胎牛血清
王秀英 (mary at labome dot com)
美国新泽西州普林斯顿合原研究有限责任公司 (Synatom Research)
译者
王秀英 (mary at labome dot com)
美国新泽西州普林斯顿合原研究有限责任公司 (Synatom Research)
DOI
http://dx.doi.org/10.13070/mm.cn.2.117
日期
更新 : 2016-09-03; 原始版 : 2012-03-22
引用
实验材料和方法 2012;2:117
摘要

深入讨论胎牛血清及其在真核细胞培养的应用。

英文摘要

An in-depth discussion of fetal bovine serum and its applications in eukaryotic cell culture.

什么是胎牛血清?

血清是血液凝固后的液体部分,不含血细胞、纤维蛋白及凝集因子等,含有多种对细胞生长必需的营养因子及大分子物质。血清白蛋白是胎牛血清的主要成分。此外,重要的是,胎牛血清中含有大量对细胞生长必需的生长因子 [1, 2] 。 胎牛血清也含有一些小分子物质,如氨基酸、糖类、脂类及激素类物质。

胎牛血清应用在哺乳动物细胞培养中;因此,胎牛血清应用很广泛。比如,胎牛血清能够用在科研、工业生产、人兽疫苗生产的质量控制及生物技术药物的质量控制领域。

相对于新生动物血清或成体动物血清来说,胚胎期动物血清有什么优点?

与其它类型血清相比,胚胎期动物血清含有更多的生长因子,及较少的gamma球蛋白含量。

胎牛血清 图 1
图 1. 美洲鲎(Horseshoe crabs)。
血清与血浆的区别

血清是由血液凝固后的上清液制备的;而血浆中含有抗凝剂,所以血浆中含有血液凝固相关的蛋白。

胎牛血清在包装销售前,需要经过哪些检测?

国际血清行业协会(www.serumindustry.org)是血清供销商的商业组织。 协会规定血清需要利用特定的方法进行检测,并且检测结果必须显示在化验证明书中。

检测项目
细菌和真菌 -- 无菌检测
支原体
细胞病变诱导剂 -- 病毒检测
血细胞吸附剂 -- 病毒检测
牛病毒性腹泻 -- 病毒检测
pH检测
渗透压检测
总蛋白量,由 双缩脲法检测
内毒素检测
血红素检测
电泳模式检测
其它特性检测,如干细胞培养
表一:最低血清试验的要求。
内毒素检测

内毒素,又叫脂多糖(LPS)或脂寡糖(LOS),存在于革兰氏阴性菌的细胞外膜。临床上,内毒素作为一些革兰氏阴性菌的诊断提示,如脑膜炎球菌。动物体内,内毒素能够导致发热或者诱导炎症反应;细胞培养中,内毒素能够诱导细胞反应异常。

通常,采用鲎阿米巴样细胞裂解实验(LAL检测)检测内毒素水平。LAL是从美洲鲎(见图一)的血液中制备的,对内毒素非常灵敏,在微量内毒素存在情况下就凝结。阿米巴样细胞相当于哺乳动物的白细胞。

特异性血清及其加工方式
热钝化

胎牛血清的常规处理方式是热钝化,即放在56°水浴30分钟;其目的是使血清中的补体成分及一些未知的细胞生长抑制分子失活。

与胎牛血清的其它加工方式一样,热钝化处理过程必须小心谨慎,因为血清中的生长因子容易在较高温度或者较长时间加热条件下失活,甚至产生沉淀。

Life Technologies公司的Invitrogen及GIBCO品牌的热钝化胎牛血清用在细胞培养中,来研究UCP-2及UCP-3 [3] ,核因子kappa-B [4] ,蚕杆状病毒表达系统中表达FMDV衣壳蛋白 [5] ,亚临床的移植物抗宿主病 [6] ,IFN-gamma的作用 [7] ,及自噬 [8] 。Thermo Fisher HyClone公司的热钝化胎牛血清用来研究能引起严重急性呼吸综合征的冠状病毒的核衣壳蛋白 [9] ,NO [10] ,激活素C [11] ,及IE1 [12] 。Sigma公司的热钝化胎牛血清用来研究化疗 [13] ,而Equitech-Bio公司的热钝化胎牛血清用来研究谷胱甘肽巯基转移酶Pi [14] 。

