二化螟味觉受体基因鉴定、克隆与表达模式分析

doi:10.3864/j.issn.0578-1752.2023.13.006

摘要/Abstract

摘要：

【背景】二化螟（Chilo suppressalis）是我国水稻上的重要害虫之一，味觉受体（gustatory receptor，GR）基因在昆虫取食、产卵等过程中发挥重要作用。【目的】基于组学数据鉴定并克隆二化螟GR基因序列，明确其在不同发育阶段和成虫不同组织中的表达特性，为后续深入研究二化螟GR基因的功能打下基础。【方法】结合二化螟转录组数据和其他昆虫的GR基因序列，通过多重序列比对，鉴定二化螟GR基因。利用RT-PCR方法克隆获得二化螟GR基因的完整开放阅读框（open reading frame，ORF）序列，运用生物信息学分析工具对二化螟GR基因编码的氨基酸序列进行分子生物学特征、结构域等分析；基于最大似然法构建二化螟GR基因蛋白序列与其他昆虫GR基因的系统进化树；利用实时荧光定量PCR（RT-qPCR）方法分析GR基因在二化螟不同发育阶段（1—6龄幼虫和雌、雄成虫）和成虫不同组织（雌、雄成虫的触角、头、翅、腹、足）中的表达模式。【结果】鉴定并克隆得到5个二化螟GR基因，分别命名为CsupGR1—5，ORF长度为1 122—1 428 bp，编码氨基酸序列长度为373—475 aa，其中CsupGR2、CsupGR4、CsupGR5具有7个跨膜结构域，CsupGR1、CsupGR3具有8个跨膜结构域。系统进化分析显示，CsupGR1、CsupGR2与黑腹果蝇DmelGR63a及小菜蛾PxylGR7、PxylGR8亲缘关系较近，属于CO₂受体家族；CsupGR3与黑腹果蝇DmelGR43a及家蚕BmorGR9、BmorGR10亲缘关系较近，属于果糖/肌醇受体家族；CsupGR4、CsupGR5与黑腹果蝇DmelGR64及家蚕BmorGR5—8亲缘关系较近，属于糖受体家族。发育期表达谱分析表明，二化螟5个GR基因在不同发育时期均有表达，其中CsupGR1、CsupGR4均在雄成虫中高表达，CsupGR2在1龄幼虫和雄成虫中表达量最高，CsupGR3在成虫中高表达，CsupGR5在4龄幼虫中表达量最高；组织表达谱分析表明，5个GR基因在雌、雄成虫的各组织均有表达，其中CsupGR1、CsupGR5在成虫头部表达量高，CsupGR2—4在成虫触角部位高表达。【结论】鉴定克隆的5个二化螟GR基因均具有昆虫味觉受体基因的典型特征，且在成虫触角或头部高表达，推测这5个基因可能与二化螟识别和适应寄主植物有关。

关键词: 二化螟, 味觉受体, 基因鉴定, 序列分析, 表达谱

Abstract:

【Background】The rice striped stem borer, Chilo suppressalis, is one of the important rice pests in China. The gustatory receptors (GRs) play important roles in insect feeding, oviposition, and so on.【Objective】Based on transcriptomic and genomic data, the gustatory receptor gene sequences of C. suppressalis were identified and cloned in this study. Then, their expression characteristics at different developmental stages and in different tissues of adults were clarified. These results will lay a foundation for further study on the function of gustatory receptor genes in C. suppressalis.【Method】Combined with the transcriptome data of C. suppressalis and the GR sequences of other insects, the GR genes in C. suppressalis were identified by multiple sequence alignment. The complete open reading frame (ORF) sequences of the GR genes in C. suppressalis were cloned by RT-PCR, and the molecular biological characteristics and domains of the amino acid sequences encoded by the GR genes in C. suppressalis were analyzed by bioinformatics analysis tools. The phylogenetic tree of GR gene protein sequences of C. suppressalis and other insect GR genes was constructed based on the maximum likelihood method. The expression patterns of GR genes at different developmental stages (1-6 instar larvae, female and male adults) and in different adult tissues (antennae, head, wing, abdomen and leg) of C. suppressalis were determined by real-time quantitative PCR (RT-qPCR).【Result】Five gustatory receptor genes were identified and named as CsupGR1-5. The complete ORFs of these GR genes were between 1 122 and 1 428 bp in length, encoding 373-475 amino acids. Among them, CsupGR2, CsupGR4, CsupGR5 had seven transmembrane domains, CsupGR1, CsupGR3 had eight transmembrane domains. Phylogenetic analysis showed that CsupGR1 and CsupGR2 were closely related to DmelGR63a in Drosophila melanogaster, PxylGR7 and PxylGR8 in Plutella xylostella, belonging to the CO₂ receptor family. CsupGR3 was closely related to DmelGR43a in D. melanogaster, BmorGR9 and BmorGR10 in Bombyx mori, belonging to the fructose/inositol receptor family. CsupGR4 and CsupGR5 were closely related to DmelGR64 in D. melanogaster and BmorGR5-8 in B. mori, belonging to the sugar receptor family. The expression profile analysis showed that the five GR genes were all expressed at different developmental stages in C. suppressalis. Among them, CsupGR1 and CsupGR4 were highly expressed in male adults, CsupGR2 had the highest expression level in the 1st instar larvae and male adults, CsupGR3 was highly expressed in adults, and CsupGR5 had the highest expression level in the 4th instar larvae. Tissue expression profile analysis showed that these five GR genes were expressed in all tissues of male and female adults. CsupGR1 and CsupGR5 were highly expressed in the head of adults, and CsupGR2-4 were highly expressed in the antennae of adults.【Conclusion】The five GR genes of C. suppressalis identified in this study have the typical characteristics of insect gustatory receptors and are highly expressed in the antennae or heads of adults. It is speculated that these five genes may be related to the recognition and adaptation of C. suppressalis to host plants.

