奶山羊乳腺组织乳脂代谢相关miRNAs的筛选及功能验证
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摘要
山羊乳中总脂肪含量,尤其是其中与人类健康密切相关的中短链脂肪酸及不饱和脂肪酸含量均高于牛乳。山羊乳的乳脂特点应与其乳腺乳脂代谢密切相关。然而,在乳脂代谢基因中,至今尚未发现山羊特异性表达或者具有特异功能的基因。据此推测山羊乳脂代谢的特点可能与其在基因不同水平(如转录后、翻译水平)的调控机制有关。MicroRNAs (miRNAs)是一类在转录后水平调控基因表达的小分子RNA,其功能几乎涉及生命活动的各个方面。已证实miRNAs是调控脂肪和肝脏组织中脂肪代谢的重要因子,然而,现有研究缺乏(?)niRNAs调控山羊乳腺乳脂代谢的研究报道。因此,本研究从miRNAs调控乳腺乳脂代谢的角度出发,通过测序技术获得西农萨能奶山羊乳腺组织miRNAs表达谱,分析其序列特征,检测其在乳腺组织不同阶段的表达水平,筛选与山羊乳脂代谢相关的miRNAs,并通过在山羊乳腺上皮细胞(GMEC)中过表达miRNAs,检测细胞中与乳脂代谢相关的指标,证实miRNAs对山羊乳脂合成和组成具有重要影响,同时发现3对可在乳腺组织中协同表达的miRNAs,发现其协同调控山羊乳腺组织乳脂代谢。本研究从miRNAs调控角度阐明山羊乳脂代谢分子调控机制、揭示山羊乳腺泌乳机理,为后期增加乳脂含量及改善乳组成等研究提供重要的理论和试验依据。论文主要结果如下：
1.利用Solexa测序技术检测山羊泌乳中期乳腺组织中的小RNAs序列,测序结果经数据库过滤后得到高质量小RNAs序列读数22,084,321个,其中长度为22个核苷酸(nt)的小RNA读数最多,约占所有小RNAs读数的33.4%。将测序所得序列与小RNAs数据库中已知序列进行比对,结果发现山羊乳腺组织中含有5种小RNA,分别为tRNAs、 rRNAs、snoRNAs、snRNAs和miRNAs,其中miRNAs的读数最多(10,534,221),约占乳腺组织小RNAs,总读数的47.7%。对miRNAs序列进行鉴定,共获得1,143个miRNAs,其中931个为保守miRNAs,212个为山羊新的候选miRNAs。将山羊931个保守miRNAs与miRBase (17.0)数据库中已注释的miRNAs序列进行比对,结果发现：(1)山羊和牛的保守niRNAs数目最多,达575个,其中,山羊与牛特有的miRNAs数目为116个；(2)335个miRNAs的391个末端碱基发生变化,其中碱基减少的数目大于增加的数目,且miRNAs3'端的碱基变化数目大于5'端；(3)109个碱基发生替换,包括63个转换和46个颠换。
2.利用定量PCR技术检测山羊30个miRNAs在泌乳中期和干奶期乳腺组织中的表达量,获得17个差异表达miRNAs。利用PicTar和TargetScan软件预测miRNAs的靶基因,并通过GO和KEGG软件预测靶基因的功能并进行靶基因富集,结果发现17个差异表达miRNAs的靶基因主要参与乳腺发育、细胞增殖、乳脂合成等代谢过程。从17个差异表达miRNAs及10个功能已知(调控其它动物脂质代谢),miRNAs中,通过催乳素实验获得4个(miR-23a、miR-27a、miR-103和miR-200a)表达量受催乳素调控的miRNAs,推测这4个miRNAs可能与山羊乳腺组织乳脂代谢有关。
3.从山羊基因组DNA上扩增出200bp至500bp长的pri-miR-23a、pri-miR-27a、 pri-miR-103和pri-miR-200a序列(皆包括pre-miRNA及侧翼序列),与牛的序列相比,山羊pre-miR-27a3'端第11个碱基由A变为G,山羊其余pre-miRNAs序列与牛相同。以pri-miRNAs为插入片段,成功构建重组腺病毒载体pAd-miRNAs。将腺病毒载体转染HEK293细胞株,病毒载体包装成功后获得Ad-miR-23a、Ad-miR-27a、Ad-miR-103和Ad-miR-200a4种腺病毒,用这4种病毒分别感染GMEC,72h后检测细胞中miRNAs表达水平,结果发现病毒介导miRNAs的表达量依次为对照(感染Ad的GMEC)的3.48、1.94、2.92和2.48倍。
4.检测感染Ad-miR-10372h时GMEC中的乳脂合成相关指标,结果显示：miR-103过表达可促进细胞中乳脂积累,与对照(感染Ad的细胞)相比,细胞中甘油三酯的含量增加0.33倍,总脂肪酸含量增加0.16倍,不饱和脂肪酸含量增加0.42倍,其中,不饱和脂肪酸c9-C18：1和c9,t11-C18：2含量大幅上调,较对照相比分别上调0.35倍和1.26倍,同时,脂肪酸和甘油三酯合成及脂滴生成相关基因FASN, DGAT1、ADRP、SLA27A6等表达量升高。在感染病毒0、24、48和72h的GMEC中,miR-103过表达可上调PPARγ、 SREBP-lc和LXRa及它们下游基因mRNAs表达水平,同时下调脂解和氧化相关基因HSL、ATGL、CPT1和ACOX1的表达量。此外,miR-103与PANK3在乳腺组织中的表达量之间具有正相关关系,相关系数达0.891,且在GMEC中进行miR-103过表达和抑制试验时,PANK3的表达量随着miR-103表达量变化而改变。
5.检测感染Ad-miR-27a72h时GMEC中的乳脂合成相关指标,结果显示：与对照(感染Ad的细胞)相比,过表达miR-27a能抑制细胞中脂滴形成,下调甘油三酯含量,降低不饱和脂肪酸/饱和脂肪酸的比例,其中,不饱和脂肪酸c9,C18：1和c9,t11-C18：2含量大幅下降,较对照分别下降0.36和0.45倍,而饱和脂肪酸C16：0和C18：0含量升高,较对照分别升高0.19和1.02倍。检测感染病毒0、24、48和72h时细胞中乳脂代谢相关基因的表达量：PPARγ蛋白表达量降低,其下游基因、负责甘油三酯合成的DGAT1的表达量下降,同时,与脂滴形成相关的基因ADRP和TIP47表达量升高,与脂解、氧化相关的基因HSL、ATGL和ACOX1表达量升高。
