淺談什么是胎牛血清?

血清是血液凝固后的液體部分，不含血細胞、纖維蛋白及凝集因子等，含有多種對細胞生長必需的營養因子及大分子物質。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對細胞生長必需的生長因子 [1, 2] 。 胎牛血清也含有一些小分子物質，如氨基酸、糖類、脂類及**類物質。

胎牛血清應用在哺乳動物細胞培養中；因此，胎牛血清應用很廣泛。比如，胎牛血清能夠用在科研、工業生產、**疫苗生產的質量控制及生物技術**的質量控制領域。

與其它類型血清相比，胚胎期動物血清含有更多的生長因子，及較少的gamma球蛋白含量。

圖 1. 美洲鱟（Horseshoe crabs）。

血清是由血液凝固后的上清液制備的；而血漿中含有抗凝劑，所以血漿中含有血液凝固相關的蛋白。

國際血清行業協會（www.serumindustry.org）是血清供銷商的商業組織。 協會規定血清需要利用特定的方法進行檢測，并且檢測結果必須顯示在化驗證明書中。

內**，又叫脂多糖（LPS）或脂寡糖（LOS），存在于革蘭氏陰性菌的細胞外膜。臨床上，內**作為一些革蘭氏陰性菌的診斷提示，如腦膜炎球菌。動物體內，內**能夠導致發熱或者誘導炎癥反應；細胞培養中，內**能夠誘導細胞反應異常。

通常，采用鱟阿米巴樣細胞裂解實驗（LAL檢測）檢測內**水平。LAL是從美洲鱟（見圖一）的血液中制備的，對內**非常靈敏，在微量內**存在情況下就凝結。阿米巴樣細胞相當于哺乳動物的白細胞。

胎牛血清的常規處理方式是熱鈍化，即放在56°水浴30分鐘；其目的是使血清中的補體成分及一些未知的細胞生長抑制分子失活。

與胎牛血清的其它加工方式一樣，熱鈍化處理過程必須小心謹慎，因為血清中的生長因子容易在較高溫度或者較長時間加熱條件下失活，甚至產生沉淀。

Life Technologies公司的Invitrogen及GIBCO品牌的熱鈍化胎牛血清用在細胞培養中，來研究UCP-2及UCP-3 [3] ，核因子kappa-B [4] ，蠶桿狀病毒表達系統中表達FMDV衣殼蛋白 [5] ，亞臨床的移植物抗宿主病 [6] ，IFN-gamma的作用 [7] ，及自噬 [8] 。Thermo Fisher HyClone公司的熱鈍化胎牛血清用來研究能引起嚴重急性呼吸綜合征的冠狀病毒的核衣殼蛋白 [9] ，NO [10] ，激活素C [11] ，及IE1 [12] 。Sigma公司的熱鈍化胎牛血清用來研究化療 [13] ，而Equitech-Bio公司的熱鈍化胎牛血清用來研究谷胱甘肽巰基轉移酶Pi [14] 。淺談什么是胎牛血清?

活性炭能夠結合脂溶性分子，因此能夠用來除去**類物質，例如，胎牛血清中的雌二醇、孕酮、皮質醇、睪酮及T3和T4分子。血清中這些脂類**能夠干擾**檢測及胰島素檢測方法。

Thermo Scientific的活性炭吸附的胎牛血清（csFCS）被用來識別TGF-beta家族成員 [15] 。

透析處理能夠除去胎牛血清中所有分子量小于10,000Da的分子。例如，**、細胞因子、葡萄糖、氨基酸還有許多其它分子，都能夠被除去。透析也能除去胎牛血清中的***及其它內源分子。

Life Technologies的透析處理的胎牛血清被用來研究驅動蛋白5B [16] 及丙酮酸激酶M2失活 [17] ，Thermo Fisher HyClone的透析處理胎牛血清用來研究KRAS信號通路的突變 [18] ，而Sigma公司的透析處理胎牛血清用來研究PAH代謝情況 [19] 。

Gamma照射能夠對胎牛血清進行**處理。胎牛血清常規的**方法是多次透過0.1um的濾膜以消除微生物污染，而gamma照射能夠鈍化牛群中常見的病毒。但是，一些類型的病毒能夠抵抗gamma照射，比如細小病毒。

低IgG的胎牛血清適用于抗體生產及相關領域。淺談什么是胎牛血清?