Key words: Chilo suppressalis, gustatory receptor (GR), gene identification, sequence analysis, expression profile

黄玉萱, 沈忱, 鞠佳菲, 杨磊, 罗光华, 方继朝. 二化螟味觉受体基因鉴定、克隆与表达模式分析[J]. 中国农业科学, 2023, 56(13): 2504-2517.

HUANG YuXuan, SHEN Chen, JU JiaFei, YANG Lei, LUO GuangHua, FANG JiChao. Identification, Cloning and Expression Profiles of Gustatory Receptor Genes in Chilo suppressalis[J]. Scientia Agricultura Sinica, 2023, 56(13): 2504-2517.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2023.13.006

https://www.chinaagrisci.com/CN/Y2023/V56/I13/2504

图/表 12

表1

图1

表2

表3

图2

图3

图4

图5

图6

图7

图8

图9

参考文献 49

[1]	MAO K K, LI W H, LIAO X, LIU C Y, QIN Y, REN Z J, QIN X Y, WAN H, SHENG F, LI J H. Dynamics of insecticide resistance in different geographical populations of Chilo suppressalis (Lepidoptera: Crambidae) in China 2016-2018. Journal of Economic Entomology, 2019, 112(4): 1866-1874. doi: 10.1093/jee/toz109
[2]	胡君, 陈文明, 张真真, 郑雪松, 靳建超, 苏建亚, 高聪芬, 沈晋良. 长江流域稻区二化螟抗药性监测. 中国水稻科学, 2010, 24(5): 509-515. doi: 10.3969/j.issn.1001-7216.2010.05.011
	HU J, CHEN W M, ZHANG Z Z, ZHENG X S, JIN J C, SU J Y, GAO C F, SHEN J L. Insecticide resistance monitoring of Chilo suppressalis in the drainage area of the Yangtze River, China. Chinese Journal of Rice Science, 2010, 24(5): 509-515. (in Chinese)
[3]	LU Y, ZHENG X, LU Z. Application of vetiver grass Vetiveria zizanioides: Poaceae (L.) as a trap plant for rice stem borer Chilo suppressalis: Crambidae (Walker) in the paddy fields. Journal of Integrative Agriculture, 2019, 18(4): 797-804. doi: 10.1016/S2095-3119(18)62088-X
[4]	ZHAO X, XU X, WANG X G, YIN Y, LI M Y, WU Y Q, LIU Y H, CHENG Q H, GONG C W, SHEN L T. Mechanisms for multiple resistances in field populations of rice stem borer, Chilo suppressalis (Lepidoptera: Crambidae) from Sichuan Province, China. Pesticide Biochemistry and Physiology, 2021, 171: 104720. doi: 10.1016/j.pestbp.2020.104720
[5]	姚诚诚, 杜立啸, 李云河. 虫害诱导植物信息化合物介导的植物间交流及机制. 植物保护, 2021, 47(6): 1-10.
	YAO C C, DU L X, LI Y H. Plant-to-plant communications medicated by herbivore-induced plant volatiles and the mechanisms. Plant Protection, 2021, 47(6): 1-10. (in Chinese)
[6]	王广利, 付小刚, 韩晓霞, 张宇瑶, 魏洪义. 二化螟雌蛾对香根草挥发物的EAG和行为反应. 应用昆虫学报, 2016, 53(1): 148-156.
	WANG G L, FU X G, HAN X X, ZHANG Y Y, WEI H Y. EAG and behavioral responses of Chilo suppressalis females to plant volatiles from Vetiveria zizanioides. Chinese Journal of Applied Entomology, 2016, 53(1): 148-156. (in Chinese)
[7]	HU X Y, SU S L, LIU Q S, JIAO Y Y, PENG Y F, LI Y H, TURLINGS T C J. Caterpillar-induced rice volatiles provide enemy-free space for the offspring of the brown planthopper. eLife, 2020, 9: e55421. doi: 10.7554/eLife.55421
