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γ生育三烯酚的生物活性及其作用機(jī)理研究進(jìn)展

2016-09-10 06:16牛玲玲徐偉麗何勝華米雅清劉巧紅魯兆新
食品工業(yè)科技 2016年5期
關(guān)鍵詞:生育膽固醇抗氧化

牛玲玲,徐偉麗,*,劉 蕾,何勝華,米雅清,劉巧紅,魯兆新

(1.哈爾濱工業(yè)大學(xué)化工學(xué)院食品科學(xué)與工程系,黑龍江哈爾濱 150090;2.佳木斯大學(xué)基礎(chǔ)醫(yī)學(xué)院,黑龍江佳木斯 154007)

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牛玲玲1,徐偉麗1,*,劉蕾2,何勝華1,米雅清1,劉巧紅1,魯兆新1

(1.哈爾濱工業(yè)大學(xué)化工學(xué)院食品科學(xué)與工程系,黑龍江哈爾濱 150090;2.佳木斯大學(xué)基礎(chǔ)醫(yī)學(xué)院,黑龍江佳木斯 154007)

維生素E包括生育酚和生育三烯酚兩個(gè)亞族,每個(gè)亞族有四種單體(α,β,γ和δ)。相較于其他維生素E單體,γ-生育三烯酚(γ-T3)具有更好的抗腫瘤、神經(jīng)保護(hù)、降低膽固醇、抗炎以及抑制脂肪產(chǎn)生的作用。本文旨在綜述γ-T3的生物活性及其作用機(jī)理,提出目前γ-T3生物活性研究中存在的問題,并對(duì)其研究方向進(jìn)行展望。

γ-生育三烯酚,生理活性,作用機(jī)理

維生素E包括生育酚和生育三烯酚兩個(gè)亞族。生育酚和生育三烯酚又可細(xì)分為四種單體(α,β,γ和δ),見表1[1]。生育三烯酚區(qū)別生育酚在于脂肪族側(cè)鏈,生育酚有一個(gè)含葉綠基的側(cè)鏈附著在色原烷醇核上,而生育三烯酚側(cè)鏈含有不飽和鍵,并形成異戊二烯鏈[2]。富含油脂的農(nóng)產(chǎn)品和菜籽油是維生素E的主要天然來源,生育三烯酚也廣泛存在于天然植物中,其中棕櫚油、大米麩皮中含量最高;椰子油、可可油、豆油、大麥和小麥中也含有一定量生育三烯酚[3]。α-生育三烯酚(α-T3)是生育三烯酚在燕麥(燕麥L.)和大麥中的主要形式,β-生育三烯酚(β-T3)是在帶殼和去殼小麥中發(fā)現(xiàn)的生育三烯酚的主要形式[4]。棕櫚油和米糠油是γ-T3重要的天然來源[5-6]。近年來,隨著對(duì)生育三烯酚在人體內(nèi)作用和功能的闡釋,生育三烯酚已成為國外研究的熱點(diǎn)之一。已有研究表明,生育三烯酚在抗癌和抗氧化等很多方面的作用都優(yōu)于生育酚,但研究并不全面和深入,而且相比于其他的維生素E單體,γ-T3擁有更好的抗腫瘤、抗氧化、降低膽固醇、抗炎和減少脂肪產(chǎn)生等作用[7-8]。所以本文將重點(diǎn)綜述γ-生育三烯酚優(yōu)越的生理活性及其可能的作用機(jī)理。

1 抗氧化作用

維生素E是一種很強(qiáng)的脂溶性抗氧化劑,具有直接猝滅自由基的能力和作為膜穩(wěn)定劑的功能[9]。研究表明,在維生素E的異構(gòu)體中,α-T3的抗氧化能力最強(qiáng),這可能與它自身的結(jié)構(gòu)有關(guān),特別是其位于側(cè)鏈上的不飽和雙鍵[10]。