活性炭处理

活性炭能够结合脂溶性分子,因此能够用来除去激素类物质,例如,胎牛血清中的雌二醇、孕酮、皮质醇、睾酮及T3和T4分子。血清中这些脂类激素能够干扰免疫检测及胰岛素检测方法。

Thermo Scientific的活性炭吸附的胎牛血清(csFCS)被用来识别TGF-beta家族成员 [15] 。

透析处理

透析处理能够除去胎牛血清中所有分子量小于10,000Da的分子。例如,激素、细胞因子、葡萄糖、氨基酸还有许多其它分子,都能够被除去。透析也能除去胎牛血清中的抗生素及其它内源分子。

Life Technologies的透析处理的胎牛血清被用来研究驱动蛋白5B [16] 及丙酮酸激酶M2失活 [17] ,Thermo Fisher HyClone的透析处理胎牛血清用来研究KRAS信号通路的突变 [18] ,而Sigma公司的透析处理胎牛血清用来研究PAH代谢情况 [19] 。

gamma照射

Gamma照射能够对胎牛血清进行灭菌处理。胎牛血清常规的灭菌方法是多次透过0.1um的滤膜以消除微生物污染,而gamma照射能够钝化牛群中常见的病毒。但是,一些类型的病毒能够抵抗gamma照射,比如细小病毒。

低IgG含量

低IgG的胎牛血清适用于抗体生产及相关领域。

胎牛血清 图 2
图 2. Gibco Life Technologies许可。
干细胞培养

与一般细胞培养相比,干细胞培养对生长因子有更严格的要求。由于胎牛血清中的一些生长因子促进干细胞分化,所以,需要检测各批胎牛血清在保持不同类型干细胞的全能性方面的性能。

来源地

胎牛血清的主要来源地就是那些大量养牛的国家,包括美国、澳大利亚、新西兰、加拿大及南美和中美洲地区,由政府机构管理血清制品的生产及收集过程。

胎牛血清分为美国农业部(USDA)级和欧盟级。其中,美国农业部级胎牛血清没有疯牛病(BSE)及口蹄疫(FMD)污染,能够出口到任何国家,而欧盟级胎牛血清只在大部分欧洲国家及亚洲国家销售。

如何保存胎牛血清?

胎牛血清一般是在-5°到-20°之间冻存,2°到8°之间融化。一个比较好的办法是,将胎牛血清分装到50ml管中,这样做就避免了反复冻融对血清造成的危害。血清融化后常出现沉淀,可能是血清蛋白变性造成的。简短离心即可除去沉淀,并且,这些沉淀一般不会影响血清质量的。

胎牛血清的选择

许多供应商提供不同等级、不同来源及不同处理的胎牛血清。而胎牛血清是一种复杂的混合物,不同供应商的不同批次产品肯定会有一些不同。因此,有必要建立一套选择、评估胎牛血清的机制。

一般来说,可以通过参考相关研究文献选择适合的胎牛血清。

Labome对212篇科研文献做了关于胎牛血清使用情况的调查。

胎牛血清 图 3
图 3. Thermo Fisher Scientific许可。
文献中的胎牛血清

这些文献是Labome的10,000篇文献中随机抽取的一小部分,选自开放访问的杂志,出版时间介于2006和2014年之间。

胎牛血清的主要供应商有Life Technologies公司,Thermo Scientific HyClone公司及Sigma-Aldrich公司(表二)。

供应商数目
Life Technologies87
Thermo Scientific HyClone43
Sigma-Aldrich28
Gemini Bio-Products10
Atlanta Biologicals6
PAA Laboratories6
PAN Biotech3
Biological Industries2
Omega Scientific2
others23
表二: 调查中的供应商及使用其胎牛血清的文献数目。
Life Technologies公司

Life Technologies公司是一家全球性的生物技术公司,总部设在加利福尼亚州Carlsbad市。该公司是由Invitrogen公司和Applied Biosystems公司于2008年合并而来。GIBCO是该公司的一个品牌,主要提供细胞培养产品,比如细胞培养基、胎牛血清及其它相关试剂等。截止2012年3月,该公司共有大概34种类型的胎牛血清产品。

