Identification of PLR gene family and expression of responsive hormones in <i>Phoebe bournei</i>

WANG Qianqing; ZHANG Yuting; ZHANG Junhong; LIU Hui; TONG Zaikang

doi:10.11833/j.issn.2095-0756.20220351

Volume 39 Issue 6

Nov. 2022

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WANG Qianqing, ZHANG Yuting, ZHANG Junhong, LIU Hui, TONG Zaikang. Identification of PLR gene family and expression of responsive hormones in Phoebe bournei[J]. Journal of Zhejiang A&F University, 2022, 39(6): 1173-1182. doi: 10.11833/j.issn.2095-0756.20220351

Citation:

WANG Qianqing, ZHANG Yuting, ZHANG Junhong, LIU Hui, TONG Zaikang. Identification of PLR gene family and expression of responsive hormones in Phoebe bournei[J]. Journal of Zhejiang A&F University, 2022, 39(6): 1173-1182. doi: 10.11833/j.issn.2095-0756.20220351

Identification of PLR gene family and expression of responsive hormones in Phoebe bournei

doi: 10.11833/j.issn.2095-0756.20220351

State Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China

Received Date: 2022-05-16
Accepted Date: 2022-08-08
Rev Recd Date: 2022-07-15

Available Online: 2022-11-21

Publish Date: 2022-12-20

Abstract

Objective Pinoresinol-laricisinol reductase (PLR) is a key enzyme gene involved in lignan biosynthesis. This study aims to clear the characteristics of PLR gene family in Phoebe bournei and the expression patterns under different hormone treatments, so as provide basis for further exploring the biological function of PLR gene family in P. bournei. Method The members of PLR gene family in P. bournei genome were identified by bioinformatics methods, and the gene structure, conserved motifs, chromosome location, phylogeny, cis acting elements of promoter and hormone response were analyzed. Result A total of 8 PLR (PbPLR) members were identified from the genome of P. bournei, which were unevenly distributed on chromosomes 1, 2, 5 and 10. Phylogenetic analysis indicated that PbPLR function may be related to substrate stereoselectivity. Promoter element analysis showed that there were hormone response elements such as bascisic acid (ABA), salicylic acid (SA), and methyl jasmonate (MeJA) in the promoter region of PbPLRs. Gene expression analysis showed that the expression of PbPLRs gene had the characteristics of tissue specificity and hormone response. The expression of PbPLRs gene was the highest in roots, followed by stems and leaves. The expression of PbPLR2 was the strongest when treated with 3 hormones. Conclusion The 8 PbPLRs gene identified from the genome of P. bournei have different gene characteristics, and different members may have different stereoselectivity of catalytic enzymes. The expression patterns of members are diverse, which may be induced by various hormones and have tissue specificity. [Ch, 6 fig. 2 tab. 25 ref.]
- Phoebe bournei,
- PLR gene family,
- phylogenetics,
- hormone treatment,
- expression analysis

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沈阳化工大学材料科学与工程学院沈阳 110142

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不同树种木材的天然耐腐性差异大，据统计每年全球因木材腐朽造成的损失占采伐利用木材的10%以上，严重影响木材的加工利用^[1]。木材腐朽主要由腐朽菌和变色菌等真菌侵蚀木材细胞所引起，而木材的天然耐腐性则依靠自身代谢物的生物杀菌、抑菌活性来实现^[2]。闽楠Phoebe bournei为樟科Lauraceae楠属Phoebe植物，是中国特有的珍稀用材与绿化树种。楠木具有极强耐腐性，阴沉木历经千年不腐，被誉为“木中之玉”，市场俗称“金丝楠乌木”^[3]。闽楠和桢楠Phoebe zhennan作为“金丝楠木”最主要的产材树种，树干挺拔，木材颜色呈淡黄色，坚硬细密，纹理美观具有芳香香气，属上等家具良材^[4]。因此，闽楠可作为研究木材天然耐腐性的良好材料，闽楠全基因组测序的完成为基因挖掘和功能研究提供了重要基础。

已有研究发现：从木材分离的木脂素(lignan)类化合物具有抗菌活性，参与植物防御系统，增强木材耐久性和抗真菌能力。例如北美乔柏Thuja plicata心材中提取的木脂素通过清除自由基和螯合亚铁来干扰真菌氧化还原循环提高木材耐腐性^[5]，南洋杉Araucaria araucana中提取的松脂素(pinoresinol)和开环落叶松脂素(secoisolariciresinol)对木材腐朽菌具有较好抗性^[6]。然而，关于木脂素生物合成调控途径尚不明确，从而限制其深入研究与生产应用。