6.检测感染Ad-miR-23a72h时GMEC中的乳脂合成相关指标,结果显示：与对照相比,过表达miR-23a的细胞中脂滴含量增加1.83倍,甘油三酯含量增加2.21倍,同时,乳脂合成相关基因FASN、DGAT1和TIP47的表达量升高,依次为对照的1.82、1.89、2.93倍,而脂解相关基因ATGL的表达量降低,仅为对照的0.6倍。此外,过表达miR-23a还可增加固醇转运相关基因ABCA1和ABCG2的表达水平。检测感染Ad-miR-200a72h时GMEC中乳脂代谢相关基因的表达量,结果发现miR-200a对乳脂合成各阶段相关基因的表达皆有影响,其可降低FASN和ADRP的表达量,提高SCD、 DGTA1和HSL的表达量,同时还可调控PPARγ和SREBP-1c的表达水平。
7.利用Pearson相关系数分析miR-23a、miR-27a、miR-103和miR-200a在泌乳中期山羊乳腺组织中表达量之间的相关性,结果发现miR-23a与miR-27a、miR-27a与miR-200a、miR-200a与miR-103三对miRNAs之间的表达量具有正相关关系,它们的相关系数依次为0.79、0.57和0.716。此外,在这三对miRNAs中,miR-23a与miR-27a、 miR-27a与miR-200a之间在过表达时均表现为相互促进作用,而miR-103与miR-200a在过表达时表现为相互抑制作用。
Goat milk has higher fat content than cow milk. Goat milk also contains greater amounts of short-chain fatty acids, medium-chain fatty acids, and unsaturated fatty acids which are considered good for human health. These characteristics of goat milk may be attributable to the mechanism of milk fat metabolism. In comparison to cow, no specific gene in goat has been identified. Furthermore, the gene's function identified in goat and cow is the same according to the current reports. It has been hypothesized that the difference in milk fat content and composition between goat and cow are caused by the difference in gene expression and regulation in each levels (i.e., post-transcription, post-translation). MicroRNAs (miRNAs) are small, non-coding RNAs that have emerged as key post-transcriptional regulators of gene expression in nearly all molecular metabolisms. MiRNAs have been reported to be important regulators for lipid and fat metabolism both in adipose and liver tissues. Little is known about the biological roles of miRNAs in goat mammary gland. To investigate the function of miRNAs regulating milk fat metabolism in goat mammary gland, we choose miRNAs as an entry point and performed a large scale miRNAs sequencing to obtain the miRNA profile in lactating mammary gland of Xinong Saanen dairy goats. We characterized goat miRNAs, measured their expression in mammary gand during different phases, and screened miRNAs associated with milk fat synthesis. Then, we over-expressed the miRNAs in goat mammary gland epithelial cells (GMEC) by using recombinant adenovirus, and measured the indices of milk fat synthesis of the GMEC. We also identified the synergistic regulation among miRNAs in mammary gland. This study is helpful with understanding the mechanism of lactation and milk fatty acid metabolism, and will provide a new subject for improving the quality of goat milk and increasing beneficial content of fatty acid in milk. The main results of this study were showed as following:
