王思嘉，孫嘉瑩，劉美琦，任偉超，于欣欣，劉秀波，馬 偉, 3*

人參基因家族生物信息學(xué)分析

王思嘉1，孫嘉瑩1，劉美琦1，任偉超1，于欣欣1，劉秀波2*，馬 偉1, 3*

1.黑龍江中醫(yī)藥大學(xué)藥學(xué)院，黑龍江哈爾濱 150040 2.黑龍江中醫(yī)藥大學(xué)佳木斯學(xué)院，黑龍江佳木斯 154007 3.教育部北藥基礎(chǔ)與應(yīng)用研究重點(diǎn)實(shí)驗(yàn)室，黑龍江哈爾濱 150040

通過(guò)人參基因家族生物信息學(xué)分析，為人參功能基因開(kāi)發(fā)利用提供理論依據(jù)。運(yùn)用PlantTFDB數(shù)據(jù)庫(kù)預(yù)測(cè)人參基因組中基因；通過(guò)ExPASy網(wǎng)站得到人參基因家族信息和特征；使用MEME網(wǎng)站對(duì)基因家族保守基因序列進(jìn)行分析；運(yùn)用MEGA軟件建立基因家族系統(tǒng)進(jìn)化關(guān)系；利用TBtools軟件進(jìn)行表達(dá)分析。人參基因組中含有157個(gè)基因家族成員，其中152個(gè)定位在細(xì)胞核，其余5個(gè)分別定位在葉綠體和內(nèi)質(zhì)網(wǎng)，相對(duì)分子質(zhì)量在14 009.93～83 440.38，等電點(diǎn)在4.53～10.05，氨基酸數(shù)目在120～760 aa，除以外，均為親水蛋白。在干旱脅迫下的表達(dá)量為83.47；在根和鹽脅迫條件下表達(dá)量均最高，分別為119.82和117.86；在葉中表達(dá)量為48.54；花中表達(dá)量最高為79.55；成熟果實(shí)中表達(dá)量最高是，為119.32；在未成熟果實(shí)中的表達(dá)量最高，且為65.32。推測(cè)與同源基因功能相似，可以調(diào)控根的分生組織活性；在干旱脅迫條件下表達(dá)量最高，推測(cè)對(duì)干旱脅迫有重要調(diào)控作用；A亞族成員/（）、（）、/（）參與干旱脅迫和鹽脅迫調(diào)控，推測(cè)與A亞族成員具有同源關(guān)系的、、、在干旱脅迫和鹽脅迫條件下具有相似調(diào)控作用。

人參；；生物信息學(xué)；基因家族；鹽脅迫；干旱脅迫

作為成員眾多的轉(zhuǎn)錄因子家族，在植物領(lǐng)域研究較為透徹[1]。該基因家族對(duì)植物的生長(zhǎng)發(fā)育有重要調(diào)節(jié)作用，參與植物次生代謝和非生物脅迫的調(diào)控[2-3]。通過(guò)前人研究成果發(fā)現(xiàn)，在擬南芥、陸地棉、花生、小麥、番茄等植物中分別含有75、151、112、156、70個(gè)基因成員[4-8]。前期研究將基因家族分為10個(gè)亞族[9]，分為A、B、C、D、E、F、G、H、I和S。其中，A亞族的和參與應(yīng)激響應(yīng)脫落酸（ABA）信號(hào)[10]，可激活多個(gè)晚期胚胎發(fā)育[11]；C亞族中的、與共同表達(dá)可調(diào)控種子特異性[12]；在D亞族中，確定花器官數(shù)目，并參與莖部和花分生組織的表達(dá)[13]；E亞族中和通過(guò)其他基因家族成員相互作用參與植株發(fā)育[14]。除此之外，在芹菜中發(fā)現(xiàn)基因在干旱、低溫、鹽處理?xiàng)l件下的表達(dá)量不同，并隨時(shí)間不同發(fā)生變化[15]；蒺藜苜蓿中基因參與鹽脅迫，和均參與干旱脅迫[16]。

人參C.A.Meyer是五加科、人參屬多年生草本植物[17]，在明代《理虛元鑒》中提到其具有大補(bǔ)元?dú)?、?fù)脈固脫之效[18]，所以對(duì)人體機(jī)能恢復(fù)有一定作用。人參用于治療神經(jīng)衰弱和抑郁型精神病，也可用于治療冠心病[19]，且人參皂苷Rg1有保護(hù)糖尿病大鼠腎臟的作用[20]，人參皂苷Rg3與紫杉醇結(jié)合可抑制腫瘤細(xì)胞增長(zhǎng)[21]，如Luminal型乳腺癌[22]。同時(shí)，人參的一些活性成分在美容領(lǐng)域有一定研究[23]。近期研究表明影響人參皂苷的合成[24]，可見(jiàn)，人參在臨床上具有較高的藥用價(jià)值且應(yīng)用廣泛。