圖 2. Gibco Life Technologies許可。

與一般細胞培養相比，干細胞培養對生長因子有更嚴格的要求。由于胎牛血清中的一些生長因子促進干細胞分化，所以，需要檢測各批胎牛血清在保持不同類型干細胞的全能性方面的性能。

胎牛血清的主要來源地就是那些大量養牛的國家，包括美國、澳大利亞、新西蘭、加拿大及南美和中美洲地區，由政府機構管理血清制品的生產及收集過程。

胎牛血清分為美國農業部（USDA）級和歐盟級。其中，美國農業部級胎牛血清沒有瘋牛?。˙SE）及口蹄疫（FMD）污染，能夠出口到任何國家，而歐盟級胎牛血清只在大部分歐洲國家及亞洲國家銷售。

胎牛血清一般是在-5°到-20°之間凍存，2°到8°之間融化。一個比較好的辦法是，將胎牛血清分裝到50ml管中，這樣做就避免了反復凍融對血清造成的危害。血清融化后常出現沉淀，可能是血清蛋白變性造成的。簡短離心即可除去沉淀，并且，這些沉淀一般不會影響血清質量的。

許多供應商提供不同等級、不同來源及不同處理的胎牛血清。而胎牛血清是一種復雜的混合物，不同供應商的不同批次產品肯定會有一些不同。因此，有必要建立一套選擇、評估胎牛血清的機制。

一般來說，可以通過參考相關研究文獻選擇適合的胎牛血清。

Labome對212篇科研文獻做了關于胎牛血清使用情況的調查。

圖 3. Thermo Fisher Scientific許可。

這些文獻是Labome的10,000篇文獻中隨機抽取的一小部分，選自開放訪問的雜志，出版時間介于2006和2014年之間。

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

Life Technologies公司是一家全球性的生物技術公司，總部設在加利福尼亞州Carlsbad市。該公司是由Invitrogen公司和Applied Biosystems公司于2008年合并而來。GIBCO是該公司的一個品牌，主要提供細胞培養產品，比如細胞培養基、胎牛血清及其它相關試劑等。截止2012年3月，該公司共有大概34種類型的胎牛血清產品。

Labome的調查結果表明，有42%（193篇中的82篇）的文獻涉及到GIBCO的胎牛血清。表三列出了這些文獻的研究領域分布情況。

生理學是熱門的研究領域，且大部分文獻聚焦在對生理過程中基因功能的研究。例如，該公司的胎牛血清產品已經被用來研究人小膠質細胞和腦組織中白蛋白的表達情況 [20] ，層粘連蛋白連接過程中alpha-肌營養**蛋白聚糖的O-甘露糖磷酸化的功能 [21] ，APE1蛋白殘基D70在對DNA的3'重末端剪切中的新作用 [22] ，果蠅中CENH3在著絲粒形成中的作用 [23] ，IGF1異構體及功能 [24] ，NO信號通路中二醇的作用 [25] ，PKC在用異氟烷預誘導產生保護功能的過程中的作用 [26] ，RARalpha上游因子對多向性遺傳網絡的調控作用 [27] ，在TNF誘導的破骨細胞的前體細胞（OCPs）轉移過程中SDF-1/CXCR4軸的作用 [28] ，C端固定的G蛋白alpha亞基對其再循環速率的影響 [29] ，SRp20和CUG-BP1在調控胰島素受體外顯子11的選擇性剪切中的功能 [30] ，生長因子對ADAMTS-4表達的影響 [31] ，在ES細胞保持一致性過程中PRC2的作用 [32] ，轉錄因子Ets-1在調控NPR-A表達過程中的作用 [33] ，在mTOR信號通路中H+-ATPase的作用 [34] ，LC細胞中ET-1對膠原合成及分泌的影響 [35] ，及**細胞中線粒體K+通道Kv1.3在Bax誘導凋亡過程中的作用 [36] 。

此外，胎牛血清產品也被用來研究一些**的效果，比如說，局部**抑制劑 [37] 以及組蛋白去乙酰化酶（HDAC）抑制劑LAQ824 [38] 。一些研究者利用胎牛血清產品研究凋亡進程。例如，Atapattu et al. [39] 和Lam et al. [40] 用胎牛血清培養細胞研究凋亡過程中的具體細節。還有一些科學家研究分子機制 [41-45] 以及某些分子在相應信號通路中的作用 [46-48] ，例如，鈣粘素依賴的粘著連接（AJ）與Wnt信號通路之間的相互關系 [47] 、CHIP在聚集小體信號通路中的作用 [48] 。

