生物技术通报 ›› 2022, Vol. 38 ›› Issue (9): 167-179.doi: 10.13560/j.cnki.biotech.bull.1985.2021-1459
袁星(), 郭彩华, 刘金明, 亢超, 全绍文, 牛建新()
收稿日期:
2021-11-12
出版日期:
2022-09-26
发布日期:
2022-10-11
作者简介:
袁星,女,硕士研究生,研究方向:园艺学;E-mail: 基金资助:
YUAN Xing(), GUO Cai-hua, LIU Jin-ming, KANG Chao, QUAN Shao-wen, NIU Jian-xin()
Received:
2021-11-12
Published:
2022-09-26
Online:
2022-10-11
摘要:
基于核桃基因组鉴定核桃COL(CONSTANS-Like)家族成员,并对其进行分析,以揭示核桃COL家族的分子进化和潜在功能。以‘新新2号'(Juglans regia cv. Xinxin No. 2)为材料,对COL基因家族全基因组进行鉴定及生物信息分析,并分析其在不同组织及不同时间雌花芽的表达。结果表明,核桃基因组中共存在55个COL基因,不均匀地分布在16个染色体和1个Scaffold上,依次命名为JrCOL1-JrCOL55,氨基酸为117-789 aa,分子量为12.99-86.08 kD,等电点为4.00-8.94,进化树分析将其分为3个亚族,且同一分支基因的保守基序和基因结构位置相似。启动子区的顺式作用元件分析发现JrCOL基因含有大量与光信号、植物激素、胁迫等相关的顺式作用元件。‘新新2号'中JrCOL家族基因具有组织特异性表达,其中JrCO1、JrCO7a-c、JrCO8、JrCOL23、JrCO31a-c、JrCO33、JrCOL35和JrCOL50a-f在叶片中表达量最高。对‘新新2号'雌花芽不同时间JrCOL基因的RT-qPCR分析表明,JrCOL1、JrCOL35、JrCOL50a-f和JrCOL52在‘新新2号'雌花芽中,具有一致的表达趋势,其中JrCOL50a-f在‘新新2号'雌花芽分化过程中起到重要作用。在核桃基因组中鉴定出55个COL家族基因,分为3个亚家族,不均匀地分布在16个染色体和1个Scaffold上,结构进化保守,组织表达模式多样。推测JrCOL家族基因在核桃叶片发育过程中发挥重要功能。
袁星, 郭彩华, 刘金明, 亢超, 全绍文, 牛建新. 核桃CONSTANS-Like基因家族全基因组鉴定及表达分析[J]. 生物技术通报, 2022, 38(9): 167-179.
YUAN Xing, GUO Cai-hua, LIU Jin-ming, KANG Chao, QUAN Shao-wen, NIU Jian-xin. Genome-wide Identification of CONSTANS-Like Family Genes and Expression Analysis in Wanlut[J]. Biotechnology Bulletin, 2022, 38(9): 167-179.
基因 Gene | 上游引物 Forward primer(5'-3') | 下游引物 Reverse primer(5'-3') |
---|---|---|
JrCOL1 | TTCGAGAAGACGATCCGATACG | CGATCAACTTCCGATCCCATCT |
JrCOL7 | ATTCCCCTTCTGTGGGTGAATC | TGGCTCATGTTGTTTTCTGCAG |
JrCOL8 | GTTATGATGCCATCGACAGCAC | TGCCATTCCATTAACGTGAGGA |
JrCOL23 | CGTGACGGATGTATCGGATTCA | TTTTCTCTGTACCTCAGCACCC |
JrCOL31 | CTGCAGAAAACAACATGAGCCA | AATGTGAGGCCCAGAATGAACT |
JrCOL33 | GGTTCACTGGATGTCCTTCTGT | TGAGAAGCATCATCCCAAACCA |
JrCOL35 | ACCCCAATTTCAGTTCCAAGGT | TATCCGTTCCCGCACTGTAAAA |
JrCOL42 | GTGGCTCTAAAGATGTCCCACA | TATTCTGTGCCAGTTGCTCCAT |
JrCOL50 | CTCGTTGCGATCATCTTGTGTC | GATGACACTGAAGAGCCTCCAA |
JrCOL52 | AGAATTGGGAAATGGCAGGGAT | CCATCGTAAGCAAAGCCGTAAG |
18S | GGTCAATCTTCTCGTTCCCTT | TCGCATTTCGCTACGTTCTT |
表1 引物序列
Table 1 Primer sequences
基因 Gene | 上游引物 Forward primer(5'-3') | 下游引物 Reverse primer(5'-3') |
---|---|---|
JrCOL1 | TTCGAGAAGACGATCCGATACG | CGATCAACTTCCGATCCCATCT |
JrCOL7 | ATTCCCCTTCTGTGGGTGAATC | TGGCTCATGTTGTTTTCTGCAG |
JrCOL8 | GTTATGATGCCATCGACAGCAC | TGCCATTCCATTAACGTGAGGA |
JrCOL23 | CGTGACGGATGTATCGGATTCA | TTTTCTCTGTACCTCAGCACCC |
JrCOL31 | CTGCAGAAAACAACATGAGCCA | AATGTGAGGCCCAGAATGAACT |
JrCOL33 | GGTTCACTGGATGTCCTTCTGT | TGAGAAGCATCATCCCAAACCA |
JrCOL35 | ACCCCAATTTCAGTTCCAAGGT | TATCCGTTCCCGCACTGTAAAA |
JrCOL42 | GTGGCTCTAAAGATGTCCCACA | TATTCTGTGCCAGTTGCTCCAT |
JrCOL50 | CTCGTTGCGATCATCTTGTGTC | GATGACACTGAAGAGCCTCCAA |
JrCOL52 | AGAATTGGGAAATGGCAGGGAT | CCATCGTAAGCAAAGCCGTAAG |
18S | GGTCAATCTTCTCGTTCCCTT | TCGCATTTCGCTACGTTCTT |
基因名称 Gene name | 基因ID Gene ID | 染色体位置 Chromosome position | 相关蛋白 Related protein | 蛋白重命名 Protein rename | 氨基酸长度 aa length | 蛋白分子量 Molecular weight/kD | 理论等电点pI | 亚细胞定位Subc- ellular localization |
---|---|---|---|---|---|---|---|---|
JrCOL1 | LOC109001350 | Chr.1:1256032..1258201 | XP_018834143.1 | JrCOL1 | 373 | 41.31 | 5.17 | 叶绿体 |
JrCOL2 | LOC109001381 | Chr.1:1488506..1490928 | XP_035544637.1 | JrCOL2 | 211 | 23.40 | 5.73 | 细胞质 |
JrCOL3 | LOC109009665 | Chr.1:3978995..3983057 | XP_018845790.2 | JrCOL3 | 350 | 39.64 | 8.72 | 细胞核 |
JrCOL4a-d | LOC109010156 | Chr.1:18366251..18370500 | XP_035543378.1 | JrCOL4a | 428 | 47.24 | 5.27 | 细胞核 |