本研究利用了生物信息學(xué)技術(shù)，對(duì)人參基因家族成員進(jìn)行鑒定，分析基因家族信息、系統(tǒng)進(jìn)化、保守基序以及基因表達(dá)等，為深入研究人參轉(zhuǎn)錄因子在干旱、鹽、低溫等協(xié)迫條件下的調(diào)控作用提供理論基礎(chǔ)。

1 材料

人參基因組序列文件下載自人參基因組數(shù)據(jù)庫(kù)（ginseng genome database，http://ginsengdb.snu.ac.kr/data.php），擬南芥基因家族來(lái)自TAIR數(shù)據(jù)庫(kù)（http://www.arabidopsis.org/index.jsp）。

2 方法

2.1 人參bZIP基因家族的鑒定

使用生信軟件TBtools（v1.086）提取人參全部基因序列，將得到的蛋白序列文件提交plantTFDB數(shù)據(jù)庫(kù)（http://planttfdb.cbi.pku.edu.cn/）進(jìn)行預(yù)測(cè)，得到人參基因家族成員，用TBtools軟件去除重復(fù)轉(zhuǎn)錄本，并利用Expasy網(wǎng)站（https://web.expasy.org/ protparam/）獲取人參的等點(diǎn)量、相對(duì)分子質(zhì)量、蛋白質(zhì)疏水性、不穩(wěn)定系數(shù)、脂肪族系數(shù)等信息，利用WoLFPSORT （https://wolfpsort.hgc.jp/）對(duì)人參基因家族成員進(jìn)行亞細(xì)胞定位。

2.2 人參bZIP保守基序分析

使用MEME網(wǎng)站（http://meme-suite.org/tools/ meme），將人參bZIP家族成員蛋白序列的文件進(jìn)行保守基序預(yù)測(cè)，最大保守基序數(shù)目設(shè)為10，其余條件默認(rèn)，并使用TBtools軟件進(jìn)行可視化。

2.3 人參bZIP基因家族的系統(tǒng)進(jìn)化

把人參bZIP蛋白序列與擬南芥bZIP蛋白序列整合，將所得文件使用MUSCLE程序進(jìn)行多序列比對(duì)，利用Mega-X軟件采用鄰接法（neighbor-joining，NJ），Bootstrap值設(shè)置為1000次重復(fù)，模型選擇Poisson model，構(gòu)建系統(tǒng)發(fā)育樹(shù)。

2.4 人參bZIP基因表達(dá)分析

根據(jù)人參的等基因數(shù)據(jù)，在TBtools中繪制熱圖，進(jìn)一步對(duì)人參基因家族進(jìn)行表達(dá)分析。

3 結(jié)果與分析

3.1 人參bZIP基因家族信息和特征

根據(jù)人參基因家族的信息（表1）可知，氨基酸數(shù)目在120 aa（PgbZIP19）～760 aa（PgbZIP22），相對(duì)分子質(zhì)量的變化范圍為14 009.93（PgbZIP19）～83 440.38（PgbZIP22），等電點(diǎn)的范圍介于4.53～10.05（PgbZIP76～PgbZIP4），亞細(xì)胞定位顯示除PgbZIP5、PgbZIP11、PgbZIP79、PgbZIP116、PgbZIP130以外均定位在細(xì)胞核，PgbZIP5、PgbZIP116定位在葉綠體，PgbZIP11、PgbZIP79、PgbZIP130定位在內(nèi)質(zhì)網(wǎng)；蛋白質(zhì)疏水性（GRAVY）除PgbZIP157外均為負(fù)值；PgbZIP5、PgbZIP115、PgbZIP126的不穩(wěn)定系數(shù)小于40，為穩(wěn)定蛋白，其余為不穩(wěn)定蛋白。