1. Solexa sequencing technique was used to profile all small RNAs in mammary gland of lactating goats. After filtering low quality data,22,084,321clean reads of small RNAs were obtained. Among these small RNAs, the number of22nucleotides (nt) small RNAs (33.4%of total small RNAs reads) is the most dominant fragments. Based on the results of sequence alignment, five categories small RNA (tRNAs, rRNAs, snoRNAs, snRNAs, and miRNAs) were identified in goat mammary gland, of which miRNAs (10,534,221reads) has the highest percentage composition accounted for47.7%of total small RNAs reads. Goat mammary gland contains1,143miRNAs including931conserved miRNAs and212novel miRNA candidates. The931conserved miRNAs were aligned with current annotated miRNAs in the miRBase (Release17.0). Results of sequence alignment showed:(1) goat and cow (575miRNAs) have the largest number of conserved miRNAs, at meanwhile, goat and cow (116miRNAs) also have the largest number of specific miRNAs which have been identified only in two species;(2) Among conserved miRNAs,335miRNAs containing391end variations were identified. Frequency of base reduction is higher than base addition.3'base variations happen more easily than5';(3)109base substitutions containing63transitions and46transversions were found.
2. PCR was used to identify the expression levels of the read top30miRNAs in mammary gland during mid-lactation and dry period.17differently expressed miRNAs were identified. PicTar and TargetScan softwares were used to predict miRNA targets. GO Term and KEGG softwares were used to enrich the miRNAs targets. The majority of target function of the17miRNAs is associated with processes of development, proliferation and fat synthesis. Among the17miRNAs and10known function (regulating lipid metabolism in other mammals) miRNAs,4miRNAs (miR-23a, miR-27a, miR-103, and miR-200a) were identified by prolactin experiment. The4miRNAs positively response to prolactin and had potential functions of regulating milk fat metabolism in goat mammary gland.
3.200-500bp lengths (including pre-miRNA and flanking sequences) of pri-miR-23a, pri-miR-27a, pri-miR-103and pri-miR-200a were amplified from goat genome, respectively. In comparison to cow, goat pre-miR-27a has one base mutation from A to G on position11of3'. The other goat pre-miRNAs have identical sequences with cow.4recombinant adenovirus plasmids were constructed with their respective pri-miRNA as an inserted sequence. The adenovirus plasmids were transfected to HEK293and packaged in the cells successfully. Adenovirus stably over-expressed miRNAs in GMEC. The miRNA expression mediated by their respective virus in GMEC were3.48-(miR-23a),1.94-(miR-27a),2.92-(miR-103), and2.48-fold (miR-200a) compared with controls (Ad-infected GMECs).