Labome的调查结果表明,有42%(193篇中的82篇)的文献涉及到GIBCO的胎牛血清。表三列出了这些文献的研究领域分布情况。

研究领域数目
生理学32
免疫学18
肿瘤学18
病理学7
神经科学6
病毒学1
表三: Life Technologies的GIBCO品牌血清在各研究领域的文献引用情况。
生理学领域

生理学是最热门的研究领域,且大部分文献聚焦在对生理过程中基因功能的研究。例如,该公司的胎牛血清产品已经被用来研究人小胶质细胞和脑组织中白蛋白的表达情况 [20] ,层粘连蛋白连接过程中alpha-肌营养不良蛋白聚糖的O-甘露糖磷酸化的功能 [21] ,APE1蛋白残基D70在对DNA的3'重末端剪切中的新作用 [22] ,果蝇中CENH3在着丝粒形成中的作用 [23] ,IGF1异构体及功能 [24] ,NO信号通路中二醇的作用 [25] ,PKC在用异氟烷预诱导产生保护功能的过程中的作用 [26] ,RARalpha上游因子对多向性遗传网络的调控作用 [27] ,在TNF诱导的破骨细胞的前体细胞(OCPs)转移过程中SDF-1/CXCR4轴的作用 [28] ,C端固定的G蛋白alpha亚基对其再循环速率的影响 [29] ,SRp20和CUG-BP1在调控胰岛素受体外显子11的选择性剪切中的功能 [30] ,生长因子对ADAMTS-4表达的影响 [31] ,在ES细胞保持一致性过程中PRC2的作用 [32] ,转录因子Ets-1在调控NPR-A表达过程中的作用 [33] ,在mTOR信号通路中H+-ATPase的作用 [34] ,LC细胞中ET-1对胶原合成及分泌的影响 [35] ,及淋巴细胞中线粒体K+通道Kv1.3在Bax诱导凋亡过程中的作用 [36] 。

此外,胎牛血清产品也被用来研究一些药物的效果,比如说,局部免疫抑制剂 [37] 以及组蛋白去乙酰化酶(HDAC)抑制剂LAQ824 [38] 。一些研究者利用胎牛血清产品研究凋亡进程。例如,Atapattu et al. [39] 和Lam et al. [40] 用胎牛血清培养细胞研究凋亡过程中的具体细节。还有一些科学家研究分子机制 [41-45] 以及某些分子在相应信号通路中的作用 [46-48] ,例如,钙粘素依赖的粘着连接(AJ)与Wnt信号通路之间的相互关系 [47] 、CHIP在聚集小体信号通路中的作用 [48] 。

免疫学领域

免疫学领域也是一个热门领域。胎牛血清产品曾被用来研究IL-23在起始炎症反应中的作用 [49] ,IL-23及IL-12在Dectin-1和TLR信号通路共激活中的作用 [50] ,炎症小体识别在适应性免疫对流感病毒识别中的作用 [51] ,特异的CD40抗体诱导的免疫激活 [52] ,IL-17的免疫缺陷导致的慢性粘膜皮肤念珠菌病(CMCD) [53] ,体内干扰素在CCL3介导的嗜中性粒细胞招募中的关键作用 [54] 。目前也有些关于免疫细胞的报道,比如,巨噬细胞中TNF-alpha的作用 [55] ,单核细胞和巨噬细胞中SP对PPAR-gamma蛋白表达的影响 [56] ,免疫球蛋白IgG的Fc片段衍生的多肽Tregitopes在激活自然调节T细胞中的作用 [57] ,NK细胞中选择素的作用 [58] 。此外,也有一些研究组关注于对免疫系统的研究,即固有免疫系统及适应性免疫系统 [59, 60] 。近年来,Kendirgi et al. 一直致力于对抵抗流感病毒A/H5N1的标准线性DNA疫苗的研究 [61] ,而Li et al. 一直在研究新型FMDV疫苗 [5] 。