松脂素还原酶Pinoresinol-lariciresinol reductase (PLR)是一种编码松脂醇的落叶松脂素还原酶基因，属木脂素生物合成的关键酶，一分子松脂素在PLR松脂素-落叶松脂素还原酶作用下经两步还原成落叶松脂素和异落叶松脂素，也是木脂素生物合成途径中重要限速步骤之一。目前已从模式植物拟南芥Arabidopsis thaliana^[7]和富含木脂素的植物(连翘Forsythia suspense^[8-9]、菘蓝Isatis indigotica^[10]、亚麻属Linum^[11-13])中克隆得到PLR基因。迄今为止，所有以重组纯化蛋白为特征的PLR均可以将松脂素还原为落叶松树脂素以及将落叶松树脂素还原为异落叶松树脂素，但AtPrR1对落叶松树脂素的活性比松脂素低约35倍，AtPrR2对作为底物的落叶松树脂素完全没有催化活性。因此，MIN等^[14]将它们命名为松脂醇还原酶，而不是松脂醇-落叶松树脂醇还原酶。PLR的表达受激素的调控，不同物种在不同激素处理下松脂醇-落叶松树脂醇还原酶的表达模式有所不同。在脱落酸(ABA)处理下，LuPLR1的表达增强，LuPLR1的表达似乎是由在种子成熟和休眠中起关键作用的植物激素ABA触发的^[12-13]。所以赤霉酸(GA)对LuPLR1的表达具有相反的调节作用^[13]。在茉莉酸甲酯(MeJA)的处理下，LuPLR2的表达量有明显提高，因为亚麻在MeJA处理和伤害后，LuPLR2基因表达和叶酸蛋白产量增加，这与这些木脂素参与植物防御反应一致。本研究选择耐腐性强的闽楠解析木脂素合成重要限速酶PLR基因，利用闽楠基因组数据，对PLR基因家族成员进行鉴定和基因结构、保守基序及染色体定位分析，构建系统发育进化树预测闽楠PLR (PbPLR)催化酶的立体选择性，为进一步研究PbPLR基因在闽楠木脂素合成过程中的生物学功能提供基础。

3. 讨论

木脂素类化合物具有较为广泛的应用前景，PLR作为植物木脂素生物合成的重要限速酶基因之一，引起了广泛的关注。例如，在亚麻中鉴定到2个PLR基因(LuPLR1和LuPLR2)^[12]，模式植物拟南芥中共有2个PrR基因(AtPrR1和AtPrR2)^[7]，台湾杉中发现了3个PLR基因(TcPLR1、TcPLR2和TcPLR3)等^[21]。本研究从闽楠基因组中共鉴定到8个PbPLRs基因，不均等分布于闽楠的4条染色体上，且均含有PLR家族保守的NADPH结合位点和赖氨酸活化位点。PLR属于short-chain dehydrogenase/reductase (SDR)超级家族，该家族广泛参与植物初级代谢和次级代谢过程，且在高等植物中具有明显功能分化^[22]。闽楠中数量较多的PLR成员可能参与不同代谢通路，并发生功能分化。

利用不同物种PLR蛋白构建系统进化树，发现裸子植物PLR蛋白单独聚为一类，而被子植物PLR系统聚类与催化酶的立体选择性相关，该结果与前人研究结果相一致。NAKATSUBO等^[7]发现被子植物PLR (FiPLR1、LaPLR1、LuPLR1、LpPLR1、AtPrR1和AtPrR2)和裸子植物PLR (TpPLR1和TpPLR2)分别聚类。PLR催化酶的底物立体选择性直接决定了下游木脂素的结构骨架，是形成木脂素类化合物结构多样性的关键节点。PLR立体选择性最早是在金钟连翘Forsythia intermedia中发现，功能研究发现2个FiPLR都偏好选择催化(+)-松脂醇和(+)-落叶松脂醇为(−)-异落叶松脂醇^[9]。随后，在其他物种中鉴定的PLRs同样具有明显立体选择性^[23]。拟南芥AtPrR1可催化2种松脂醇异构体，而AtPrR2仅能催化(−)-松脂醇形成(−)-落叶松脂醇^[7]。根据已有催化酶的功能研究，推测闽楠PbPLR1、PbPLR6、PbPLR7和PbPLR8偏好选择(+)-松脂醇和(+)-落叶松脂醇，而PbPLR2、PbPLR3、PbPLR4和PbPLR5聚类的PLR蛋白的立体选择性不同，无法通过系统进化树推测其底物偏好性。FUJITA等^[24]研究发现：北美乔柏的TpPLRs聚类为一个分支却具有相反的立体选择性。PLR家族成员的蛋白序列差异与底物立体选择性并不完全等同，对底物的选择偏好性可能随物种共同经历多次进化；或形成于物种进化的早期，随后的碱基突变并不影响其催化功能^[25]。