Newaz MA等[11]研究了γ-T3對(duì)自發(fā)性高血壓大鼠(SHR)脂質(zhì)過氧化和總抗氧化狀態(tài)的作用效果。實(shí)驗(yàn)分為正常小鼠組、未經(jīng)處理的高血壓小鼠組和15、30、150 mg/kg的γ-T3處理組,每兩周測量一次血壓,為期3個(gè)月。結(jié)果表明,γ-T3作用三個(gè)月后,3個(gè)處理組高血壓小鼠血管中脂質(zhì)過氧化物濃度都顯著降低(處理組1:p<0.05;處理組2:p<0.001;處理組3:p<0.005);血漿中的脂質(zhì)過氧化物濃度都降低,其中處理組1顯著降低(p=0.034);血漿超氧化物歧化酶(SOD)活性得到顯著改善(處理組1:p<0.001;處理組2:p<0.05;處理組3:p<0.001)。在正常小鼠和SHR中總抗氧化狀態(tài)(TAS)與SOD水平呈顯著負(fù)相關(guān),血管和血漿脂質(zhì)過氧化物濃度和血壓成顯著正相關(guān)。表明γ-T3具有顯著降低血漿脂質(zhì)過氧化物濃度和提高總抗氧化狀態(tài)的能力。

表1 生育酚和生育三烯酚的化學(xué)結(jié)構(gòu)[1]Table 1 The chemical structure of tocopherol and tocotrienols[1]

Abd Manan N等[12]探討了γ-T3對(duì)脂質(zhì)過氧化、抗氧化酶活性、暴露在過氧化氫(H2O2)環(huán)境中的成骨細(xì)胞凋亡的影響。成骨細(xì)胞預(yù)先經(jīng)1、10、100 μmol/L的γ-T3處理24 h,然后暴露在490 μmol/L(IC50)H2O2環(huán)境。結(jié)果表明,γ-T3可以防止通過H2O2誘導(dǎo)的丙二醛(MDA)含量的升高,并且呈劑量依賴性。所有劑量的γ-T3都能夠防止SOD、過氧化氫酶(CAT)活性的降低。1、10 μmol/L的γ-T3能夠減少成骨細(xì)胞凋亡,而高劑量的(100 μmol/L)γ-T3與H2O2相比具有更強(qiáng)的凋亡誘導(dǎo)作用。即低劑量的γ-T3能夠降低H2O2毒性,對(duì)成骨細(xì)胞提供保護(hù),但高劑量則會(huì)引起細(xì)胞毒性。

2 抗腫瘤作用

γ-T3可以誘導(dǎo)多種類型癌細(xì)胞發(fā)生凋亡[13-14]。其抑制腫瘤作用在不同類型癌細(xì)胞中也表現(xiàn)出不同的作用機(jī)制[15-17]。目前,對(duì)于γ-T3抗腫瘤的研究主要集中在人乳腺癌[18]、肝癌[19]、結(jié)腸癌[20]、胃癌[21]、前列腺癌[22]等方面。

Patacsil D等[23]的研究證明在MDA-MB231和MCF-7乳腺癌細(xì)胞中γ-T3明顯是通過PARP裂解和caspase-7活化誘導(dǎo)細(xì)胞凋亡。γ-T3處理后,MCF-7細(xì)胞中參與細(xì)胞生長增殖、細(xì)胞死亡、細(xì)胞周期、細(xì)胞發(fā)育和細(xì)胞運(yùn)動(dòng)的多個(gè)基因的表達(dá)發(fā)生改變。表明γ-T3能夠激活PERK和pIRE1α途徑來誘導(dǎo)ER應(yīng)激反應(yīng)。同時(shí)采用siRNA法證明激活轉(zhuǎn)錄因子3(ATF3)在γ-T3誘導(dǎo)的細(xì)胞凋亡中起重要作用。研究結(jié)果證明γ-T3調(diào)控內(nèi)質(zhì)網(wǎng)應(yīng)激信號(hào)傳導(dǎo),而且確定ATF3為γ-T3在乳腺癌細(xì)胞中的分子靶標(biāo)。