XP_018846422.1 | JrCOL4b | 428 | 47.24 | 5.27 | 细胞核 | |||
XP_018846423.1 | JrCOL4c | 428 | 47.24 | 5.27 | 细胞核 | |||
XP_018846424.1 | JrCOL4d | 428 | 47.24 | 5.27 | 细胞核 | |||
JrCOL5 | LOC109015440 | Chr.1:18372462..18372860 | XP_018853453.2 | JrCOL5 | 132 | 14.50 | 6.85 | 细胞核 |
JrCOL6a-b | LOC108991987 | Chr.1:19317524..19318826 | XP_018821963.2 | JrCOL6a | 237 | 25.91 | 4.42 | 细胞核 |
XP_018821964.2 | JrCOL6b | 236 | 25.78 | 4.42 | 细胞核 | |||
JrCOL7a-c | LOC109000761 | Chr.1:44957519..44963229 | XP_018833298.1 | JrCOL7a | 416 | 45.11 | 5.19 | 细胞核 |
XP_018833299.1 | JrCOL7b | 416 | 45.11 | 5.19 | 细胞核 | |||
XP_018833303.1 | JrCOL7c | 416 | 45.11 | 5.19 | 细胞核 | |||
JrCOL8 | LOC109013302 | Chr.2:7141021..7147381 | XP_018850883.1 | JrCOL8 | 539 | 60.48 | 6.22 | 细胞核 |
JrCOL9 | LOC108984132 | Chr.2:11707792..11709460 | XP_018811512.1 | JrCOL9 | 277 | 31.23 | 4.94 | 细胞核 |
JrCO10 | LOC109014521 | Chr.2:30680768..30682741 | XP_018852564.1 | JrCOL10 | 333 | 37.49 | 4.97 | 细胞核 |
JrCOL11a-b | LOC109014085 | Chr.2:35047335..35048788 | XP_018851954.1 | JrCOL11a | 258 | 27.76 | 4.28 | 细胞核 |
XP_018851962.1 | JrCOL11b | 255 | 27.37 | 4.25 | 细胞核 | |||
JrCOL12a-b | LOC108999038 | Chr.3:18899526..18901860 | XP_018831379.1 | JrCOL12a | 150 | 16.21 | 4.00 | 细胞核,叶绿体 |
XP_035544064.1 | JrCOL12b | 150 | 16.21 | 4.00 | 细胞核,叶绿体 | |||
JrCOL13 | LOC109017893 | Chr.4:14653613..14655876 | XP_035544914.1 | JrCOL13 | 170 | 18.24 | 4.11 | 细胞核 |
JrCOL14 | LOC108989855 | Chr.5:6297905..6299253 | XP_018819161.1 | JrCOL14 | 220 | 26.37 | 4.94 | 细胞核 |
JrCOL15a-b | LOC109022307 | Chr.5:9188186..9197431 | XP_018860727.1 | JrCOL15a | 789 | 86.08 | 6.30 | 叶绿体 |
XP_018860748.1 | JrCOL15b | 778 | 84.89 | 6.35 | 叶绿体 | |||
JrCOL16 | LOC108994549 | Chr.6:15543376..15545802 | XP_018825341.2 | JrCOL16 | 444 | 48.81 | 5.88 | 细胞核 |
JrCOL17 | LOC108998057 | Chr.6:29988891..29991120 | XP_018830038.2 | JrCOL17 | 407 | 46.32 | 5.46 | 细胞核 |
JrCOL18 | LOC108992642 | Chr.7:12544651..12554366 | XP_018822785.2 | JrCOL18 | 685 | 75.72 | 6.71 | 细胞核 |
JrCOL19 | LOC108983659 | Chr.7:14608063..14609540 | XP_018810913.1 | JrCOL19 | 266 | 28.49 | 8.73 | 叶绿体 |
JrCOL20 | LOC109005469 | Chr.7:32506953..32517320 | XP_018839979.2 | JrCOL20 | 238 | 26.36 | 5.08 | 细胞核 |
JrCOL21 | LOC109019524 | Chr.7:38617705..38622031 | XP_018857358.1 | JrCOL21 | 432 | 46.81 | 4.35 | 叶绿体 |
JrCOL22a-b | LOC109014549 | Chr.7:49644025..49658252 | XP_035548321.1 | JrCOL22a | 780 | 85.75 | 6.42 | 叶绿体 |
XP_035548322.1 | JrCOL22b | 778 | 85.50 | 6.42 | 叶绿体 | |||
JrCOL23 | LOC109000996 | Chr.8:600112..601757 | XP_018833641.1 | JrCOL23 | 345 | 37.98 | 6.16 | 细胞质 |
JrCOL24 | LOC108984911 | Chr.8:6608517..6610109 | XP_018812557.1 | JrCOL24 | 301 | 33.04 | 6.25 | 细胞核 |
JrCOL25a-b | LOC108980667 | Chr.8:24595513..24597399 | XP_035549568.1 | JrCOL25a | 158 | 17.12 | 5.62 | 细胞质 |
XP_035549569.1 | JrCOL25b | 117 | 12.99 | 6.49 | 细胞质 | |||
JrCOL26 | LOC108997344 | Chr.9:19840907..19842615 | XP_018829103.1 | JrCOL26 | 331 | 37.53 | 5.83 | 细胞核 |
JrCOL27 | LOC108985349 | Chr.9:21400481..21403221 | XP_018813169.1 | JrCOL27 | 295 | 31.92 | 5.55 | 叶绿体 |
JrCOL28 | LOC108986213 | Chr.10:2554388..2557653 | XP_018814300.1 | JrCOL28 | 299 | 34.08 | 6.43 | 细胞核 |
JrCOL29a-c | LOC108984383 | Chr.10:13263191..13268808 | XP_018811858.2 | JrCOL29a | 436 | 47.97 | 6.55 | 细胞质 |
XP_035550266.1 | JrCOL29b | 436 | 47.97 | 6.55 | 细胞质 | |||
XP_035550267.1 | JrCOL29c | 408 | 44.52 | 5.70 | 细胞核 | |||
JrCOL30a-b | LOC109017472 | Chr.10:13911798..13913088 | XP_018855279.1 | JrCOL30a | 260 | 28.60 | 4.37 | 叶绿体 |
