Terrell Anne M, Anand Deepti, Smith Sylvie F, Dang Christine A, Waters Stephanie M, Pathania Mallika, Beebe David C, Lachke Salil A
Department of Biological Sciences, University of Delaware, Newark, DE, USA.
Department of Ophthalmology and Visual Sciences, Washington University, St. Louis, MO, USA.
Exp Eye Res. 2015 Feb;131:42-55. doi: 10.1016/j.exer.2014.12.011. Epub 2014 Dec 19.
The discovery of cytosolic RNA granule (RG) component proteins associated with human cataract has initiated investigations on post-transcriptional mechanisms of gene expression control in the lens. Application of established mouse lens epithelial cell lines (LECs) can provide rapid insights on RG function in lens cells, especially because mouse mutants in several RG components are not available. However, although these LECs represent potential reagents for such analyses, they are uncharacterized for lens gene expression or RG formation. Therefore, a detailed molecular and cellular characterization of three permanent mouse LECs 17EM15, 21EM15 and αTN4 is performed in this study. Comparative analysis between microarray gene expression datasets on LEC 21EM15 and iSyTE lens tissue demonstrates that 30% of top 200 iSyTE identified lens-enriched genes are expressed in these cells. Majority of these candidates are independently validated to either have lens expression, function or linkage to cataract. Moreover, analysis of microarray data with genes described in Cat-Map, an online database of cataract associated genes and loci, demonstrates that 131 genes linked to cataract loci are expressed in 21EM15 cells. Furthermore, gene expression in LECs is compared to isolated lens epithelium or fiber cells by qRT-PCR and by comparative analyses with publically available epithelium or fiber-specific microarray and RNA-seq (sequencing) datasets. Expression of select candidate genes was validated by regular and real-time quantitative RT-PCR. Expression of lens epithelium-enriched genes Foxe3, Pax6, Anxa4 and Mcm4 is up-regulated in LEC lines, compared to isolated lens fiber cells. Moreover, similar to isolated lens epithelium, all three LECs exhibit down-regulation of fiber cell-expressed genes Crybb1, Mip and Prox1 when compared to fiber cells. These data indicate that the LEC lines exhibit greater similarity to lens epithelium than to fiber cells. Compared to non-lens cell line NIH3T3, LECs exhibit significantly enriched expression of transcription factors with important function in the lens, namely Pax6, Foxe3 and Prox1. In addition to these genes, all three LECs also express key lens- and cataract-associated genes, namely Dkk3, Epha2, Hsf4, Jag1, Mab21l1, Meis1, Pknox1, Pou2f1, Sfrp1, Sparc, Tdrd7 and Trpm3. Additionally, 21EM15 microarrays indicate expression of Chmp4b, Cryab and Tcfap2a among others important genes. Immunostaining with makers for Processing bodies (P-bodies) and Stress granules (SGs) demonstrates that these classes of RGs are robustly expressed in all three LECs. Moreover, under conditions of stress, 17EM15 and αTN4 exhibit significantly higher numbers of P-bodies and SGs compared to NIH3T3 cells. In sum, these data indicate that mouse LECs 21EM15, 17EM15 and αTN4 express key lens or cataract genes, are similar to lens epithelium than fiber cells, and exhibit high levels of P-bodies and SGs, indicating their suitability for investigating gene expression control and RG function in lens-derived cells.
与人类白内障相关的胞质RNA颗粒(RG)组成蛋白的发现,引发了对晶状体中基因表达控制的转录后机制的研究。应用已建立的小鼠晶状体上皮细胞系(LECs)可以快速洞察RG在晶状体细胞中的功能,特别是因为缺乏几种RG组分的小鼠突变体。然而,尽管这些LECs是进行此类分析的潜在试剂,但它们在晶状体基因表达或RG形成方面尚未得到表征。因此,本研究对三种永久性小鼠LECs 17EM15、21EM15和αTN4进行了详细的分子和细胞表征。对LEC 21EM15和iSyTE晶状体组织的微阵列基因表达数据集进行比较分析表明,iSyTE鉴定的前200个晶状体富集基因中有30%在这些细胞中表达。这些候选基因中的大多数经独立验证,要么在晶状体中有表达、具有功能,要么与白内障有关联。此外,利用白内障相关基因和位点的在线数据库Cat-Map中描述的基因对微阵列数据进行分析表明,与白内障位点相关的131个基因在21EM15细胞中表达。此外,通过qRT-PCR以及与公开可用的上皮或纤维特异性微阵列和RNA测序(测序)数据集进行比较分析,将LECs中的基因表达与分离的晶状体上皮或纤维细胞进行比较。通过常规和实时定量RT-PCR验证了所选候选基因的表达。与分离的晶状体纤维细胞相比,晶状体上皮富集基因Foxe3、Pax6、Anxa4和Mcm4在LEC系中的表达上调。此外,与分离的晶状体上皮相似,与纤维细胞相比,所有三种LECs均表现出纤维细胞表达基因Crybb1、Mip和Prox1的下调。这些数据表明,LEC系与晶状体上皮的相似性高于与纤维细胞的相似性。与非晶状体细胞系NIH3T3相比,LECs中在晶状体中具有重要功能的转录因子Pax6、Foxe3和Prox1的表达显著富集。除了这些基因外,所有三种LECs还表达关键的晶状体和白内障相关基因,即Dkk3、Epha2、Hsf4、Jag1、Mab21l1、Meis1、Pknox1、Pou2f1、Sfrp1、Sparc、Tdrd7和Trpm3。此外,21EM15微阵列表明Chmp4b、Cryab和Tcfap2a等其他重要基因的表达。用加工体(P小体)和应激颗粒(SGs)的标志物进行免疫染色表明,这些类型的RG在所有三种LECs中均有强烈表达。此外,在应激条件下,与NIH3T3细胞相比,17EM15和αTN4表现出显著更多的P小体和SGs。总之,这些数据表明,小鼠LECs 21EM15、17EM15和αTN4表达关键的晶状体或白内障基因,与晶状体上皮的相似性高于与纤维细胞的相似性,并且表现出高水平的P小体和SGs,表明它们适合用于研究晶状体来源细胞中的基因表达控制和RG功能。