表1 人參bZIP基因家族信息和特征

Table 1 Information and characteristics of ginseng bZIP gene family

基因名稱(chēng)基因序列不穩(wěn)定系數(shù)氨基酸數(shù)量相對(duì)分子質(zhì)量等電點(diǎn)蛋白質(zhì)疏水性亞細(xì)胞定位脂肪族系數(shù) PgbZIP1Pg_S0015.258.5142047 358.916.49?0.838細(xì)胞核67.62 PgbZIP2Pg_S0054.554.5340746 070.726.10?0.814細(xì)胞核71.43 PgbZIP3Pg_S0157.158.8332636 432.058.65?0.489細(xì)胞核86.01 PgbZIP4Pg_S0161.2961.5320421 865.7510.05?0.750細(xì)胞核61.23 PgbZIP5Pg_S0185.134.0616718 520.479.85?0.278葉綠體87.07 PgbZIP6Pg_S0227.3247.5136340 718.146.94?0.886細(xì)胞核57.36 PgbZIP7Pg_S0266.1766.6237040 533.445.57?0.791細(xì)胞核61.03 PgbZIP8Pg_S0270.157.8445549 459.836.20?0.753細(xì)胞核64.15 PgbZIP9Pg_S0292.4145.3833737 695.697.75?0.736細(xì)胞核74.07 PgbZIP10Pg_S0431.2552.7333036 872.618.34?0.814細(xì)胞核69.73 PgbZIP11Pg_S0431.545.9475682 233.396.04?0.632內(nèi)質(zhì)網(wǎng)63.43 PgbZIP12Pg_S0447.753.3338341 604.956.24?0.840細(xì)胞核58.36 PgbZIP13Pg_S0500.3441.0020123 151.389.52?0.040細(xì)胞核97.01 PgbZIP14Pg_S0602.2541.5736140 739.216.09?0.419細(xì)胞核83.88 PgbZIP15Pg_S0624.1961.2415517 986.097.91?1.008細(xì)胞核64.84 PgbZIP16Pg_S0659.1055.9946051 724.917.06?0.482細(xì)胞核75.57 PgbZIP17Pg_S0662.3860.8150455 852.426.88?0.586細(xì)胞核74.42 PgbZIP18Pg_S0662.4051.2933137 878.636.37?0.888細(xì)胞核64.20 PgbZIP19Pg_S0662.4355.7712014 009.939.64?0.828細(xì)胞核70.75 PgbZIP20Pg_S0684.2852.3331635 490.074.63?0.456細(xì)胞核88.83 PgbZIP21Pg_S0687.150.7016918 223.945.94?0.691細(xì)胞核64.67 PgbZIP22Pg_S0741.1645.6376383 440.386.28?0.487細(xì)胞核75.48 PgbZIP23Pg_S0741.3858.4643047 802.186.85?0.658細(xì)胞核69.70 PgbZIP24Pg_S0745.2167.2937642 193.557.82?0.891細(xì)胞核63.40 PgbZIP25Pg_S0762.1964.2937640 738.815.72?0.649細(xì)胞核65.48 PgbZIP26Pg_S0788.342.2213315 153.859.57?0.377細(xì)胞核87.29 PgbZIP27Pg_S0872.672.6418320 453.586.59?0.997細(xì)胞核51.86 PgbZIP28Pg_S0884.151.7628731 741.486.67?0.206細(xì)胞核79.16 PgbZIP29Pg_S0889.6139.6525527 585.928.95?0.634細(xì)胞核76.94 PgbZIP30Pg_S0905.134.6127529 893.275.86?0.683細(xì)胞核64.18 PgbZIP31Pg_S0938.951.1029533 135.474.72?0.467細(xì)胞核86.88 PgbZIP32Pg_S0967.2773.7025228 878.759.91?0.616細(xì)胞核73.45 PgbZIP33Pg_S0981.1549.6640142 806.986.04?0.818細(xì)胞核51.17 PgbZIP34Pg_S0992.2254.1859765 001.448.66?0.727細(xì)胞核61.68 PgbZIP35Pg_S1022.244.7219921 669.626.15?0.778細(xì)胞核65.68 PgbZIP36Pg_S1055.2752.2729033 092.188.72?0.890細(xì)胞核63.55 PgbZIP37Pg_S1084.852.5445449 502.749.53?0.649細(xì)胞核72.80 PgbZIP38Pg_S1097.3269.3914316 053.836.96?0.694細(xì)胞核55.24 PgbZIP39Pg_S1144.256.8543546 958.299.11?0.694細(xì)胞核65.49 PgbZIP40Pg_S1164.663.1117220 021.326.16?0.836細(xì)胞核77.15 PgbZIP41Pg_S1176.1459.0241243 666.805.86?0.942細(xì)胞核46.04 PgbZIP42Pg_S1222.150.0333738 200.436.03?0.433細(xì)胞核83.18 PgbZIP43Pg_S1241.249.2813515 270.995.91?0.696細(xì)胞核57.78 PgbZIP44Pg_S1242.2368.0833837 650.935.89?0.715細(xì)胞核67.87 PgbZIP45Pg_S1290.3562.9144148 732.357.15?0.655細(xì)胞核68.82 PgbZIP46Pg_S1338.2056.0458965 153.926.35?0.527細(xì)胞核76.53 PgbZIP47Pg_S1348.1064.9336339 935.995.82?0.796細(xì)胞核64.10 PgbZIP48Pg_S1349.160.0013215 826.019.46?0.883細(xì)胞核74.55 PgbZIP49Pg_S1389.3351.3040843 739.046.19?0.884細(xì)胞核49.80