4. Indices of milk fat synthesis were measured in Ad-miR-103-infected cells at72h post infection. In comparison to Ad-infected cells, Ad-miR-103-infected cells accumulated more fat droplets. Triglyceride content increased by0.33-fold, total fatty acid content increased by0.16-fold, and unsaturated fatty acid content increased by0.42-fold. Specifically, c9-C18:1and c9,t11-C18:2contents were increased by1.35-and2.16-fold. The expression of FASN, DGAT1, ADRP and SLA27A6associated with milk fat synthesis increased by1.15-,1.21-,18.02-and14.0-fold, respectively. Furthermore, throughout the observation period (0,24,48and72h post infection), expression of PPARy, SREBP-1c, LXRa and the downstream genes of them were all up-regulated. At meanwhile, expression of HSL, ATGL, CPT1and ACOX1 related to fatty acid lipolysis and oxidation was down-regulated. In addition, a strong correlation was identified between miR-103and its host gene PANK3(correlation coefficient R=0.891) in mammary gland, and miR-103can transcriptionally regulate PANK3expression in GMEC.
5. Indices of milk fat synthesis were measured in Ad-miR-27a-infected cells at72h post infection. In comparison to Ad-infected cells, fat droplet accumulation in Ad-miR-27a-infected cells was suppressed. The triglyceride content and total fatty acid content both decreased. Specifically, c9,C18:1and c9,t11-C18:2contents decreased by0.64-and0.55-fold, whereas, C16:0and C18:0contents increased by1.19and2.02-fold, resulting the decreased ratio of unsaturated/saturated fatty acid content. Furthermore, throughout the observation period (0,24,48and72h post infection), the expression of PPARy protein was suppressed, and the expression of DGAT1, which is related to triglyceride synthesis and also a downstream gene of PPARy, was down-regulated, too. Additionally, HSL, ATGL and ACOX1expression associated with oxidation and lipolysis was increased, as well as ADRP and TIP47involved in fat droplet formation.
6. In comparison to Ad-infected cells, fat droplet and triglyceride contents in Ad-miR-23a-infected cells increased by0.83-and1.21-fold. The expression of FASN, DGAT1and T1P47associated with milk fat synthesis increased by0.82-,0.89-and1.93-fold, whereas, the expression of ATGL related to lipolysis decreased by0.40-fold. Additionally, over-expression of miR-23a up-regulated the expression of ABCA1and ABCG2involved in cholesterol transport. Over-expression of miR-200a in GMEC affected the expression of genes involved in all phases of milk fat synthesis. MiR-200a down-regulated the expression of FASN and ADRP, whereas, up-regulated the expression of SCD, DGAT1and HSL. Furthermore, miR-200a altered the expression of PPARy and SREBP-1c.
7. Pearson correlation coefficient (R) was used to analysize the relationship between miRNAs in mammary gland of lactating goats. A strong correlation was identified between3pairs of miRNAs, miR-103and miR-200a (R=0.716), miR-27a and miR-200a (R=0.57), miR-23a and miR-27a (R=0.79). Furthermore, miR-103and miR-200a can mutually down-regulate expression, and the other two pairs can mutually up-regulated expression.