Jiang Q等[24]的研究表明γ-T3處理可以誘導(dǎo)人前列腺癌細(xì)胞PC-3和LNCaP的凋亡、壞死和自噬。他們在探索γ-T3引發(fā)效應(yīng)的潛在機(jī)制時(shí)發(fā)現(xiàn),γ-T3處理可導(dǎo)致細(xì)胞內(nèi)的二氫鞘氨醇和二氫神經(jīng)酰胺含量顯著增長,但是初始鞘脂合成途徑中鞘脂的中間體對(duì)神經(jīng)酰胺或鞘氨醇沒有影響。通過γ-T3誘導(dǎo)產(chǎn)生的鞘脂遠(yuǎn)比γ-生育酚誘導(dǎo)產(chǎn)生的含量高。這些發(fā)現(xiàn)提供了一種γ-T3誘導(dǎo)癌細(xì)胞死亡的分子機(jī)制,認(rèn)為細(xì)胞內(nèi)二氫鞘氨醇和二氫神經(jīng)鞘氨醇含量的升高可能是一個(gè)新的抗癌機(jī)制。

孫文廣等[21]的研究發(fā)現(xiàn)γ-T3不僅抑制SGC-7901細(xì)胞的核分裂,降低細(xì)胞的增殖活力,而且抑制DNA的合成。其抑制機(jī)理與γ-T3誘導(dǎo)caspase-3活力,參與調(diào)節(jié)Bax及Bcl-2蛋白表達(dá)有關(guān)。已有的研究發(fā)現(xiàn)caspase-3酶切Bcl-2產(chǎn)生的N34Bcl-2片斷被轉(zhuǎn)染到BHK細(xì)胞中,也可以易位到線粒體上,促進(jìn)細(xì)胞色素C的釋放,加速細(xì)胞凋亡。有報(bào)道稱富含γ-T3的棕櫚油可以增強(qiáng)caspase-3活力,誘導(dǎo)人乳腺癌和人結(jié)腸癌細(xì)胞凋亡[25-26]。γ-T3可以通過促進(jìn)Bax蛋白表達(dá),降低Bcl-2蛋白表達(dá),影響二者比值,引起細(xì)胞凋亡,這與Agarwal MK等[16]報(bào)道的富含γ-T3的棕櫚油能夠升高Bax/Bcl-2比值,誘導(dǎo)人結(jié)腸癌細(xì)胞凋亡相一致。

3 神經(jīng)保護(hù)作用

研究表明,生育三烯酚可保護(hù)神經(jīng)細(xì)胞免受氧化損傷[27]。納摩爾濃度的α-生育三烯酚是唯一可通過調(diào)節(jié)細(xì)胞死亡的特定介質(zhì)防止誘導(dǎo)的神經(jīng)變性的試劑[28]。微摩爾數(shù)量的生育三烯酚能抑制HMG-CoA還原酶活性和肝酶負(fù)責(zé)的膽固醇合成[29]。其中,γ-T3對(duì)于神經(jīng)保護(hù)作用的研究較少。

Then等[30]研究了γ-T3對(duì)由H2O2誘導(dǎo)的氧化應(yīng)激的唐氏綜合癥(DS)神經(jīng)元的保護(hù)作用。實(shí)驗(yàn)分為6組,對(duì)照組、H2O2組、γ-T3預(yù)處理的H2O2組、γ-T3處理組、α-生育酚預(yù)處理的H2O2組和α-生育酚處理組。γ-T3培養(yǎng)24 h能夠使DS神經(jīng)元的凋亡減少10%,但是γ-T3在較長的潛伏期(14 d)和在濃度≥100 μmol/L時(shí)有細(xì)胞毒性。γ-T3通過減少H2O2生成的活性氧來作為自由基清除劑。但是DS神經(jīng)元先經(jīng)H2O2處理后,再采用γ-T3或α-生育酚進(jìn)行處理,Bcl-2/Bax比值都降低。這表明,γ-T3或α-生育酚預(yù)處理都不足以保護(hù)DS神經(jīng)元來自H2O2誘導(dǎo)的氧化傷害,不能促進(jìn)DS細(xì)胞的存活,而是誘導(dǎo)DS細(xì)胞的凋亡。