肿瘤学领域

胎牛血清产品也被用来研究dasatinib(Src激酶抑制剂)对黑色素瘤的治疗效果 [62] ,celecoxib对ICAM-1和VCAM-1表达的影响 [63] ,在人类癌细胞中Foxp3的表达情况 [64] ,人的多发性骨髓瘤中FAS的表达情况 [65] ,雄性激素受体敲除的细胞在前列腺癌骨转移治疗中的作用 [66] ,NO释放的药物JS-K对乳腺癌侵染的作用 [67] ,gankyrin蛋白对p53及Mdm2的作用 [68] ,正常人肾细胞及肾癌中SOSTDC1的表达情况 [69] ,卵巢癌中NF-kappaB对原癌基因PIK3CA的转录调控 [70] ,C6胶质瘤细胞中鼠GFG异构体的表达及亚细胞分布情况 [71] ,胃癌细胞中Aurora激酶A对GSK-3beta的调控机制 [72] 。此外,仍有一些研究组致力于对肿瘤治疗的研究。例如,Barkan et al. 以对细胞骨架的研究为切入点,抑制孤立的休眠肿瘤细胞的迁移生长,达到对肿瘤治疗的目的 [73] ,Sahin et al. 利用计算方法模拟,并实验检测模拟效果,最终针对具有耐trastuzumab抗性的乳腺癌提出了一种可能的治疗方案 [74] 。

病理学领域

一些研究组利用GIBCO品牌胎牛血清产品进行病理学方面的研究。Chae et al. 研究人AQP5蛋白在慢性粒细胞性白血病发病过程中的作用 [75] ,Roberts et al. 研究内皮蛋白pUL36和pUL37在疱疹单一病毒1感染中的作用 [76] ,Konigshoff et al. 一直在研究Wnt/beta-catenin信号通路在先天肺纤维化(IPF)中的表达及活性情况 [77] 。此外,其他一些研究组也在进行相似的研究 [78, 79] 。

神经科学领域

胎牛血清产品也被用来培养神经细胞来研究神经细胞中雌二醇在激活beta-连环蛋白依赖的转录中的作用 [80] ,神经PC12细胞中传播锥虫疾病的寄生虫来源的神经营养因子对类胆碱基因表达的作用 [81] ,中枢神经系统中dysbindin-1蛋白的生理作用 [82] ,软骨酸蛋白1B对轴突延伸的调控作用 [83] ,及核因子kappa-B在MPTP诱导的轻微炎症反应中对选择性诱导神经NO合成酶的作用 [4] 。

病毒学领域

GIBCO品牌的胎牛血清被用来培养细胞,研究疫苗病毒p37与宿主体内LE来源的运输小体生物合成相关蛋白的相互关系 [84] 。

Thermo Scientific HyClone公司

Thermo Fisher Scientific公司是科学研究领域最大的设备及试剂供应商。截止2012年3月22日,该公司的Thermo Scientific HyClone品牌共有19种类型的胎牛血清产品。Thermo Scientific HyClone的胎牛血清产品广泛应用在生物医学研究中。本次调查中,193篇文献中的40篇涉及到该公司的胎牛血清产品。 表格 3列出了这些文献的研究领域分布情况。

研究领域数目
生理学24
免疫学5
肿瘤学5
病毒学3
神经科学1
表四: Thermo Scientific Hyclone品牌血清在各研究领域的文献引用情况。
生理学领域

生理学领域是Thermo Scientific Hyclone的胎牛血清产品应用最多的领域。一些科研工作者利用该公司的胎牛血清来研究细胞内GDPD5在甘油磷酸胆碱(GPC)的渗透调节过程中的作用 [85] ,反式作用位点连接的AP-1元件对BRCA1转录的影响 [86] ,17beta-E2 对线粒体相关的锰超氧化物歧化酶(MnSOD)表达及活性的影响 [87] ,多肽纳米纤维骨架的自装配对伤口愈合的加速作用 [88] ,PM胆固醇对PIP2调节的细胞骨架结构的影响 [89] ,TRBP对PACT激活的调控作用 [90] ,人角膜基质纤维母细胞及肌成纤维细胞中maspin合成的下调机制 [91] ,YKL-40在调节基本纤维母细胞生长因子的生物活性中的作用 [92] ,血清素5HT2受体的生化性质及其在信号传导过程中的作用 [93] ,饱和脂肪酸在DNA损伤反应中的作用 [94] ,SV40大T抗原对基因组完整性及DNA损伤反应的影响 [95] ,及循环张力与核心蛋白聚糖缺陷对3D多细胞胶原基质的影响 [96] 。