1.利用Solexa测序技术检测山羊泌乳中期乳腺组织中的小RNAs序列,测序结果经数据库过滤后得到高质量小RNAs序列读数22,084,321个,其中长度为22个核苷酸(nt)的小RNA读数最多,约占所有小RNAs读数的33.4%。将测序所得序列与小RNAs数据库中已知序列进行比对,结果发现山羊乳腺组织中含有5种小RNA,分别为tRNAs、 rRNAs、snoRNAs、snRNAs和miRNAs,其中miRNAs的读数最多(10,534,221),约占乳腺组织小RNAs,总读数的47.7%。对miRNAs序列进行鉴定,共获得1,143个miRNAs,其中931个为保守miRNAs,212个为山羊新的候选miRNAs。将山羊931个保守miRNAs与miRBase (17.0)数据库中已注释的miRNAs序列进行比对,结果发现：(1)山羊和牛的保守niRNAs数目最多,达575个,其中,山羊与牛特有的miRNAs数目为116个；(2)335个miRNAs的391个末端碱基发生变化,其中碱基减少的数目大于增加的数目,且miRNAs3'端的碱基变化数目大于5'端；(3)109个碱基发生替换,包括63个转换和46个颠换。
2.利用定量PCR技术检测山羊30个miRNAs在泌乳中期和干奶期乳腺组织中的表达量,获得17个差异表达miRNAs。利用PicTar和TargetScan软件预测miRNAs的靶基因,并通过GO和KEGG软件预测靶基因的功能并进行靶基因富集,结果发现17个差异表达miRNAs的靶基因主要参与乳腺发育、细胞增殖、乳脂合成等代谢过程。从17个差异表达miRNAs及10个功能已知(调控其它动物脂质代谢),miRNAs中,通过催乳素实验获得4个(miR-23a、miR-27a、miR-103和miR-200a)表达量受催乳素调控的miRNAs,推测这4个miRNAs可能与山羊乳腺组织乳脂代谢有关。
3.从山羊基因组DNA上扩增出200bp至500bp长的pri-miR-23a、pri-miR-27a、 pri-miR-103和pri-miR-200a序列(皆包括pre-miRNA及侧翼序列),与牛的序列相比,山羊pre-miR-27a3'端第11个碱基由A变为G,山羊其余pre-miRNAs序列与牛相同。以pri-miRNAs为插入片段,成功构建重组腺病毒载体pAd-miRNAs。将腺病毒载体转染HEK293细胞株,病毒载体包装成功后获得Ad-miR-23a、Ad-miR-27a、Ad-miR-103和Ad-miR-200a4种腺病毒,用这4种病毒分别感染GMEC,72h后检测细胞中miRNAs表达水平,结果发现病毒介导miRNAs的表达量依次为对照(感染Ad的GMEC)的3.48、1.94、2.92和2.48倍。
4.检测感染Ad-miR-10372h时GMEC中的乳脂合成相关指标,结果显示：miR-103过表达可促进细胞中乳脂积累,与对照(感染Ad的细胞)相比,细胞中甘油三酯的含量增加0.33倍,总脂肪酸含量增加0.16倍,不饱和脂肪酸含量增加0.42倍,其中,不饱和脂肪酸c9-C18：1和c9,t11-C18：2含量大幅上调,较对照相比分别上调0.35倍和1.26倍,同时,脂肪酸和甘油三酯合成及脂滴生成相关基因FASN, DGAT1、ADRP、SLA27A6等表达量升高。在感染病毒0、24、48和72h的GMEC中,miR-103过表达可上调PPARγ、 SREBP-lc和LXRa及它们下游基因mRNAs表达水平,同时下调脂解和氧化相关基因HSL、ATGL、CPT1和ACOX1的表达量。此外,miR-103与PANK3在乳腺组织中的表达量之间具有正相关关系,相关系数达0.891,且在GMEC中进行miR-103过表达和抑制试验时,PANK3的表达量随着miR-103表达量变化而改变。
5.检测感染Ad-miR-27a72h时GMEC中的乳脂合成相关指标,结果显示：与对照(感染Ad的细胞)相比,过表达miR-27a能抑制细胞中脂滴形成,下调甘油三酯含量,降低不饱和脂肪酸/饱和脂肪酸的比例,其中,不饱和脂肪酸c9,C18：1和c9,t11-C18：2含量大幅下降,较对照分别下降0.36和0.45倍,而饱和脂肪酸C16：0和C18：0含量升高,较对照分别升高0.19和1.02倍。检测感染病毒0、24、48和72h时细胞中乳脂代谢相关基因的表达量：PPARγ蛋白表达量降低,其下游基因、负责甘油三酯合成的DGAT1的表达量下降,同时,与脂滴形成相关的基因ADRP和TIP47表达量升高,与脂解、氧化相关的基因HSL、ATGL和ACOX1表达量升高。
6.检测感染Ad-miR-23a72h时GMEC中的乳脂合成相关指标,结果显示：与对照相比,过表达miR-23a的细胞中脂滴含量增加1.83倍,甘油三酯含量增加2.21倍,同时,乳脂合成相关基因FASN、DGAT1和TIP47的表达量升高,依次为对照的1.82、1.89、2.93倍,而脂解相关基因ATGL的表达量降低,仅为对照的0.6倍。此外,过表达miR-23a还可增加固醇转运相关基因ABCA1和ABCG2的表达水平。检测感染Ad-miR-200a72h时GMEC中乳脂代谢相关基因的表达量,结果发现miR-200a对乳脂合成各阶段相关基因的表达皆有影响,其可降低FASN和ADRP的表达量,提高SCD、 DGTA1和HSL的表达量,同时还可调控PPARγ和SREBP-1c的表达水平。
7.利用Pearson相关系数分析miR-23a、miR-27a、miR-103和miR-200a在泌乳中期山羊乳腺组织中表达量之间的相关性,结果发现miR-23a与miR-27a、miR-27a与miR-200a、miR-200a与miR-103三对miRNAs之间的表达量具有正相关关系,它们的相关系数依次为0.79、0.57和0.716。此外,在这三对miRNAs中,miR-23a与miR-27a、 miR-27a与miR-200a之间在过表达时均表现为相互促进作用,而miR-103与miR-200a在过表达时表现为相互抑制作用。
Goat milk has higher fat content than cow milk. Goat milk also contains greater amounts of short-chain fatty acids, medium-chain fatty acids, and unsaturated fatty acids which are considered good for human health. These characteristics of goat milk may be attributable to the mechanism of milk fat metabolism. In comparison to cow, no specific gene in goat has been identified. Furthermore, the gene's function identified in goat and cow is the same according to the current reports. It has been hypothesized that the difference in milk fat content and composition between goat and cow are caused by the difference in gene expression and regulation in each levels (i.e., post-transcription, post-translation). MicroRNAs (miRNAs) are