XP_018855280.1 | JrCOL30b | 259 | 28.48 | 4.37 | 叶绿体 | |||
JrCOL31a-c | LOC109000741 | Chr.10:37518173..37524266 | XP_018833263.1 | JrCOL31a | 409 | 44.45 | 5.96 | 细胞核 |
XP_018833265.1 | JrCOL31b | 409 | 44.45 | 5.96 | 细胞核 | |||
XP_018833266.1 | JrCOL31c | 409 | 44.45 | 5.96 | 细胞核 | |||
JrCOL32a-d | LOC108992026 | Chr.11:8894066..8897365 | XP_018822007.1 | JrCOL32a | 194 | 21.42 | 6.43 | 叶绿体 |
XP_018822008.1 | JrCOL32b | 192 | 21.17 | 6.23 | 叶绿体 | |||
XP_018822009.1 | JrCOL32c | 186 | 20.48 | 6.43 | 叶绿体 | |||
XP_018822010.1 | JrCOL32d | 184 | 20.22 | 6.23 | 叶绿体 | |||
JrCOL33 | LOC108992144 | Chr.11:23645881..23650290 | XP_018822151.1 | JrCOL33 | 364 | 40.75 | 5.51 | 细胞核 |
JrCOL34 | LOC108991312 | Chr.11:29101394..29103034 | XP_018821038.1 | JrCOL34 | 300 | 33.48 | 5.99 | 细胞核 |
JrCOL35 | LOC108998881 | Chr.11:32536877..32538774 | XP_018831163.1 | JrCOL35 | 373 | 40.72 | 5.86 | 叶绿体 |
JrCOL36 | LOC109003090 | Chr.11:33364576..33368847 | XP_018836611.1 | JrCOL36 | 409 | 45.46 | 8.94 | 细胞核 |
JrCOL37 | LOC109005190 | Chr.11:36684126..36685787 | XP_018839540.1 | JrCOL37 | 341 | 37.58 | 6.87 | 叶绿体 |
JrCOL38a-b | LOC108982143 | Chr.12:8537584..8542475 | XP_018808983.1 | JrCOL38a | 671 | 74.38 | 5.97 | 细胞核 |
XP_035539295.1 | JrCOL38b | 652 | 72.23 | 6.08 | 细胞核 | |||
JrCOL39 | LOC108992474 | Chr.12:13436453..13439714 | XP_018822597.1 | JrCOL39 | 237 | 26.10 | 4.91 | 细胞核 |
JrCOL40a-b | LOC108981163 | Chr.12:23463905..23468842 | XP_018807797.1 | JrCOL40a | 423 | 46.11 | 4.45 | 细胞核 |
XP_018807806.1 | JrCOL40b | 418 | 45.53 | 4.53 | 细胞核 | |||
JrCOL41a-b | LOC108986927 | Chr.13:5686536..5688137 | XP_018815283.1 | JrCOL41a | 293 | 32.61 | 5.70 | 细胞核 |
XP_018815285.1 | JrCOL41b | 206 | 22.72 | 5.57 | 细胞核 | |||
JrCOL42 | LOC109008345 | Chr.13:8537826..8540900 | XP_018843947.2 | JrCOL42 | 475 | 51.86 | 5.92 | 细胞核 |
JrCOL43a-b | LOC108992288 | Chr.13:31296667..31301250 | XP_018822350.1 | JrCOL43a | 691 | 77.57 | 6.80 | 细胞核 |
XP_018822351.1 | JrCOL43b | 689 | 77.43 | 6.80 | 细胞核 | |||
JrCOL44a-d | LOC109007733 | Chr.13:37236625..37242278 | XP_018843087.1 | JrCOL44a | 518 | 57.78 | 5.61 | 细胞核 |
XP_018843095.1 | JrCOL44b | 518 | 57.78 | 5.61 | 细胞核 | |||
XP_018843104.1 | JrCOL44c | 518 | 57.78 | 5.61 | 细胞核 | |||
XP_018843113.1 | JrCOL44d | 518 | 57.78 | 5.61 | 细胞核 | |||
JrCOL45 | LOC109013207 | Chr.13:38161618..38165624 | XP_018850755.1 | JrCOL45 | 287 | 32.38 | 5.25 | 细胞核 |
JrCOL46 | LOC108996999 | Chr.14:6471961..6473893 | XP_018828617.1 | JrCOL46 | 218 | 25.99 | 5.27 | 细胞核 |
JrCOL47 | LOC108993945 | Chr.14:10350983..10353374 | XP_018824556.1 | JrCOL47 | 251 | 28.23 | 8.27 | 细胞核 |
JrCOL48 | LOC109013626 | Chr.15:14881124..14883084 | XP_018851315.1 | JrCOL48 | 464 | 52.39 | 5.09 | 细胞核,细胞质 |
JrCOL49 | LOC109004589 | Chr.16:1780812..1784882 | XP_018838732.1 | JrCOL49 | 279 | 31.45 | 5.27 | 细胞核 |
JrCOL50a-f | LOC109012501 | Chr.16:2630759..2636833 | XP_035542414.1 | JrCOL50a | 518 | 57.90 | 5.72 | 细胞核 |
XP_035542415.1 | JrCOL50b | 492 | 54.89 | 5.87 | 细胞核 | |||
XP_035542416.1 | JrCOL50c | 489 | 54.51 | 5.83 | 细胞核 | |||
XP_035542417.1 | JrCOL50d | 453 | 50.39 | 5.37 | 细胞核 | |||
XP_018849735.1 | JrCOL50e | 518 | 57.90 | 5.72 | 细胞核 | |||
XP_018849736.1 | JrCOL50f | 518 | 57.90 | 5.72 | 细胞核 | |||
JrCOL51 | LOC109007532 | Chr.16:6094342..6099323 | XP_018842768.2 | JrCOL51 | 709 | 78.92 | 7.28 | 细胞核 |
JrCOL52 | LOC109008337 | Chr.16:22543220..22546364 | XP_018843939.1 | JrCOL52 | 475 | 52.12 | 6.67 | 细胞核 |