續(xù)表1

基因名稱(chēng)基因序列不穩(wěn)定系數(shù)氨基酸數(shù)量相對(duì)分子質(zhì)量等電點(diǎn)蛋白質(zhì)疏水性亞細(xì)胞定位脂肪族系數(shù) PgbZIP50Pg_S1420.156.6046050 405.539.41?0.707細(xì)胞核71.17 PgbZIP51Pg_S1456.247.4926728 784.606.24?0.675細(xì)胞核64.31 PgbZIP52Pg_S1457.1359.9629433 033.726.27?0.906細(xì)胞核61.77 PgbZIP53Pg_S1511.260.7139744 477.497.71?0.577細(xì)胞核80.83 PgbZIP54Pg_S1514.651.1429532 949.184.66?0.474細(xì)胞核86.54 PgbZIP55Pg_S1544.549.3135039 660.196.68?0.405細(xì)胞核88.69 PgbZIP56Pg_S1552.2561.4134537 802.025.59?0.731細(xì)胞核67.39 PgbZIP57Pg_S1563.1855.9418821 978.335.88?0.973細(xì)胞核64.84 PgbZIP58Pg_S1565.159.0034338 577.779.00?0.678細(xì)胞核79.91 PgbZIP59Pg_S1583.1460.7416818 710.599.69?1.162細(xì)胞核52.92 PgbZIP60Pg_S1622.461.6929232 892.646.27?0.887細(xì)胞核62.53 PgbZIP61Pg_S1649.248.0935039 141.396.57?0.847細(xì)胞核60.60 PgbZIP62Pg_S1650.555.0020823 519.519.54?0.853細(xì)胞核69.47 PgbZIP63Pg_S1658.2046.0748052 700.005.62?0.471細(xì)胞核74.83 PgbZIP64Pg_S1668.1455.7217220 028.355.62?0.734細(xì)胞核79.42 PgbZIP65Pg_S1678.943.5416819 224.617.99?0.878細(xì)胞核58.86 PgbZIP66Pg_S1679.4154.5615517 984.116.52?0.954細(xì)胞核66.71 PgbZIP67Pg_S1693.2669.1521024 377.308.54?0.851細(xì)胞核67.24 PgbZIP68Pg_S1717.661.7142446 545.546.13?0.793細(xì)胞核64.25 PgbZIP69Pg_S1721.958.4451256 857.386.81?0.516細(xì)胞核73.01 PgbZIP70Pg_S1800.161.3735739 424.489.07?0.437細(xì)胞核84.31 PgbZIP71Pg_S1864.159.7149855 340.856.86?0.524細(xì)胞核73.45 PgbZIP72Pg_S1869.757.8350855 745.978.39?0.507細(xì)胞核76.22 PgbZIP73Pg_S1900.367.6815817 294.219.64?1.116細(xì)胞核64.24 PgbZIP74Pg_S1915.1751.3544148 531.729.72?0.723細(xì)胞核67.71 PgbZIP75Pg_S1933.569.9235540 525.187.32?0.885細(xì)胞核68.73 PgbZIP76Pg_S1993.2851.7021824 386.154.53?0.621細(xì)胞核88.01 PgbZIP77Pg_S2000.1168.2646950 995.106.43?1.158細(xì)胞核45.20 PgbZIP78Pg_S2014.560.4917520 042.415.72?0.786細(xì)胞核61.43 PgbZIP79Pg_S2057.2546.0975281 850.906.36?0.657內(nèi)質(zhì)網(wǎng)63.51 PgbZIP80Pg_S2057.353.6533036 880.537.10?0.808細(xì)胞核70.61 PgbZIP81Pg_S2086.353.4237041 530.805.96?0.751細(xì)胞核79.95 PgbZIP82Pg_S2109.144.6326728 731.586.71?0.633細(xì)胞核64.31 PgbZIP83Pg_S2176.163.2024827 929.969.20?1.026細(xì)胞核54.84 PgbZIP84Pg_S2176.266.7623726 141.146.35?0.705細(xì)胞核62.32 PgbZIP85Pg_S2207.2453.1616218 637.776.29?0.916細(xì)胞核61.48 PgbZIP86Pg_S2245.4137.0529732 562.615.74?0.513細(xì)胞核71.85 PgbZIP87Pg_S2306.247.7047151 983.506.40?0.490細(xì)胞核75.20 PgbZIP88Pg_S2466.160.0217719 957.388.38?0.802細(xì)胞核72.66 PgbZIP89Pg_S2496.2052.9849753 511.616.40?0.710細(xì)胞核63.08 PgbZIP90Pg_S2533.261.0315817 289.108.96?1.124細(xì)胞核59.94 PgbZIP91Pg_S2631.1035.4927830 366.925.39?0.564細(xì)胞核73.67 PgbZIP92Pg_S2754.461.2535639 287.319.07?0.423細(xì)胞核86.74 PgbZIP93Pg_S2757.562.8116117 636.529.68?1.115細(xì)胞核59.44 PgbZIP94Pg_S2874.474.0135540 591.358.48?0.872細(xì)胞核71.21 PgbZIP95Pg_S2936.2468.6837541 992.327.81?0.910細(xì)胞核62.53 PgbZIP96Pg_S2971.3358.3114016 833.149.89?1.040細(xì)胞核58.57 PgbZIP97Pg_S2990.265.3419822 967.385.64?0.968細(xì)胞核69.13 PgbZIP98Pg_S3098.2156.6242344 644.976.24?0.891細(xì)胞核45.56 PgbZIP99Pg_S3109.1757.2518721 425.135.72?0.710細(xì)胞核61.66 PgbZIP100Pg_S3138.2158.0025928 290.966.38?0.568細(xì)胞核73.40 PgbZIP101Pg_S3210.142.6430033 871.757.72?0.858細(xì)胞核64.07 PgbZIP102Pg_S3210.252.1033137 937.737.02?0.883細(xì)胞核64.17 PgbZIP103Pg_S3210.559.9457063 743.438.27?0.639細(xì)胞核73.18 PgbZIP104Pg_S3356.1451.3941043 461.267.82?0.768細(xì)胞核60.05 PgbZIP105Pg_S3402.152.1236140 840.446.62?0.445細(xì)胞核81.16 PgbZIP106Pg_S3475.156.4815417 766.859.00?0.882細(xì)胞核67.79 PgbZIP107Pg_S3530.959.4219822 184.806.12?0.586細(xì)胞核81.87 PgbZIP108Pg_S3584.373.9914316 078.926.96?0.677細(xì)胞核57.97