4 降低膽固醇作用

膽固醇生物合成的關(guān)鍵酶是3-羥基-3-甲基戊二酰輔酶A還原酶(HMGR),該酶位于內(nèi)質(zhì)網(wǎng)上。有研究表明,生育三烯酚降低膽固醇的作用并不是作為合成途徑中的抑制因子,而是通過抑制肝臟中HMGR的活性來實(shí)現(xiàn)的[31]。生育三烯酚能有效抑制甲羥戊酸的合成,進(jìn)一步減少肝組織中膽固醇合成中間產(chǎn)物焦磷酸法尼酯和焦磷酸牻牛兒酯的含量,從而減少膽固醇的合成[32]。

目前,細(xì)胞實(shí)驗(yàn)表明γ-T3具有降脂作用,特別是通過抑制HMGR活性來降低膽固醇含量[33]。體外實(shí)驗(yàn)表明γ-T3降低膽固醇合成的能力比α-T3高30倍[34]。另外γ-T3還能夠促進(jìn)膽固醇的代謝[35]。最近的研究發(fā)現(xiàn)γ-、δ-T3能激活HMGR蛋白泛素化,從而促進(jìn)其降解,并具有阻斷膽固醇調(diào)節(jié)因子結(jié)合蛋白的能力[36]。

然而,Hasselwander 等[37]探究了生育三烯酚在膽固醇喂養(yǎng)的兔子中降膽固醇、抗動(dòng)脈粥樣硬化的效果,給兔飼喂基礎(chǔ)日糧(對(duì)照組),實(shí)驗(yàn)組輔以γ-T3,α-生育三烯酚乙酸酯,混合生育三烯酚或α-生育三烯酚,飼喂12周。結(jié)果表明,所有處理組均導(dǎo)致血漿中生育三烯酚含量顯著增加,但是對(duì)血脂和動(dòng)脈粥樣硬化病變沒有顯著的影響。血漿中γ-和α-T3含量顯著低于α-生育酚。此研究對(duì)γ-T3是否具有降低膽固醇能力給出了不明確、模棱兩可的結(jié)果,γ-T3能夠降低膽固醇能力存在爭議。生育三烯酚降低膽固醇的能力有待驗(yàn)證,推測這可能與血漿生育三烯酚水平有關(guān)。

5 抗炎作用

角鯊烯氫過氧化物(SQ-OOH)是主要的角鯊烯(SQ)過氧化產(chǎn)物。SQ-OOH能夠促進(jìn)炎癥基因如白介素和環(huán)氧合酶-2(COX-2)的表達(dá)。NakagawaK等[38]的研究表明,γ-T3可以抑制HaCaT細(xì)胞中SQ-OOH誘導(dǎo)的活性氧(ROS)產(chǎn)生、核因子κB(NF-κB)活化、COX-2的mRNA和蛋白表達(dá),以及前列腺素E-2(PGE2)的產(chǎn)生。即γ-T3可通過抑制SQ-OOH誘導(dǎo)的ROS和炎性介質(zhì)發(fā)揮抗炎作用。

作為預(yù)防的策略,含有γ-T3的飲食可以減輕炎癥。另外,動(dòng)物喂養(yǎng)含有γ-T3的飲食后γ-T3會(huì)積聚在皮膚,能夠抑制皮膚由紫外和臭氧引起的氧化應(yīng)激反應(yīng)[39-41]。大鼠損傷的研究中γ-T3可能具有保護(hù)皮膚免受紫外線引起的炎癥的作用[42-43]。研究發(fā)現(xiàn),某些病癥在進(jìn)展的重要階段(如糖尿病性視網(wǎng)膜病,類風(fēng)濕關(guān)節(jié)炎和癌癥),γ-T3可以抑制病理性血管產(chǎn)生[44]。

6 抑制脂肪形成,減輕人類增生性肥胖

Zhao L等[45]證實(shí)γ-T3可通過作用于人脂肪干細(xì)胞(hASCs)的C/EBP下游和C/EBP上游特異性抑制脂肪細(xì)胞分化的早期階段,從而抑制脂肪生成。利用AMPK的顯性負(fù)突變體阻斷AMPK不能使γ-T3介導(dǎo)的自噬作用正常化,表明γ-T3增強(qiáng)的自噬活動(dòng)獨(dú)立于AMPK激活。結(jié)果表明,γ-T3激活A(yù)MPK和自噬作用有助于抑制hASCs細(xì)胞分化成脂肪細(xì)胞,從而發(fā)揮抑制脂肪形成作用,這提供了一種對(duì)分子機(jī)制研究的新穎見解。因此,γ-T3可能會(huì)構(gòu)成一個(gè)新的飲食途徑來減輕人類增生性肥胖。