JrCOL53a-d | LOC109006537 | Chr.16:24449761..24454424 | XP_018841385.2 | JrCOL53a | 209 | 23.20 | 6.65 | 细胞核,细胞质 |
XP_035542160.1 | JrCOL53b | 244 | 27.19 | 7.48 | 细胞质 | |||
XP_035542161.1 | JrCOL53c | 244 | 27.19 | 7.48 | 细胞质 | |||
XP_035542162.1 | JrCOL53d | 209 | 23.20 | 6.65 | 细胞核 | |||
JrCOL54a-b | LOC109006510 | Chr.16:24684660..24686424 | XP_018841344.1 | JrCOL54a | 298 | 33.48 | 6.31 | 细胞核 |
XP_018841345.1 | JrCOL54b | 215 | 23.72 | 6.05 | 细胞核 | |||
JrCOL55 | LOC108984240 | Scaffold_15:88..2233 | XP_018811678.1 | JrCOL55 | 170 | 18.25 | 4.08 | 细胞核 |
表2 核桃的COL基因家族鉴定
Table 2 COL gene family identified in Juglans regia
基因名称 Gene name | 基因ID Gene ID | 染色体位置 Chromosome position | 相关蛋白 Related protein | 蛋白重命名 Protein rename | 氨基酸长度 aa length | 蛋白分子量 Molecular weight/kD | 理论等电点pI | 亚细胞定位Subc- ellular localization |
---|---|---|---|---|---|---|---|---|
JrCOL1 | LOC109001350 | Chr.1:1256032..1258201 | XP_018834143.1 | JrCOL1 | 373 | 41.31 | 5.17 | 叶绿体 |
JrCOL2 | LOC109001381 | Chr.1:1488506..1490928 | XP_035544637.1 | JrCOL2 | 211 | 23.40 | 5.73 | 细胞质 |
JrCOL3 | LOC109009665 | Chr.1:3978995..3983057 | XP_018845790.2 | JrCOL3 | 350 | 39.64 | 8.72 | 细胞核 |
JrCOL4a-d | LOC109010156 | Chr.1:18366251..18370500 | XP_035543378.1 | JrCOL4a | 428 | 47.24 | 5.27 | 细胞核 |
XP_018846422.1 | JrCOL4b | 428 | 47.24 | 5.27 | 细胞核 | |||
XP_018846423.1 | JrCOL4c | 428 | 47.24 | 5.27 | 细胞核 | |||
XP_018846424.1 | JrCOL4d | 428 | 47.24 | 5.27 | 细胞核 | |||
JrCOL5 | LOC109015440 | Chr.1:18372462..18372860 | XP_018853453.2 | JrCOL5 | 132 | 14.50 | 6.85 | 细胞核 |
JrCOL6a-b | LOC108991987 | Chr.1:19317524..19318826 | XP_018821963.2 | JrCOL6a | 237 | 25.91 | 4.42 | 细胞核 |
XP_018821964.2 | JrCOL6b | 236 | 25.78 | 4.42 | 细胞核 | |||
JrCOL7a-c | LOC109000761 | Chr.1:44957519..44963229 | XP_018833298.1 | JrCOL7a | 416 | 45.11 | 5.19 | 细胞核 |
XP_018833299.1 | JrCOL7b | 416 | 45.11 | 5.19 | 细胞核 | |||
XP_018833303.1 | JrCOL7c | 416 | 45.11 | 5.19 | 细胞核 | |||
JrCOL8 | LOC109013302 | Chr.2:7141021..7147381 | XP_018850883.1 | JrCOL8 | 539 | 60.48 | 6.22 | 细胞核 |
JrCOL9 | LOC108984132 | Chr.2:11707792..11709460 | XP_018811512.1 | JrCOL9 | 277 | 31.23 | 4.94 | 细胞核 |
JrCO10 | LOC109014521 | Chr.2:30680768..30682741 | XP_018852564.1 | JrCOL10 | 333 | 37.49 | 4.97 | 细胞核 |
JrCOL11a-b | LOC109014085 | Chr.2:35047335..35048788 | XP_018851954.1 | JrCOL11a | 258 | 27.76 | 4.28 | 细胞核 |
XP_018851962.1 | JrCOL11b | 255 | 27.37 | 4.25 | 细胞核 | |||
JrCOL12a-b | LOC108999038 | Chr.3:18899526..18901860 | XP_018831379.1 | JrCOL12a | 150 | 16.21 | 4.00 | 细胞核,叶绿体 |
XP_035544064.1 | JrCOL12b | 150 | 16.21 | 4.00 | 细胞核,叶绿体 | |||
JrCOL13 | LOC109017893 | Chr.4:14653613..14655876 | XP_035544914.1 | JrCOL13 | 170 | 18.24 | 4.11 | 细胞核 |
JrCOL14 | LOC108989855 | Chr.5:6297905..6299253 | XP_018819161.1 | JrCOL14 | 220 | 26.37 | 4.94 | 细胞核 |
JrCOL15a-b | LOC109022307 | Chr.5:9188186..9197431 | XP_018860727.1 | JrCOL15a | 789 | 86.08 | 6.30 | 叶绿体 |
XP_018860748.1 | JrCOL15b | 778 | 84.89 | 6.35 | 叶绿体 | |||
JrCOL16 | LOC108994549 | Chr.6:15543376..15545802 | XP_018825341.2 | JrCOL16 | 444 | 48.81 | 5.88 | 细胞核 |
JrCOL17 | LOC108998057 | Chr.6:29988891..29991120 | XP_018830038.2 | JrCOL17 | 407 | 46.32 | 5.46 | 细胞核 |
JrCOL18 | LOC108992642 | Chr.7:12544651..12554366 | XP_018822785.2 | JrCOL18 | 685 | 75.72 | 6.71 | 细胞核 |
JrCOL19 | LOC108983659 | Chr.7:14608063..14609540 | XP_018810913.1 | JrCOL19 | 266 | 28.49 | 8.73 | 叶绿体 |
JrCOL20 | LOC109005469 | Chr.7:32506953..32517320 | XP_018839979.2 | JrCOL20 | 238 | 26.36 | 5.08 | 细胞核 |