續(xù)表1

3.2 人參bZIP家族保守基序分析

使用MEME網(wǎng)站查找人參bZIP蛋白保守基序，可知PgbZIP的保守基序個(gè)數(shù)為1～6個(gè)。Motif9只在A(yíng)亞族中存在一小部分；D亞族中普遍含有6個(gè)保守基序；Motif5只存在于I、E家族中；PgbZIP中均含有Motif1，且少數(shù)PgbZIP成員中只含有Motif1，見(jiàn)圖1。

3.3 人參bZIP基因家族的系統(tǒng)進(jìn)化關(guān)系

將人參的157個(gè)bZIP蛋白和擬南芥中74個(gè)蛋白進(jìn)行系統(tǒng)進(jìn)化樹(shù)的構(gòu)建（圖2），bZIP家族分為10個(gè)亞家族，分別為A、B、C、D、E、F、G、H、I和S，其中A亞族中包括33個(gè)人參bZIP成員；B亞族中只含有2個(gè)成員PgbZIP53、PgbZIP144；C亞族中包括12個(gè)PgbZIP成員；D亞族中含有22個(gè)PgbZIP成員；E亞族中含有15個(gè)PgbZIP成員；F亞族中含有4個(gè)PgbZIP成員：PgbZIP151、PgbZIP91、PgbZIP86、PgbZIP30；G亞族中含有8個(gè)PgbZIP成員；H亞族中含有10個(gè)PgbZIP成員；I亞族中含有16個(gè)成員；S亞族含有35個(gè)成員，是含有人參bZIP成員數(shù)最多的亞族。

圖1 人參bZIP蛋白基序以及Motif分析

圖2 人參和擬南芥bZIP轉(zhuǎn)錄因子的系統(tǒng)發(fā)育樹(shù)

3.4 人參bZIP基因在植物不同器官中的表達(dá)

通過(guò)TBtools軟件制作熱圖，對(duì)基因家族的表達(dá)進(jìn)行分析。如圖3，表明、在成熟果實(shí)中表達(dá)量較高，表達(dá)最明顯，在未成熟果實(shí)中表達(dá)量最高；在花中表達(dá)最明顯；、在根中表達(dá)量較高；在根和鹽脅迫條件下中的表達(dá)量最高，在鹽脅迫下表達(dá)量較高；在干旱脅迫下表達(dá)量最高。