免疫学领域

整个调查中,有几篇文献集中在对宿主-微生物相互关系的研究。例如,Mendez-Samperio et al. 研究人上皮细胞中分歧杆菌属芽孢杆菌Calmette-Gu rin对抗菌肽LL-37表达及分泌的影响 [97] ,Round et al. 研究宿主-微生物共生状态建立过程中脆弱拟杆菌的作用 [98] ,及Vaishnava et al. 主要研究宿主-微生物互利共生现象 [99] 。还有一些研究组集中在对实验诱导的自免疫性脑脊髓炎中基质金属蛋白酶-7功能的研究上 [100] ,一些促炎症因子的功能 [101] ,及MNV-1感染控制中MDA5的作用 [102] 。

肿瘤学领域

Beauvais et al. 用该公司的胎牛血清制品研究血管生成及肿瘤生成过程中syndecan-1对alphaVbeta3和alphaVbeta3整合素活化的调控作用 [103] ,Lee et al. 研究针对肿瘤的蒽环霉素化疗方法 [104] ,Shim et al. 研究VES处理的人前列腺癌细胞PC-3中NAG-1表达的调控 [105] ,Casalbore et al. 研究人嗅球来源的成体神经干细胞的肿瘤生成与TERT和NOTCH1激活之间的关系 [106] 。

神经科学领域

只有一篇文章牵涉到神经科学研究,该文章重在研究鼠原代感觉神经元释放END-2的活性依赖机制 [107] 。

病毒学领域

Schultz et al. 用该公司的胎牛血清检测杆状病毒早期复制事件中IE1是否作为基因选择性转录激活子 [12] ,Corti et al. 检测一种新型中和抗体能否与群1及群2 流感A病毒的血凝素相互作用 [108] , Han et al. 研究HIV-1复制过程中APOBEC3G及APOBEC3F和一个内源辅因子的相互作用 [109] 。

Sigma-Aldrich公司

本调查中,有22篇文章提到了Sigma-Aldrich公司的胎牛血清(见表五)。

研究领域数目
肿瘤学9
生理学8
病理学4
免疫学1
表五: Sigma-Aldrich公司胎牛血清在各研究领域的文献引用情况。
肿瘤学领域

共有9篇文献集中在分子肿瘤学研究领域。例如,该公司的胎牛血清产品曾用来研究甲状腺癌中NF-kappaB的靶向基因 [110] ,缺氧在p38 MAPK信号通路激活中的关键作用 [111] ,人肿瘤细胞中miR-27a和miR-451对MDR1/P-糖蛋白表达的调控作用 [112] ,受体酪氨酸激酶(RTKs)及IGF-1信号通路激活的作用 [113] ,乳腺癌中核心蛋白聚糖的抗迁移作用 [114] ,及膀胱癌中FGFR3的致癌性 [115] 。

此外,有些研究组则致力于肿瘤的治疗。例如,Xie et al. 表明通过转录/转录后合成调控环路能够靶向特异的肿瘤细胞 [116] 。近来,Ni Chonghaile et al. 也一直在研究高度线粒体priming对化疗的影响 [13] 。

生理学领域

该公司的胎牛血清产品曾被用于研究人四旋膜蛋白CD81的结构及功能 [117] ,骨骼肌中GNE和alpha-辅肌动蛋白连接的作用 [118] ,octanoylated五肽对饥饿素O-乙酰转移酶(GOAT)的作用 [119] ,钙透过通道TRPC3和TRPC7的上调表达对组成型分泌的影响 [120] ,FUSE连接蛋白对丙型肝炎病毒复制的影响 [121] ,及肌醇焦磷酸酯对ATP浓度的调控作用 [122] 。