Muto等[46]研究了維生素E單體(特別是γ-T3)對(duì)肝甘油三酯(TG)積累和三種與大鼠原代肝細(xì)胞有關(guān)的脂肪酸代謝酶的效果。研究結(jié)果表明,γ-T3能顯著降低正常肝細(xì)胞中TG的含量,抑制棕櫚酸(PA)誘導(dǎo)的兩個(gè)C/EBP同源蛋白質(zhì)(CHOP)和SREBP-1c的基因表達(dá),增加肉堿棕櫚1(CPT1A)mRNA的表達(dá),減少固醇調(diào)節(jié)元件結(jié)合蛋白1C(SREBP-1c)mRNA的表達(dá)??傊?γ-T3可阻止肝細(xì)胞脂肪變性,并改善內(nèi)質(zhì)網(wǎng)應(yīng)激作用和肝臟后續(xù)的炎癥。

7 結(jié)語

相比于其他的維生素E單體,抗氧化方面α-生育三烯酚研究較多且抗氧化能力最強(qiáng),γ-T3表現(xiàn)出更好的抗癌作用[7-8]。而γ-T3抗氧化、神經(jīng)保護(hù)、抗炎和減少脂肪產(chǎn)生等方面的作用研究較少。其中,γ-T3發(fā)揮抗癌和抗氧化作用與其自身濃度相關(guān),低濃度促進(jìn)細(xì)胞生長發(fā)育,高濃度則具有細(xì)胞毒性。γ-T3降低膽固醇的作用還沒有取得明確的結(jié)果。另外,γ-T3在腸道內(nèi)通透性較差,導(dǎo)致生物利用率很低,不利于它在人體中發(fā)揮作用。因此,應(yīng)總結(jié)γ-T3生物利用率較低的原因,尋找提高γ-T3生物利用率的方法,進(jìn)一步研究γ-T3發(fā)揮各種作用的機(jī)制,為通過膳食干預(yù)而預(yù)防和治療癌癥、動(dòng)脈粥樣硬化和肥胖等疾病提供依據(jù),加強(qiáng)植物資源的開發(fā)利用,提高人類生存質(zhì)量。

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Advances in biological activity ofγ-tocotrienol and its mechanism

NIU Ling-ling1,XU Wei-li1,*,LIU Lei2,HE Sheng-hua1,MI Ya-qing1,LIU Qiao-hong1,LU Zhao-xin1

(1.Department of Food Science and Engineering,School of Chemical Engineering and technology,Harbin Institute of Technology,Harbin 150090,China;2.College of Basic Medical Sciences,Jiamusi University,Jiamusi 154007,China)

Vitamin E has two subfamilies,including tocopherols and tocotrienols and each subfamily has four monomers(α,β,γandδ). Compared to other vitamin E monomers,γ-tocotrienol(γ-T3)has a better anti-tumor,neuroprotection,lower cholesterol,inhibition to fat production and other anti-inflammatory effect. The biological activity and mechanism ofγ-T3 were reviewed,the existing problems ofγ-T3 biological activity was put forward and the research direction was forecast.

γ-tocotrienol;biological activity;mechanism

2015-05-19

牛玲玲(1991-),女,在讀碩士研究生,從事營養(yǎng)學(xué)領(lǐng)域研究,E-mail:mrgirlishere@163.com。

徐偉麗(1977-),女,博士,講師,研究方向:食物中生物活性成分的功能研究及食品安全,E-mail:weilixu698@163.com。

2013年黑龍江省博士后科研啟動(dòng)金資助(LBH-Q13086);國家自然科學(xué)基金(31501481,31371685)。

TS201.1

A

1002-0306(2016)05-0374-05

10.13386/j.issn1002-0306.2016.05.068

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