4 討論

bZIP是一類(lèi)參與植物生長(zhǎng)、生物與非生物脅迫的轉(zhuǎn)錄因子，通過(guò)圖2和圖3分析，人參中在根中的表達(dá)量最高，在莖和花中的表達(dá)量逐漸降低，與其同源的擬南芥基因具有抑制分生組織活性與根生長(zhǎng)的功能[25]，推測(cè)其在也具有負(fù)調(diào)控作用。（）、（）、（）、（）、（）、（）為擬南芥中TGA中的成員，相互作用參與植物疾病調(diào)控[26-27]，推測(cè)其同源基因、、、也有相似的功能。

非生物脅迫包括干旱脅迫、鹽脅迫、低溫脅迫等[28]，Yoshida等[29]發(fā)現(xiàn)bZIP轉(zhuǎn)錄因子對(duì)抗逆性有重要作用。在干旱脅迫條件下表達(dá)量最高，屬于S亞族，該亞族對(duì)非生物脅迫有較強(qiáng)響應(yīng)，其中AtbZIP44在干旱脅迫中發(fā)揮重要作用[30]，因與為同源基因，推測(cè)在人參中起到相似作用。

ABA在植物的生長(zhǎng)發(fā)育中對(duì)環(huán)境脅迫發(fā)揮重要作用[31]，其中擬南芥成員/（）、（）、/（）主要參與ABA、鹽脅迫、干旱脅迫、熱脅迫等[32]，成員、、與其為同源基因，且在鹽脅迫條件下表達(dá)較高，推測(cè)、、可能參與鹽脅迫調(diào)控。對(duì)鹽脅迫有較強(qiáng)反應(yīng)[31]，、在鹽脅迫條件下的表達(dá)量較高，推測(cè)其在人參鹽脅迫條件下發(fā)揮重要作用。Fujita等[32]發(fā)現(xiàn)擬南芥中的ABRE結(jié)合因子家族成員依賴(lài)ABA信號(hào)，增強(qiáng)植物營(yíng)養(yǎng)組織的抗旱性。煙草研究在A(yíng)BA的處理下使A亞族中的成員表達(dá)量上升[33]，結(jié)合圖3發(fā)現(xiàn)成員、、、、、表達(dá)量在葉、花、干旱脅迫下或鹽脅迫有明顯增加，推測(cè)在A(yíng)BA處理下也有相似功能，參與人參的生長(zhǎng)發(fā)育。

圖3 人參bZIP基因在植物不同器官和非脅迫條件下的表達(dá)分析

通過(guò)人參基因家族的生物信息學(xué)分析，對(duì)可能參與人參非生物脅迫的基因進(jìn)行功能預(yù)測(cè)，為深入研究基因提供分子基礎(chǔ)，為進(jìn)一步探索人參次生代謝調(diào)控以及藥用活性成分臨床應(yīng)用提供理論依據(jù)。

利益沖突 所有作者均聲明不存在利益沖突

[1] 劉莉.DREBs、轉(zhuǎn)錄因子與植物抗旱性研究進(jìn)展 [J].浙江農(nóng)業(yè)科學(xué), 2013, 54(1): 98-102.

[2] 沈迪, 陳龍正, 陶建平, 等.芹菜bZIP轉(zhuǎn)錄因子基因AgbZIP16的逆境響應(yīng)分析 [J].植物生理學(xué)報(bào), 2019, 55(12): 1817-1826.

[3] 曹紅利.茶樹(shù)家族基因的非生物脅迫響應(yīng)及C亞家族CsbZIP6和CsbZIP4的功能初步分析 [D].北京: 中國(guó)農(nóng)業(yè)科學(xué)院, 2016.

[4] Jakoby M, Weisshaar B, Dr?ge-Laser W,.bZIP transcription factors in[J]., 2002, 7(3): 106-111.

[5] 邢宇鵬.棉花基因家族全基因組鑒定及分析 [D].泰安: 山東農(nóng)業(yè)大學(xué), 2020.

[6] 高斌, 陳娟娟, 崔順立, 等.花生基因家族全基因組鑒定及抗旱表達(dá)分析 [J].植物遺傳資源學(xué)報(bào), 2020, 21(1): 174-191.

[7] 李雪垠.小麥轉(zhuǎn)錄因子家族中花藥發(fā)育及抗逆相關(guān)基因的功能研究 [D].楊凌: 西北農(nóng)林科技大學(xué), 2016.

