相关。HNF1α基因敲除小鼠能够存活,这表明该因子对于活跃肝实质的形成并非绝对必需。相反,HNF4α基因敲除小鼠在胚胎发育过程中死亡。此外,最近使用四倍体聚集的体外实验表明,HNF4α对于肝细胞分化是不可或缺的。然而,HNF4α在维持肝细胞分化和功能方面的作用尚不太清楚。为了研究HNF4α在成熟肝细胞中的功能,利用Cre-loxP系统构建了条件性基因敲除小鼠。缺乏肝脏HNF4α表达的小鼠肝脏中积累了脂质,血清胆固醇和甘油三酯水平大幅降低,血清胆汁酸浓度升高。观察到的表型可能由以下原因解释:(i)由于编码载脂蛋白B和微粒体甘油三酯转移蛋白的基因表达降低,导致极低密度脂蛋白分泌的选择性破坏;(ii)由于主要高密度脂蛋白受体清道夫受体BI的表达增加,肝脏胆固醇摄取增加;(iii)由于主要基底外侧胆汁酸转运蛋白牛磺胆酸钠共转运蛋白和有机阴离子转运蛋白1的下调,肝脏胆汁酸摄取减少。这些数据表明,HNF4α是维持肝细胞分化的核心,并且是体内参与脂质稳态控制的基因的主要调节因子。