**學領域也是一個熱門領域。胎牛血清產品曾被用來研究IL-23在起始炎癥反應中的作用 [49] ，IL-23及IL-12在Dectin-1和TLR信號通路共激活中的作用 [50] ，炎癥小體識別在適應性**對流感病毒識別中的作用 [51] ，特異的CD40抗體誘導的**激活 [52] ，IL-17的**缺陷導致的慢性粘膜皮膚***病（CMCD） [53] ，體內干擾素在CCL3介導的嗜中性粒細胞招募中的關鍵作用 [54] 。目前也有些關于**細胞的報道，比如，巨噬細胞中TNF-alpha的作用 [55] ，單核細胞和巨噬細胞中SP對PPAR-gamma蛋白表達的影響 [56] ，**球蛋白IgG的Fc片段衍生的多肽Tregitopes在激活自然調節T細胞中的作用 [57] ，NK細胞中選擇素的作用 [58] 。此外，也有一些研究組關注于對**系統的研究，即固有**系統及適應性**系統 [59, 60] 。近年來，Kendirgi et al. 一直致力于對抵抗流感病毒A/H5N1的標準線性DNA疫苗的研究 [61] ，而Li et al. 一直在研究新型FMDV疫苗 [5] 。

胎牛血清產品也被用來研究dasatinib（Src激酶抑制劑）對黑色素瘤的**效果 [62] ，celecoxib對ICAM-1和VCAM-1表達的影響 [63] ，在人類癌細胞中Foxp3的表達情況 [64] ，人的多發性骨髓瘤中FAS的表達情況 [65] ，雄性**受體敲除的細胞在前列腺癌骨轉移**中的作用 [66] ，NO釋放的**JS-K對乳腺癌侵染的作用 [67] ，gankyrin蛋白對p53及Mdm2的作用 [68] ，正常人腎細胞及腎癌中SOSTDC1的表達情況 [69] ，卵巢癌中NF-kappaB對原癌基因PIK3CA的轉錄調控 [70] ，C6膠質瘤細胞中鼠GFG異構體的表達及亞細胞分布情況 [71] ，胃癌細胞中Aurora激酶A對GSK-3beta的調控機制 [72] 。此外，仍有一些研究組致力于對腫瘤**的研究。例如，Barkan et al. 以對細胞骨架的研究為切入點，抑制孤立的休眠腫瘤細胞的遷移生長，達到對腫瘤**的目的 [73] ，Sahin et al. 利用計算方法模擬，并實驗檢測模擬效果，終針對具有耐trastuzumab抗性的乳腺癌提出了一種可能的**方案 [74] 。

一些研究組利用GIBCO品牌胎牛血清產品進行病理學方面的研究。Chae et al. 研究人AQP5蛋白在慢性粒細胞性白血病發病過程中的作用 [75] ，Roberts et al. 研究內皮蛋白pUL36和pUL37在皰疹單一病毒1感染中的作用 [76] ，Konigshoff et al. 一直在研究Wnt/beta-catenin信號通路在先天肺纖維化（IPF）中的表達及活性情況 [77] 。此外，其他一些研究組也在進行相似的研究 [78, 79] 。

胎牛血清產品也被用來培養神經細胞來研究神經細胞中雌二醇在激活beta-連環蛋白依賴的轉錄中的作用 [80] ，神經PC12細胞中傳播錐蟲**的寄生蟲來源的神經營養因子對類膽堿基因表達的作用 [81] ，**神經系統中dysbindin-1蛋白的生理作用 [82] ，軟骨酸蛋白1B對軸突延伸的調控作用 [83] ，及核因子kappa-B在MPTP誘導的輕微炎癥反應中對選擇性誘導神經NO合成酶的作用 [4] 。

GIBCO品牌的胎牛血清被用來培養細胞，研究疫苗病毒p37與宿主體內LE來源的運輸小體生物合成相關蛋白的相互關系 [84] 。

Thermo Fisher Scientific公司是科學研究領域大的設備及試劑供應商。截止2012年3月22日，該公司的Thermo Scientific HyClone品牌共有19種類型的胎牛血清產品。Thermo Scientific HyClone的胎牛血清產品廣泛應用在生物醫學研究中。本次調查中，193篇文獻中的40篇涉及到該公司的胎牛血清產品。 表格 3列出了這些文獻的研究領域分布情況。