JrCOL21 | LOC109019524 | Chr.7:38617705..38622031 | XP_018857358.1 | JrCOL21 | 432 | 46.81 | 4.35 | 叶绿体 |
JrCOL22a-b | LOC109014549 | Chr.7:49644025..49658252 | XP_035548321.1 | JrCOL22a | 780 | 85.75 | 6.42 | 叶绿体 |
XP_035548322.1 | JrCOL22b | 778 | 85.50 | 6.42 | 叶绿体 | |||
JrCOL23 | LOC109000996 | Chr.8:600112..601757 | XP_018833641.1 | JrCOL23 | 345 | 37.98 | 6.16 | 细胞质 |
JrCOL24 | LOC108984911 | Chr.8:6608517..6610109 | XP_018812557.1 | JrCOL24 | 301 | 33.04 | 6.25 | 细胞核 |
JrCOL25a-b | LOC108980667 | Chr.8:24595513..24597399 | XP_035549568.1 | JrCOL25a | 158 | 17.12 | 5.62 | 细胞质 |
XP_035549569.1 | JrCOL25b | 117 | 12.99 | 6.49 | 细胞质 | |||
JrCOL26 | LOC108997344 | Chr.9:19840907..19842615 | XP_018829103.1 | JrCOL26 | 331 | 37.53 | 5.83 | 细胞核 |
JrCOL27 | LOC108985349 | Chr.9:21400481..21403221 | XP_018813169.1 | JrCOL27 | 295 | 31.92 | 5.55 | 叶绿体 |
JrCOL28 | LOC108986213 | Chr.10:2554388..2557653 | XP_018814300.1 | JrCOL28 | 299 | 34.08 | 6.43 | 细胞核 |
JrCOL29a-c | LOC108984383 | Chr.10:13263191..13268808 | XP_018811858.2 | JrCOL29a | 436 | 47.97 | 6.55 | 细胞质 |
XP_035550266.1 | JrCOL29b | 436 | 47.97 | 6.55 | 细胞质 | |||
XP_035550267.1 | JrCOL29c | 408 | 44.52 | 5.70 | 细胞核 | |||
JrCOL30a-b | LOC109017472 | Chr.10:13911798..13913088 | XP_018855279.1 | JrCOL30a | 260 | 28.60 | 4.37 | 叶绿体 |
XP_018855280.1 | JrCOL30b | 259 | 28.48 | 4.37 | 叶绿体 | |||
JrCOL31a-c | LOC109000741 | Chr.10:37518173..37524266 | XP_018833263.1 | JrCOL31a | 409 | 44.45 | 5.96 | 细胞核 |
XP_018833265.1 | JrCOL31b | 409 | 44.45 | 5.96 | 细胞核 | |||
XP_018833266.1 | JrCOL31c | 409 | 44.45 | 5.96 | 细胞核 | |||
JrCOL32a-d | LOC108992026 | Chr.11:8894066..8897365 | XP_018822007.1 | JrCOL32a | 194 | 21.42 | 6.43 | 叶绿体 |
XP_018822008.1 | JrCOL32b | 192 | 21.17 | 6.23 | 叶绿体 | |||
XP_018822009.1 | JrCOL32c | 186 | 20.48 | 6.43 | 叶绿体 | |||
XP_018822010.1 | JrCOL32d | 184 | 20.22 | 6.23 | 叶绿体 | |||
JrCOL33 | LOC108992144 | Chr.11:23645881..23650290 | XP_018822151.1 | JrCOL33 | 364 | 40.75 | 5.51 | 细胞核 |
JrCOL34 | LOC108991312 | Chr.11:29101394..29103034 | XP_018821038.1 | JrCOL34 | 300 | 33.48 | 5.99 | 细胞核 |
JrCOL35 | LOC108998881 | Chr.11:32536877..32538774 | XP_018831163.1 | JrCOL35 | 373 | 40.72 | 5.86 | 叶绿体 |
JrCOL36 | LOC109003090 | Chr.11:33364576..33368847 | XP_018836611.1 | JrCOL36 | 409 | 45.46 | 8.94 | 细胞核 |
JrCOL37 | LOC109005190 | Chr.11:36684126..36685787 | XP_018839540.1 | JrCOL37 | 341 | 37.58 | 6.87 | 叶绿体 |
JrCOL38a-b | LOC108982143 | Chr.12:8537584..8542475 | XP_018808983.1 | JrCOL38a | 671 | 74.38 | 5.97 | 细胞核 |
XP_035539295.1 | JrCOL38b | 652 | 72.23 | 6.08 | 细胞核 | |||
JrCOL39 | LOC108992474 | Chr.12:13436453..13439714 | XP_018822597.1 | JrCOL39 | 237 | 26.10 | 4.91 | 细胞核 |
JrCOL40a-b | LOC108981163 | Chr.12:23463905..23468842 | XP_018807797.1 | JrCOL40a | 423 | 46.11 | 4.45 | 细胞核 |
XP_018807806.1 | JrCOL40b | 418 | 45.53 | 4.53 | 细胞核 | |||
JrCOL41a-b | LOC108986927 | Chr.13:5686536..5688137 | XP_018815283.1 | JrCOL41a | 293 | 32.61 | 5.70 | 细胞核 |
XP_018815285.1 | JrCOL41b | 206 | 22.72 | 5.57 | 细胞核 | |||
JrCOL42 | LOC109008345 | Chr.13:8537826..8540900 | XP_018843947.2 | JrCOL42 | 475 | 51.86 | 5.92 | 细胞核 |
JrCOL43a-b | LOC108992288 | Chr.13:31296667..31301250 | XP_018822350.1 | JrCOL43a | 691 | 77.57 | 6.80 | 细胞核 |
XP_018822351.1 | JrCOL43b | 689 | 77.43 | 6.80 | 细胞核 | |||
JrCOL44a-d | LOC109007733 | Chr.13:37236625..37242278 | XP_018843087.1 | JrCOL44a | 518 | 57.78 | 5.61 | 细胞核 |
XP_018843095.1 | JrCOL44b | 518 | 57.78 | 5.61 | 细胞核 | |||
XP_018843104.1 | JrCOL44c | 518 | 57.78 | 5.61 | 细胞核 | |||