病理学领域

2008年间,Verma et al. 以Ccl2-/-/Cx3cr1-/-小鼠为模型,研究年龄相关的斑点再生(AMD)过程中伴侣蛋白ERp29的表达情况 [123] ,Kowalski et al. 也在研究TLR2和TLR4刺激对PBMCs诱导的促炎症因子的表达的影响 [124] ,Kami et al. 在研究心肌再生中gremlin的效果 [125] 。

免疫学领域

Bansal et al. 研究表明,巨噬细胞中PI3K和Notch1信号能够促进PIM2诱导的COX-2和MMP-9的表达 [126] 。

Gemini Bio-Products公司

本调查中,9篇文献提及该公司的产品。Hamanaka et al. 利用该公司的胎牛血清培养细胞来研究ER压力中PERK的作用 [127] ,Kitareewan et al. 研究G0S2在APL的类视黄醇反应中的作用 [128] 。此外,其它相关文献研究的内容如下,T细胞激活 [129] ,CD8+ T细胞反应 [130] ,血管生成及肿瘤生成过程中syndecan-1在alphaVbeta3和alphaVbeta3 integrin激活中的作用 [103] ,神经退行性疾病相关的HFE变异及铁离子调节的机制 [131] ,及皮肤感染的发病机理 [132] 。

Atlanta Biologicals公司

Atlanta Biologicals公司的胎牛血清产品曾被用来研究二次RSV感染中gamma干扰素的有利和不利效应 [133] ,检测密码子12、13及61之外的RAS突变对骨髓恶性肿瘤癌变的影响 [134] ,证明在ER及NF-kappaB信号通路中CBP的剂量是基因特异性信号整合的关键调节子 [135] ,研究MNNG处理引起的DNA修复蛋白向染色质DNA的快速聚集过程 [136] 。

PAA Laboratories公司

PAA Laboratories公司现在已经成为GE医疗的一部分。

该公司的胎牛血清曾被用来研究人绒毛膜癌中MAPK信号通路 [137] ,Hsp90与CDC37分子共同保持致癌的蛋白激酶的活性 [138] 。Burdak-Rothkamm et al. 实验识别ATM作为作为放射诱导DNA损伤的复杂信号通路的一个组成成分 [139] ,Marschner et al. [140] 和 Sprynski et al. [141] 也用该公司的胎牛血清从事生理学研究。

Biological Industries公司

Cardoso et al. 用该公司的产品研究癌症细胞中驱动蛋白5B的作用 [16] ,而Avinoam et al. 研究秀丽线虫融合家族蛋白成员的AFF-1的结构与功能 [142] 。

PAN-Biotech GmbH公司

该公司的胎牛血清制品已被用来研究载脂蛋白对丙型肝炎病毒颗粒合成及装配的影响 [143] ,识别巨噬细胞及树突细胞的克隆源骨髓前体细胞 [144] ,以及证明能够通过光基因转录装置控制术血糖水平 [145] 。

Omega Scientific公司

Witherden et al. 用该公司的胎牛血清产品研究在gamma-deltaT细胞激活过程中粘着结合分子类蛋白作为协同刺激受体的作用 [146] ,Epting et al. 也利用该公司产品研究肌原性前体细胞中Sca-1的定位 [147] 。

其它公司的胎牛血清产品

本调查中提到的胎牛血清产品供应商除了上述公司外,还有其它一些公司,如Equitech-Bio、Biochrom、 Lonza、 Biowest及Clontech等。

Hussain et al. 用Clontech公司的胎牛血清研究活化的突变AKT2能够引发人低血糖症 [148] ,Nold-Petry et al. 用Lonza的胎牛血清研究IL-32对内皮细胞功能的影响 [149] ,Hokaiwado et al. 用Equitech-Bio的胎牛血清研究GST-Pi对雄性激素无关的前列腺癌细胞增殖的影响 [14] , Slipicevic et al. 用Biochrom的胎牛血清研究脂肪酸连接蛋白(FABP7)对黑色素瘤细胞增殖及侵染的影响 [150] , Sato et al. 用Biowest的胎牛血清培养鸡DRG神经元细胞以研究软骨酸蛋白1B在NgR1抑制情况下发挥调控轴突发射的效果 [83] 。

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