[8] 朱蕓曄, 薛冰, 王安全, 等.番茄轉(zhuǎn)錄因子家族的生物信息學(xué)分析 [J].應(yīng)用與環(huán)境生物學(xué)報(bào), 2014, 20(5): 767-774.

[9] Dr?ge-Laser W, Snoek B L, Snel B,.Thetranscription factor family-an update [J]., 2018, 45(Pt A): 36-49.

[10] Kang J Y, Choi H I, Im M Y,.basic leucine zipper proteins that mediate stress-responsive abscisic acid signaling [J]., 2002, 14(2): 343-357.

[11] Bensmihen S, Rippa S, Lambert G,.The homologousandtranscription factors function antagonistically to fine-tune gene expression during late embryogenesis [J]., 2002, 14(6): 1391-1403.

[12] Lara P, O?ate-Sánchez L, Abraham Z,.Synergistic activation of seed storage protein gene expression inby ABI3and two bZIPs related to OPAQUE2 [J]., 2003, 278(23): 21003-21011.

[13] Fletcher J C.The ULTRAPETALA gene controls shoot and floral meristem size in[J]., 2001, 128(8): 1323-1333.

[14] Shen H S, Cao K M, Wang X P.A conserved proline residue in the leucine zipper region of AtbZIP34 and AtbZIP61 ininterferes with the formation of homodimer [J]., 2007, 362(2): 425-430.

[15] 沈迪, 陳龍正, 陶建平, 等.芹菜轉(zhuǎn)錄因子基因的逆境響應(yīng)分析 [J].植物生理學(xué)報(bào), 2019, 55(12): 1817-1826.

[16] 齊曉, 張正社, 閔學(xué)陽(yáng), 等.紫花苜蓿基因家族的鑒定、進(jìn)化及表達(dá)分析 [J].草業(yè)科學(xué), 2017, 34(8): 1635-1648.

[17] 楊秀偉, 富力.人參中三萜類(lèi)化學(xué)成分的生物學(xué)活性和藥理學(xué)作用 [J].中國(guó)現(xiàn)代中藥, 2016, 18(1): 36-55.

[18] 龍玉婷, 孫志會(huì), 王志文, 等.人參大補(bǔ)元?dú)獾墓π?nèi)涵淺析 [J].中國(guó)中醫(yī)急癥, 2019, 28(9): 1679-1682.

[19] 柳良燕.人參西洋參功效比較及用法研究 [J].實(shí)用醫(yī)技雜志, 2008, 15(34): 41-43.

[20] 馬小芬, 謝席勝, 左川, 等.人參皂甙Rg1對(duì)糖尿病腎病大鼠腎臟保護(hù)作用的機(jī)制研究 [J].生物醫(yī)學(xué)工程學(xué)雜志, 2010, 27(2): 342-347.

[21] 張奉海, 張瑞榮, 陳淑娟, 等.人參皂苷Rg3聯(lián)合紫杉醇抑制人肝癌細(xì)胞HepG2增殖和對(duì)裸鼠移植瘤模型的作用機(jī)制研究 [J].中醫(yī)藥學(xué)報(bào), 2020, 48(10): 16-20.

[22] 鄭穎娟, 王輝, 劉曉靜, 等.人參皂苷對(duì)Luminal型乳腺癌中Fas、FasL表達(dá)水平的影響 [J].遼寧中醫(yī)雜志, 2020, 47(10): 185-189.

[23] 熊晨陽(yáng), 許明良, 易帆, 等.人參不同部位主要活性成分及其在美容護(hù)膚方面的研究進(jìn)展 [J].日用化學(xué)工業(yè), 2019, 49(3): 193-198.

[24] 李宏杰.吉林人參轉(zhuǎn)錄因子基因家族系統(tǒng)分析及PgbZIP48-3基因功能的初步驗(yàn)證 [D].長(zhǎng)春: 吉林農(nóng)業(yè)大學(xué), 2020.

[25] Weiste C, Pedrotti L, Selvanayagam J,.Thetranscription factor links low-energy signalling to auxin-mediated control of primary root growth [J]., 2017, 13(2): e1006607.

[26] Kesarwani M, Yoo J, Dong X N.Genetic interactions of TGA transcription factors in the regulation of pathogenesis-related genes and disease resistance in[J]., 2007, 144(1): 336-346.

[27] 田義, 張彩霞, 康國(guó)棟, 等.植物TGA轉(zhuǎn)錄因子研究進(jìn)展 [J].中國(guó)農(nóng)業(yè)科學(xué), 2016, 49(4): 632-642.