XP_018843113.1 | JrCOL44d | 518 | 57.78 | 5.61 | 细胞核 | |||
JrCOL45 | LOC109013207 | Chr.13:38161618..38165624 | XP_018850755.1 | JrCOL45 | 287 | 32.38 | 5.25 | 细胞核 |
JrCOL46 | LOC108996999 | Chr.14:6471961..6473893 | XP_018828617.1 | JrCOL46 | 218 | 25.99 | 5.27 | 细胞核 |
JrCOL47 | LOC108993945 | Chr.14:10350983..10353374 | XP_018824556.1 | JrCOL47 | 251 | 28.23 | 8.27 | 细胞核 |
JrCOL48 | LOC109013626 | Chr.15:14881124..14883084 | XP_018851315.1 | JrCOL48 | 464 | 52.39 | 5.09 | 细胞核,细胞质 |
JrCOL49 | LOC109004589 | Chr.16:1780812..1784882 | XP_018838732.1 | JrCOL49 | 279 | 31.45 | 5.27 | 细胞核 |
JrCOL50a-f | LOC109012501 | Chr.16:2630759..2636833 | XP_035542414.1 | JrCOL50a | 518 | 57.90 | 5.72 | 细胞核 |
XP_035542415.1 | JrCOL50b | 492 | 54.89 | 5.87 | 细胞核 | |||
XP_035542416.1 | JrCOL50c | 489 | 54.51 | 5.83 | 细胞核 | |||
XP_035542417.1 | JrCOL50d | 453 | 50.39 | 5.37 | 细胞核 | |||
XP_018849735.1 | JrCOL50e | 518 | 57.90 | 5.72 | 细胞核 | |||
XP_018849736.1 | JrCOL50f | 518 | 57.90 | 5.72 | 细胞核 | |||
JrCOL51 | LOC109007532 | Chr.16:6094342..6099323 | XP_018842768.2 | JrCOL51 | 709 | 78.92 | 7.28 | 细胞核 |
JrCOL52 | LOC109008337 | Chr.16:22543220..22546364 | XP_018843939.1 | JrCOL52 | 475 | 52.12 | 6.67 | 细胞核 |
JrCOL53a-d | LOC109006537 | Chr.16:24449761..24454424 | XP_018841385.2 | JrCOL53a | 209 | 23.20 | 6.65 | 细胞核,细胞质 |
XP_035542160.1 | JrCOL53b | 244 | 27.19 | 7.48 | 细胞质 | |||
XP_035542161.1 | JrCOL53c | 244 | 27.19 | 7.48 | 细胞质 | |||
XP_035542162.1 | JrCOL53d | 209 | 23.20 | 6.65 | 细胞核 | |||
JrCOL54a-b | LOC109006510 | Chr.16:24684660..24686424 | XP_018841344.1 | JrCOL54a | 298 | 33.48 | 6.31 | 细胞核 |
XP_018841345.1 | JrCOL54b | 215 | 23.72 | 6.05 | 细胞核 | |||
JrCOL55 | LOC108984240 | Scaffold_15:88..2233 | XP_018811678.1 | JrCOL55 | 170 | 18.25 | 4.08 | 细胞核 |
图1 拟南芥、核桃COL基因家族的系统发育关系 图中红色实心圆代表核桃COL蛋白,蓝色实心圆代表拟南芥COL蛋白
Fig. 1 Phylogenetic relationship of the COL gene family of Arabidopsis and walnut(Juglans regia) The red solid dot represents walnut COL protein,blue solid dot represents Arabidopsis COL protein
图2 JrCOL蛋白进化(A)、保守基序分布(B)、基因结构(C)及保守结构域(D)分析
Fig. 2 Analysis of JrCOL protein evolution(A),conserved motif(B),gene structure(C)and conserved domain(D)
顺式作用元件 Cis-element | 基因数目 Gene number | 序列 Sequence | 功能 Function | 顺式作用元件类型 Cis-element type |
---|---|---|---|---|
ARE | 46 | AAACCA | 厌氧诱导所必需的顺式作用调控元件 cis-elements for anaerobic induction | 胁迫相关响应元件 Abiotic and biotic stresses |
DRE core | 10 | GCCGAC | 脱水响应元件Dehydration-responsive element | |
LTR | 18 | CCGAAA | 参与低温反应的顺式作用元件 Low-temperature responsive cis-element | |
MBS | 17 | CAACTG | MYB结合位点参与干旱诱导 MYB binding site involved in drought induction | |
MYB | 50 | CAACAG | MYB响应元件MYB-responsive element | |
MYC | 53 | CAATTG | MYC响应元件MYC-responsive element | |
STRE | 40 | AGGGG | 胁迫响应元件Stress-responsive element | |
TC-rich repeat | 10 | GTTTTCTTAC | 参与防御和胁迫反应的顺式作用元素 Defense and stress-responsive element | |
WUN-motif | 21 | AAATTACT | 创伤响应元件Mechanical wounding element | |
ABRE | 43 | ACGTG | 脱落酸反应中的顺式作用元件ABA-responsive element | 激素相关响应元件 Phytohormone responsive |
ERE | 32 | ATTTTAAA | 乙烯响应元件Ethephon-responsive element | |
TGA-element | 14 | AACGAC | 生长素响应元件Auxin-responsive element | |
TCA-element | 16 | CAGAAAAGGA | 参与水杨酸反应的顺式作用元件 Salicylic-acid-responsive element | |
TGACG-motif | 33 | TGACG | 参与MeJA反应的顺调控元件MeJA-responsive element | |
CGTCA-motif | 34 | CGTCA | 参与MeJA反应的顺调控元件MeJA-responsive element | |
P-box | 16 | CCTTTTG | 赤霉素响应元件Gibberellin-responsive element | |
TATC-box | 9 | TATCCCA | 赤霉素响应元件Gibberellin-responsive element | |