生理學領域是Thermo Scientific Hyclone的胎牛血清產品應用多的領域。一些科研工作者利用該公司的胎牛血清來研究細胞內GDPD5在甘油磷酸膽堿（GPC）的滲透調節過程中的作用 [85] ，反式作用位點連接的AP-1元件對BRCA1轉錄的影響 [86] ，17beta-E2 對線粒體相關的錳超氧化物歧化酶（MnSOD）表達及活性的影響 [87] ，多肽納米纖維骨架的自裝配對傷口**的加速作用 [88] ，PM膽固醇對PIP2調節的細胞骨架結構的影響 [89] ，TRBP對PACT激活的調控作用 [90] ，人角膜基質纖維母細胞及肌成纖維細胞中maspin合成的下調機制 [91] ，YKL-40在調節基本纖維母細胞生長因子的生物活性中的作用 [92] ，血清素5HT2受體的生化性質及其在信號傳導過程中的作用 [93] ，飽和脂肪酸在DNA損傷反應中的作用 [94] ，SV40大T抗原對基因組完整性及DNA損傷反應的影響 [95] ，及循環張力與核心蛋白聚糖缺陷對3D多細胞膠原基質的影響 [96] 。

整個調查中，有幾篇文獻集中在對宿主-微生物相互關系的研究。例如，Mendez-Samperio et al. 研究人上皮細胞中分歧桿菌屬芽孢桿菌Calmette-Gu rin對**肽LL-37表達及分泌的影響 [97] ，Round et al. 研究宿主-微生物共生狀態建立過程中脆弱擬桿菌的作用 [98] ，及Vaishnava et al. 主要研究宿主-微生物互利共生現象 [99] 。還有一些研究組集中在對實驗誘導的自**性腦脊髓炎中基質金屬蛋白酶-7功能的研究上 [100] ，一些促炎癥因子的功能 [101] ，及MNV-1感染控制中MDA5的作用 [102] 。

Beauvais et al. 用該公司的胎牛血清制品研究血管生成及腫瘤生成過程中syndecan-1對alphaVbeta3和alphaVbeta3整合素活化的調控作用 [103] ，Lee et al. 研究針對腫瘤的蒽環霉素化療方法 [104] ，Shim et al. 研究VES處理的人前列腺癌細胞PC-3中NAG-1表達的調控 [105] ，Casalbore et al. 研究人嗅球來源的成體神經干細胞的腫瘤生成與TERT和NOTCH1激活之間的關系 [106] 。

只有一篇文章牽涉到神經科學研究，該文章重在研究鼠原代感覺神經元釋放END-2的活性依賴機制 [107] 。

Schultz et al. 用該公司的胎牛血清檢測桿狀病毒早期復制事件中IE1是否作為基因選擇性轉錄激活子 [12] ，Corti et al. 檢測一種新型中和抗體能否與群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復制過程中APOBEC3G及APOBEC3F和一個內源輔因子的相互作用 [109] 。

本調查中，有22篇文章提到了Sigma-Aldrich公司的胎牛血清（見表五）。

共有9篇文獻集中在分子腫瘤學研究領域。例如，該公司的胎牛血清產品曾用來研究甲狀腺癌中NF-kappaB的靶向基因 [110] ，缺氧在p38 MAPK信號通路激活中的關鍵作用 [111] ，人腫瘤細胞中miR-27a和miR-451對MDR1/P-糖蛋白表達的調控作用 [112] ，受體酪氨酸激酶（RTKs）及IGF-1信號通路激活的作用 [113] ，乳腺癌中核心蛋白聚糖的抗遷移作用 [114] ，及膀胱癌中FGFR3的致癌性 [115] 。

此外，有些研究組則致力于腫瘤的**。例如，Xie et al. 表明通過轉錄/轉錄后合成調控環路能夠靶向特異的腫瘤細胞 [116] 。近來，Ni Chonghaile et al. 也一直在研究高度線粒體priming對化療的影響 [13] 。