[28] 王冰, 程憲國(guó).干旱、高鹽及低溫脅迫下植物生理及轉(zhuǎn)錄因子的應(yīng)答調(diào)控 [J].植物營(yíng)養(yǎng)與肥料學(xué)報(bào), 2017, 23(6): 1565-1574.

[29] Yoshida T, Fujita Y, Sayama H,.AREB1, AREB2, and ABF3are master transcription factors that cooperatively regulate ABRE-dependent ABA signaling involved in drought stress tolerance and require ABA for full activation [J]., 2010, 61(4): 672-685.

[30] Weltmeier F, Rahmani F, Ehlert A,.Expression patterns within theC/S1transcription factor network: Availability of heterodimerization partners controls gene expression during stress response and development [J]., 2009, 69(1/2): 107-119.

[31] Sun X L, Li Y, Cai H,.Thetranscription factor is a positive regulator of plant tolerance to salt, osmotic and drought stresses [J]., 2012, 125(3): 429-438.

[32] Fujita Y, Fujita M, Satoh R,.AREB1 is a transcription activator of novel ABRE-dependent ABA signaling that enhances drought stress tolerance in[J]., 2005, 17(12): 3470-3488.

[33] Choi H I, Hong J H, Ha J O,.ABFs, a Family of ABA-responsive element binding factors [J]., 2000, 275(3): 1723-1730.

Bioinformatics analysis ofgene family of

WANG Si-jia1, SUN Jia-ying1, LIU Mei-qi1, REN Wei-chao1, YU Xin-xin1, LIU Xiu-bo2, MA Wei1, 3

1.College of Pharmaceutical Sciences, Heilongjiang University of Chinese Medicine, Harbin 150040, China 2.College of Jiamusi, Heilongjiang University of Chinese Medicine, Jiamusi 154007, China 3.Key Laboratory of Basic and Application Research of Beiyao (Heilongjiang University of Chinese Medicine), Ministry of Education, Harbin 150040, China

To analyzegene family bioinformatics of Renshen (), so as to provide theoretical basis for the development and utilization of ginseng functional genes.Using PlantTFDB database to predict ginsenggene; The genetic family information and characteristics of ginsengwere obtained through ExPASy website; The conserved gene sequences ofgene family were analyzed by MEME website; The phylogenetic relationship ofgene family were established by using MEGA software;expression analysis were conducted by TBtools software.The ginseng genome contained 157 bZIP gene family members, of which 152 were located in the nucleus, and the remaining five were located in the chloroplast and endoplasmic reticulum, respectively.The relative molecular mass was between 14 009.93 and 83 440.38, and the isoelectric point was in the range of 4.53—10.05, the number of amino acids was between 120 aa—760 aa, except for, all were hydrophilic proteins.The expression level ofunder drought stress was 83.47; the expression level ofwas the highest under root (119.82) and salt stress (117.86) conditions, respectively; the expression level ofin leaves was 48.54; the highest expression level ofin flowers was 79.55; The highest expression level ofin mature fruits was 119.32; the highest expression level ofin immature fruits was 65.32.It is speculated thathas similar functions to the homologous gene, and can regulate root meristem activity;has the highest expression under drought stress conditions, and it is speculated thathas an important regulatory effect on drought stress; A subfamily members,,/is involved in the regulation of drought stress and salt stress.It is speculated that,,and, which have a homologous relationship with members of subfamily A, have similar regulatory effects under drought stress and salt stress.

C.A.Meyer;; bioinformatics; gene family; salt stress; drought stress

R282.12

A

0253 - 2670(2022)09 - 2786 - 09

10.7501/j.issn.0253-2670.2022.09.022

2021-11-09

黑龍江省“頭雁”團(tuán)隊(duì)項(xiàng)目（黑龍江省頭雁行動(dòng)領(lǐng)導(dǎo)小組文件 [2019] 5號(hào)）；黑龍江中醫(yī)藥大學(xué)科研基金項(xiàng)目（中藥健康相關(guān)產(chǎn)品研發(fā)及產(chǎn)業(yè)化專(zhuān)項(xiàng)）（2019BJP06）

王思嘉，女，碩士研究生，研究方向?yàn)樗幱弥参锷锕こ萄芯?。Tel: (0451)87266988 E-mail: 2077737519@qq.com

通信作者：馬 偉，研究員，博士生導(dǎo)師，主要從事藥用植物生物工程研究。Tel: (0451)87266988 E-mail: 88788891@qq.com

劉秀波，教授，碩士生導(dǎo)師，主要從事中藥資源與中藥化學(xué)。Tel: 13796353268 E-mail: 358270831@qq.com

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