Box 4 | 42 | ATTAAT | 与光反应有关的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness | 生长发育相关响应元件 Plant growth and development |
CAT-box | 13 | GCCACT | 与分生组织表达相关的调控元件 Involved in Meristem expression element | |
GA-motif | 9 | ATAGATAA | 光响应模块的一部分Part of a light-responsive module | |
GATA-motif | 21 | AAGGATAAGG | 光响应元件的一部分Part of a light-responsive element | |
G-box | 49 | CACGTC | 参与光响应的调控元件Involved in light-responsive element | |
GT1-motif | 23 | GGTTAAT | 光响应元件Light-responsive element |
表3 JrCOL启动子上顺式作用元件的预测
Table 3 Prediction of cis-element in the JrCOL promoter
顺式作用元件 Cis-element | 基因数目 Gene number | 序列 Sequence | 功能 Function | 顺式作用元件类型 Cis-element type |
---|---|---|---|---|
ARE | 46 | AAACCA | 厌氧诱导所必需的顺式作用调控元件 cis-elements for anaerobic induction | 胁迫相关响应元件 Abiotic and biotic stresses |
DRE core | 10 | GCCGAC | 脱水响应元件Dehydration-responsive element | |
LTR | 18 | CCGAAA | 参与低温反应的顺式作用元件 Low-temperature responsive cis-element | |
MBS | 17 | CAACTG | MYB结合位点参与干旱诱导 MYB binding site involved in drought induction | |
MYB | 50 | CAACAG | MYB响应元件MYB-responsive element | |
MYC | 53 | CAATTG | MYC响应元件MYC-responsive element | |
STRE | 40 | AGGGG | 胁迫响应元件Stress-responsive element | |
TC-rich repeat | 10 | GTTTTCTTAC | 参与防御和胁迫反应的顺式作用元素 Defense and stress-responsive element | |
WUN-motif | 21 | AAATTACT | 创伤响应元件Mechanical wounding element | |
ABRE | 43 | ACGTG | 脱落酸反应中的顺式作用元件ABA-responsive element | 激素相关响应元件 Phytohormone responsive |
ERE | 32 | ATTTTAAA | 乙烯响应元件Ethephon-responsive element | |
TGA-element | 14 | AACGAC | 生长素响应元件Auxin-responsive element | |
TCA-element | 16 | CAGAAAAGGA | 参与水杨酸反应的顺式作用元件 Salicylic-acid-responsive element | |
TGACG-motif | 33 | TGACG | 参与MeJA反应的顺调控元件MeJA-responsive element | |
CGTCA-motif | 34 | CGTCA | 参与MeJA反应的顺调控元件MeJA-responsive element | |
P-box | 16 | CCTTTTG | 赤霉素响应元件Gibberellin-responsive element | |
TATC-box | 9 | TATCCCA | 赤霉素响应元件Gibberellin-responsive element | |
Box 4 | 42 | ATTAAT | 与光反应有关的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness | 生长发育相关响应元件 Plant growth and development |
CAT-box | 13 | GCCACT | 与分生组织表达相关的调控元件 Involved in Meristem expression element | |
GA-motif | 9 | ATAGATAA | 光响应模块的一部分Part of a light-responsive module | |
GATA-motif | 21 | AAGGATAAGG | 光响应元件的一部分Part of a light-responsive element | |
G-box | 49 | CACGTC | 参与光响应的调控元件Involved in light-responsive element | |
GT1-motif | 23 | GGTTAAT | 光响应元件Light-responsive element |
图3 核桃COL基因的复制模式分析 A:核桃基因组内JrCOL基因的同源性分析,灰色线表示核桃基因组中的所有共线性区块,而绿色线表示发生复制的COL基因对;B:拟南芥、核桃和葡萄COL基因的共线性分析,背景中的灰色线表示核桃和其他植物基因组中的共线性区块,而红色和紫色的连接线则突出显示了COL相关的共线性基因对
Fig. 3 Duplication models of the COL genes in walnut A:Synteny analysis of JrCOL genes. Gray lines indicate all synteny blocks in the walnut genome,whereas segmental duplicated homologous genes are indicated with green line. B:Synteny analysis of COL genes between Arabidopsis,walnut and grape. The gray lines in the background indicate the collinear blocks within walnut and other plant genomes,while the red and purple lines highlight the syntenic COL gene pairs
同源基因对 Homologous gene pair | Ka | Ks | Ka/Ks |
---|---|---|---|
JrCOL3 & JrCOL28 | 0.164 | 0.426 | 0.386 |
JrCOL4a-d & JrCOL29a-c | 0.120 | 0.310 | 0.388 |
JrCOL6a-b & JrCOL11a-b | 0.371 | 1.753 | 0.212 |
JrCOL6a-b & JrCOL12a-b | —— | —— | —— |
JrCOL6a-b & JrCOL13 | —— | —— | —— |
JrCOL6a-b & JrCOL30a-b | 0.163 | 0.405 | 0.402 |
JrCOL7a-c & JrCOL31a-c | 0.082 | 0.316 | 0.258 |
JrCOL10 & JrCOL26 | 0.105 | 0.349 | 0.301 |