small, non-coding RNAs that have emerged as key post-transcriptional regulators of gene expression in nearly all molecular metabolisms. MiRNAs have been reported to be important regulators for lipid and fat metabolism both in adipose and liver tissues. Little is known about the biological roles of miRNAs in goat mammary gland. To investigate the function of miRNAs regulating milk fat metabolism in goat mammary gland, we choose miRNAs as an entry point and performed a large scale miRNAs sequencing to obtain the miRNA profile in lactating mammary gland of Xinong Saanen dairy goats. We characterized goat miRNAs, measured their expression in mammary gand during different phases, and screened miRNAs associated with milk fat synthesis. Then, we over-expressed the miRNAs in goat mammary gland epithelial cells (GMEC) by using recombinant adenovirus, and measured the indices of milk fat synthesis of the GMEC. We also identified the synergistic regulation among miRNAs in mammary gland. This study is helpful with understanding the mechanism of lactation and milk fatty acid metabolism, and will provide a new subject for improving the quality of goat milk and increasing beneficial content of fatty acid in milk. The main results of this study were showed as following:
1. Solexa sequencing technique was used to profile all small RNAs in mammary gland of lactating goats. After filtering low quality data,22,084,321clean reads of small RNAs were obtained. Among these small RNAs, the number of22nucleotides (nt) small RNAs (33.4%of total small RNAs reads) is the most dominant fragments. Based on the results of sequence alignment, five categories small RNA (tRNAs, rRNAs, snoRNAs, snRNAs, and miRNAs) were identified in goat mammary gland, of which miRNAs (10,534,221reads) has the highest percentage composition accounted for47.7%of total small RNAs reads. Goat mammary gland contains1,143miRNAs including931conserved miRNAs and212novel miRNA candidates. The931conserved miRNAs were aligned with current annotated miRNAs in the miRBase (Release17.0). Results of sequence alignment showed:(1) goat and cow (575miRNAs) have the largest number of conserved miRNAs, at meanwhile, goat and cow (116miRNAs) also have the largest number of specific miRNAs which have been identified only in two species;(2) Among conserved miRNAs,335miRNAs containing391end variations were identified. Frequency of base reduction is higher than base addition.3'base variations happen more easily than5';(3)109base substitutions containing63transitions and46transversions were found.
2. PCR was used to identify the expression levels of the read top30miRNAs in mammary gland during mid-lactation and dry period.17differently expressed miRNAs were identified. PicTar and TargetScan softwares were used to predict miRNA targets. GO Term and KEGG softwares were used to enrich the miRNAs targets. The majority of target function of the17miRNAs is associated with processes of development, proliferation and fat synthesis. Among the17miRNAs and10known function (regulating lipid metabolism in other mammals) miRNAs,4miRNAs (miR-23a, miR-27a, miR-103, and miR-200a) were identified by prolactin experiment. The4miRNAs positively response to prolactin and had potential functions of regulating milk fat metabolism in goat mammary gland.