該公司的胎牛血清產品曾被用于研究人四旋膜蛋白CD81的結構及功能 [117] ，骨骼肌中GNE和alpha-輔肌動蛋白連接的作用 [118] ，octanoylated五肽對饑餓素O-乙酰轉移酶（GOAT）的作用 [119] ，鈣透過通道TRPC3和TRPC7的上調表達對組成型分泌的影響 [120] ，FUSE連接蛋白對丙型肝炎病毒復制的影響 [121] ，及肌醇焦磷酸酯對ATP濃度的調控作用 [122] 。

2008年間，Verma et al. 以Ccl2-/-/Cx3cr1-/-小鼠為模型，研究年齡相關的斑點再生（AMD）過程中伴侶蛋白ERp29的表達情況 [123] ，Kowalski et al. 也在研究TLR2和TLR4刺激對PBMCs誘導的促炎癥因子的表達的影響 [124] ，Kami et al. 在研究心肌再生中gremlin的效果 [125] 。

Bansal et al. 研究表明，巨噬細胞中PI3K和Notch1信號能夠促進PIM2誘導的COX-2和MMP-9的表達 [126] 。

本調查中，9篇文獻提及該公司的產品。Hamanaka et al. 利用該公司的胎牛血清培養細胞來研究ER壓力中PERK的作用 [127] ，Kitareewan et al. 研究G0S2在APL的類視黃醇反應中的作用 [128] 。此外，其它相關文獻研究的內容如下，T細胞激活 [129] ，CD8+ T細胞反應 [130] ，血管生成及腫瘤生成過程中syndecan-1在alphaVbeta3和alphaVbeta3 integrin激活中的作用 [103] ，神經退行性**相關的HFE變異及鐵離子調節的機制 [131] ，及****的發病機理 [132] 。

Atlanta Biologicals公司的胎牛血清產品曾被用來研究二次RSV感染中gamma干擾素的有利和不利效應 [133] ，檢測密碼子12、13及61之外的RAS突變對骨髓惡性腫瘤癌變的影響 [134] ，證明在ER及NF-kappaB信號通路中CBP的劑量是基因特異性信號整合的關鍵調節子 [135] ，研究MNNG處理引起的DNA修復蛋白向染色質DNA的快速聚集過程 [136] 。

PAA Laboratories公司現在已經成為GE醫療的一部分。

該公司的胎牛血清曾被用來研究人絨毛膜癌中MAPK信號通路 [137] ，Hsp90與CDC37分子共同保持致癌的蛋白激酶的活性 [138] 。Burdak-Rothkamm et al. 實驗識別ATM作為作為放射誘導DNA損傷的復雜信號通路的一個組成成分 [139] ，Marschner et al. [140] 和 Sprynski et al. [141] 也用該公司的胎牛血清從事生理學研究。

Cardoso et al. 用該公司的產品研究癌癥細胞中驅動蛋白5B的作用 [16] ，而Avinoam et al. 研究秀麗線蟲融合家族蛋白成員的AFF-1的結構與功能 [142] 。

該公司的胎牛血清制品已被用來研究載脂蛋白對丙型肝炎病毒顆粒合成及裝配的影響 [143] ，識別巨噬細胞及樹突細胞的克隆源骨髓前體細胞 [144] ，以及證明能夠通過光基因轉錄裝置控制術血糖水平 [145] 。

Witherden et al. 用該公司的胎牛血清產品研究在gamma-deltaT細胞激活過程中粘著結合分子類蛋白作為協同刺激受體的作用 [146] ，Epting et al. 也利用該公司產品研究肌原性前體細胞中Sca-1的定位 [147] 。

本調查中提到的胎牛血清產品供應商除了上述公司外，還有其它一些公司，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變AKT2能夠引發人低血糖癥 [148] ，Nold-Petry et al. 用Lonza的胎牛血清研究IL-32對內皮細胞功能的影響 [149] ，Hokaiwado et al. 用Equitech-Bio的胎牛血清研究GST-Pi對雄性**無關的前列腺癌細胞增殖的影響 [14] ， Slipicevic et al. 用Biochrom的胎牛血清研究脂肪酸連接蛋白（FABP7）對黑色素瘤細胞增殖及侵染的影響 [150] ， Sato et al. 用Biowest的胎牛血清培養雞DRG神經元細胞以研究軟骨酸蛋白1B在NgR1抑制情況下發揮調控軸突發射的效果 [83] 。

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