JrCOL11a-b & JrCOL12a-b | —— | —— | —— |
JrCOL11a-b & JrCOL13 | —— | —— | —— |
JrCOL12a-b & JrCOL13 | 0.136 | 0.690 | 0.197 |
JrCOL14 & JrCOL46 | 0.095 | 0.467 | 0.203 |
JrCOL16 & JrCOL17 | 0.315 | 2.218 | 0.142 |
JrCOL17 & JrCOL48 | 0.158 | 0.352 | 0.449 |
JrCOL18 & JrCOL38a-b | 0.164 | 0.339 | 0.484 |
JrCOL18 & JrCOL43a-b | 0.445 | 1.224 | 0.364 |
JrCOL21 & JrCOL40a-b | 0.081 | 0.309 | 0.263 |
JrCOL23 & JrCOL37 | 0.111 | 0.506 | 0.220 |
JrCOL24 & JrCOL25a-b | —— | —— | —— |
JrCOL24 & JrCOL34 | 0.123 | 0.347 | 0.354 |
JrCOL24 & JrCOL41a-b | 0.351 | 5.257 | 0.067 |
JrCOL24 & JrCOL54a-b | 0.338 | 2.787 | 0.121 |
JrCOL25a-b & JrCOL32a-d | 0.061 | 0.341 | 0.179 |
JrCOL25a-b & JrCOL53a-d | 0.164 | 1.270 | 0.129 |
JrCOL32a-d & JrCOL34 | 0.627 | 3.152 | 0.199 |
JrCOL32a-d & JrCOL53a-d | 0.223 | 1.214 | 0.184 |
JrCOL34 & JrCOL41a-b | 0.385 | 1.660 | 0.232 |
JrCOL34 & JrCOL54a-b | 0.342 | 1.956 | 0.175 |
JrCOL38a-b & JrCOL43a-b | 0.418 | 1.346 | 0.310 |
JrCOL38a-b & JrCOL51 | 0.429 | 1.279 | 0.336 |
JrCOL41a-b & JrCOL54a-b | 0.163 | 0.466 | 0.349 |
JrCOL42 & JrCOL52 | 0.111 | 0.433 | 0.256 |
JrCOL43a-b & JrCOL51 | 0.136 | 0.329 | 0.413 |
JrCOL44a-d & JrCOL50a-f | 0.140 | 0.316 | 0.442 |
JrCOL45 & JrCOL49 | 0.069 | 0.426 | 0.161 |
表4 JrCOL基因家族Ka/Ks值反映的复制时间分析
Table 4 Analysis of duplication time reflected from Ka/Ks values of JrCOL gene family
同源基因对 Homologous gene pair | Ka | Ks | Ka/Ks |
---|---|---|---|
JrCOL3 & JrCOL28 | 0.164 | 0.426 | 0.386 |
JrCOL4a-d & JrCOL29a-c | 0.120 | 0.310 | 0.388 |
JrCOL6a-b & JrCOL11a-b | 0.371 | 1.753 | 0.212 |
JrCOL6a-b & JrCOL12a-b | —— | —— | —— |
JrCOL6a-b & JrCOL13 | —— | —— | —— |
JrCOL6a-b & JrCOL30a-b | 0.163 | 0.405 | 0.402 |
JrCOL7a-c & JrCOL31a-c | 0.082 | 0.316 | 0.258 |
JrCOL10 & JrCOL26 | 0.105 | 0.349 | 0.301 |
JrCOL11a-b & JrCOL12a-b | —— | —— | —— |
JrCOL11a-b & JrCOL13 | —— | —— | —— |
JrCOL12a-b & JrCOL13 | 0.136 | 0.690 | 0.197 |
JrCOL14 & JrCOL46 | 0.095 | 0.467 | 0.203 |
JrCOL16 & JrCOL17 | 0.315 | 2.218 | 0.142 |
JrCOL17 & JrCOL48 | 0.158 | 0.352 | 0.449 |
JrCOL18 & JrCOL38a-b | 0.164 | 0.339 | 0.484 |
JrCOL18 & JrCOL43a-b | 0.445 | 1.224 | 0.364 |
JrCOL21 & JrCOL40a-b | 0.081 | 0.309 | 0.263 |
JrCOL23 & JrCOL37 | 0.111 | 0.506 | 0.220 |
JrCOL24 & JrCOL25a-b | —— | —— | —— |
JrCOL24 & JrCOL34 | 0.123 | 0.347 | 0.354 |
JrCOL24 & JrCOL41a-b | 0.351 | 5.257 | 0.067 |
JrCOL24 & JrCOL54a-b | 0.338 | 2.787 | 0.121 |
JrCOL25a-b & JrCOL32a-d | 0.061 | 0.341 | 0.179 |
JrCOL25a-b & JrCOL53a-d | 0.164 | 1.270 | 0.129 |
JrCOL32a-d & JrCOL34 | 0.627 | 3.152 | 0.199 |
JrCOL32a-d & JrCOL53a-d | 0.223 | 1.214 | 0.184 |
JrCOL34 & JrCOL41a-b | 0.385 | 1.660 | 0.232 |
JrCOL34 & JrCOL54a-b | 0.342 | 1.956 | 0.175 |
JrCOL38a-b & JrCOL43a-b | 0.418 | 1.346 | 0.310 |
JrCOL38a-b & JrCOL51 | 0.429 | 1.279 | 0.336 |
JrCOL41a-b & JrCOL54a-b | 0.163 | 0.466 | 0.349 |
JrCOL42 & JrCOL52 | 0.111 | 0.433 | 0.256 |
JrCOL43a-b & JrCOL51 | 0.136 | 0.329 | 0.413 |
JrCOL44a-d & JrCOL50a-f | 0.140 | 0.316 | 0.442 |
JrCOL45 & JrCOL49 | 0.069 | 0.426 | 0.161 |
图5 ‘新新2号'JrCOL基因在不同组织中的相对表达量 不同小写字母字母表示处理间差异显著(P<0.05),下同
Fig. 5 Relative expressions of JrCOL genes in Juglans regia cv. Xinxin No. 2 in different tissues Different lowercase letters indicate significant differences between treatments(P<0.05). The same below
图6 ‘新新2号'雌花芽不同发育阶段JrCOL基因家族表达量及其回归分析
Fig. 6 Expressions and regression analysis of the JrCOL gene families at different developmental stages of Juglans regia cv. Xinxin No. 2
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