通过启动子顺式作用元件分析发现：PbPLR基因的启动子区域具有多种响应元件，推测PbPLR表达模式可能受生物及非生物因素调控。例如光响应、厌氧诱导、耐旱和防御等响应元件，推测PbPLR表达可能受光信号及非生物胁迫等因素影响，并通过次生代谢物的催化合成而参与植物防御系统。此外，PbPLRs基因启动子区域具有多种激素响应调控元件，其中ABA、MeJA和SA等3种响应元件数目最多。不同物种PLR基因表达受不同激素调控。例如，ABA处理下亚麻LuPLR1和菘蓝LiPLR1表达显著上调^{[10, 12-13]}；MeJA诱导LuPLR2、IiPLR1和鬼臼草Podophyllum hexandrum PhPLR表达量上调^{[10, 25]}；而GA对LuPLR1表达起抑制作用^[21]。对PbPLR表达分析表明：PbPLR对ABA、MeJA和SA均有响应，这与启动子元件分析相吻合。MeJA处理下PbPLR2响应最显著，在处理48 h根部表达量最高；ABA处理下PbPLR4响应最为显著，在处理48 h时叶片表达量最高；SA处理下PbPLR6响应最为显著，在处理48 h时茎表达量最高。本研究表明：PbPLR调控模式较为复杂，可能受不同植物激素诱导，同时PbPLRs组织特异性表达可能与其结构功能分化相关但与系统进化分支无关。推测闽楠木脂素合成具有较为复杂的调控网络，需进一步研究其生物合成调控机制。

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引物名称	引物序列(5′→3′)
PbPLR1	F: GGCACAAATCTCATCAGCAA; R: GCCAGCTTTTATGGCCTGTA
PbPLR2	F: TCTGGATTACGTGCAACAGG; R: AGCGTTTGAGATAGTCCTTCG
PbPLR3	F: AGGTCTGGATTACGCACAGC; R: CGTTTGAGATAGTCCTTCACTCG
PbPLR4	F: GGAATTCCTTGCTCAAATGAAA; R: TTTCGAAATTTGCCAGACAAC
PbPLR5	F: GTGGAACAAGAGCCCAAATG; R: CTGGCAAGTGGGTGAAAAAT
PbPLR6	F: ATGGGGTGGATAGCAATGAC; R: CCATAACACGCTCTTTATATGAACC
PbPLR7	F: AAGCAAGCAGTCAAACAACC; R: AAAGAAGCCTACGGCATGG
PbPLR8	F: GTCGCCGGAAACCACTTG; R: AGTTATTTGGTAAACCAACCTTGA
PbEF1α	F: CATTCAAGTATGCGTGGGT; R: ACGGTGACCAGGAGCA

基因名称	平均分子量/kDa	等电点	起始位置	染色体位置	氨基酸数量/个
PbPLR1	35 441.98	5.90	82085409~82088341	chr01	336
PbPLR2	35 320.24	6.23	82091009~82098484	chr01	312
PbPLR3	35 378.05	6.32	111385794~111390362	chr02	312
PbPLR4	37 844.40	7.74	80076163~80082267	chr05	312
PbPLR5	35 378.05	6.32	81056124~81058143	chr05	284
PbPLR6	34 970.24	6.41	52893843~52901318	chr10	316
PbPLR7	34 910.23	6.10	52876836~52885817	chr10	316
PbPLR8	13 320.63	8.76	52801277~52802274	chr10	123

Identification of PLR gene family and expression of responsive hormones in Phoebe bournei

doi: 10.11833/j.issn.2095-0756.20220351

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通讯作者: 陈斌, bchen63@163.com

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Proportional views

Identification of PLR gene family and expression of responsive hormones in Phoebe bournei

doi: 10.11833/j.issn.2095-0756.20220351

State Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China

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1.1. 闽楠PLR基因家族成员鉴定

1.2. PbPLR基因染色体定位与基因复制分析

1.3. PbPLR基因结构及蛋白保守基序分析

1.4. PbPLR家族成员的系统发育树构建及聚类分析

1.5. PbPLR基因的启动子分析

1.6. PbPLR基因对激素处理的响应

2.1. 闽楠PLR家族成员鉴定

2.2. PbPLR基因染色体定位与基因复制分析

2.3. PbPLR基因结构及蛋白保守基序的分析

2.4. PbPLR家族系统进化分析

2.5. PbPLR基因顺式作用元件分析

2.6. 不同激素处理下的PbPLR表达分析

Catalog

Identification of PLR gene family and expression of responsive hormones in Phoebe bournei

doi: 10.11833/j.issn.2095-0756.20220351

Abstract

Relative Article

References

Proportional views

通讯作者: 陈斌, bchen63@163.com

Related

Proportional views

Identification of PLR gene family and expression of responsive hormones in Phoebe bournei

doi: 10.11833/j.issn.2095-0756.20220351

State Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China

HTML

1.1. 闽楠PLR基因家族成员鉴定

1.2. PbPLR基因染色体定位与基因复制分析

1.3. PbPLR基因结构及蛋白保守基序分析

1.4. PbPLR家族成员的系统发育树构建及聚类分析

1.5. PbPLR基因的启动子分析

1.6. PbPLR基因对激素处理的响应

2.1. 闽楠PLR家族成员鉴定

2.2. PbPLR基因染色体定位与基因复制分析

2.3. PbPLR基因结构及蛋白保守基序的分析

2.4. PbPLR家族系统进化分析

2.5. PbPLR基因顺式作用元件分析

2.6. 不同激素处理下的PbPLR表达分析

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