3.200-500bp lengths (including pre-miRNA and flanking sequences) of pri-miR-23a, pri-miR-27a, pri-miR-103and pri-miR-200a were amplified from goat genome, respectively. In comparison to cow, goat pre-miR-27a has one base mutation from A to G on position11of3'. The other goat pre-miRNAs have identical sequences with cow.4recombinant adenovirus plasmids were constructed with their respective pri-miRNA as an inserted sequence. The adenovirus plasmids were transfected to HEK293and packaged in the cells successfully. Adenovirus stably over-expressed miRNAs in GMEC. The miRNA expression mediated by their respective virus in GMEC were3.48-(miR-23a),1.94-(miR-27a),2.92-(miR-103), and2.48-fold (miR-200a) compared with controls (Ad-infected GMECs).
4. Indices of milk fat synthesis were measured in Ad-miR-103-infected cells at72h post infection. In comparison to Ad-infected cells, Ad-miR-103-infected cells accumulated more fat droplets. Triglyceride content increased by0.33-fold, total fatty acid content increased by0.16-fold, and unsaturated fatty acid content increased by0.42-fold. Specifically, c9-C18:1and c9,t11-C18:2contents were increased by1.35-and2.16-fold. The expression of FASN, DGAT1, ADRP and SLA27A6associated with milk fat synthesis increased by1.15-,1.21-,18.02-and14.0-fold, respectively. Furthermore, throughout the observation period (0,24,48and72h post infection), expression of PPARy, SREBP-1c, LXRa and the downstream genes of them were all up-regulated. At meanwhile, expression of HSL, ATGL, CPT1and ACOX1 related to fatty acid lipolysis and oxidation was down-regulated. In addition, a strong correlation was identified between miR-103and its host gene PANK3(correlation coefficient R=0.891) in mammary gland, and miR-103can transcriptionally regulate PANK3expression in GMEC.
5. Indices of milk fat synthesis were measured in Ad-miR-27a-infected cells at72h post infection. In comparison to Ad-infected cells, fat droplet accumulation in Ad-miR-27a-infected cells was suppressed. The triglyceride content and total fatty acid content both decreased. Specifically, c9,C18:1and c9,t11-C18:2contents decreased by0.64-and0.55-fold, whereas, C16:0and C18:0contents increased by1.19and2.02-fold, resulting the decreased ratio of unsaturated/saturated fatty acid content. Furthermore, throughout the observation period (0,24,48and72h post infection), the expression of PPARy protein was suppressed, and the expression of DGAT1, which is related to triglyceride synthesis and also a downstream gene of PPARy, was down-regulated, too. Additionally, HSL, ATGL and ACOX1expression associated with oxidation and lipolysis was increased, as well as ADRP and TIP47involved in fat droplet formation.
6. In comparison to Ad-infected cells, fat droplet and triglyceride contents in Ad-miR-23a-infected cells increased by0.83-and1.21-fold. The expression of FASN, DGAT1and T1P47associated with milk fat synthesis increased by0.82-,0.89-and1.93-fold, whereas, the expression of ATGL related to lipolysis decreased by0.40-fold. Additionally, over-expression of miR-23a up-regulated the expression of ABCA1and ABCG2involved in cholesterol transport. Over-expression of miR-200a in GMEC affected the expression of genes involved in all phases of milk fat synthesis. MiR-200a down-regulated the expression of FASN and ADRP, whereas, up-regulated the expression of SCD, DGAT1and HSL. Furthermore, miR-200a altered the expression of PPARy and SREBP-1c.
7. Pearson correlation coefficient (R) was used to analysize the relationship between miRNAs in mammary gland of lactating goats. A strong correlation was identified between3pairs of miRNAs, miR-103and miR-200a (R=0.716), miR-27a and miR-200a (R=0.57), miR-23a and miR-27a (R=0.79). Furthermore, miR-103and miR-200a can mutually down-regulate expression, and the other two pairs can mutually up-regulated expression.
引文
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