[8]	OZAKI K, RYUDA M, YAMADA A, UTOGUCHI A, ISHIMOTO H, CALAS D, MARION-POLL F, TANIMURA T, YOSHIKAWA H. A gustatory receptor involved in host plant recognition for oviposition of a swallowtail butterfly. Nature Communications, 2011, 2: 542. doi: 10.1038/ncomms1548
[9]	SCHOONHOVEN L M, VAN LOON J J A. An inventory of taste in caterpillars: Each species its own key. Acta Zoologica Academiae Scientiarum Hungaricae, 2002, 48(Suppl. 1): 215-263.
[10]	ZHANG Z J, ZHANG S S, NIU B L, JI D F, LIU X J, LI M W, BAI H, PALLI S R, WANG C Z, TAN A J. A determining factor for insect feeding preference in the silkworm, Bombyx mori. PLoS Biology, 2019, 17(2): e3000162.
[11]	YANG J, GUO H, JIANG N J, TANG R, LI G C, HUANG L Q, VAN LOON J J A, WANG C Z. Identification of a gustatory receptor tuned to sinigrin in the cabbage butterfly Pieris rapae. PLoS Genetics, 2021, 17(7): e1009527. doi: 10.1371/journal.pgen.1009527
[12]	SINGH R N. Neurobiology of the gustatory systems of Drosophila and some terrestrial insects. Microscopy Research and Technique, 1997, 39(6): 547-563. doi: 10.1002/(ISSN)1097-0029
[13]	CLYNE P J, WARR C G, CARLSON J R. Candidate taste receptors in Drosophila. Science, 2000, 287(5459): 1830-1834. doi: 10.1126/science.287.5459.1830
[14]	AGNIHOTRI A R, ROY A A, JOSHI R S. Gustatory receptors in Lepidoptera: Chemosensation and beyond. Insect Molecular Biology, 2016, 25(5): 519-529. doi: 10.1111/imb.12246 pmid: 27228010
[15]	SANCHEZ-GRACIA A, VIEIRA F G, ROZAS J. Molecular evolution of the major chemosensory gene families in insects. Heredity, 2009, 103(3): 208-216. doi: 10.1038/hdy.2009.55
[16]	ROBERTSON H M, WARR C G, CARLSON J R. Molecular evolution of the insect chemoreceptor gene superfamily in Drosophila melanogaster. Proceedings of the National Academy of Sciences of the United States of America, 2003, 100(Suppl. 2): 14537-14542.
[17]	GUO H, CHENG T, CHEN Z, JIANG L, GUO Y, LIU J, LI S, TANIAI K, ASAOKA K, KADONO-OKUDA K, et al. Expression map of a complete set of gustatory receptor genes in chemosensory organs of Bombyx mori. Insect Biochemistry and Molecular Biology, 2017, 82: 74-82. doi: 10.1016/j.ibmb.2017.02.001
[18]	ZHAN S, MERLIN C, BOORE J L, REPPERT S M. The monarch butterfly genome yields insights into long-distance migration. Cell, 2011, 147(5): 1171-1185. doi: 10.1016/j.cell.2011.09.052 pmid: 22118469
[19]	BRISCOE A D, MACIAS-MUÑOZ A, KOZAK K M, WALTERS J R, YUAN F, JAMIE G A, MARTIN S H, DASMAHAPATRA K K, FERGUSON L C, MALLET J, JACQUIN-JOLY E, JIGGINS C D. Female behaviour drives expression and evolution of gustatory receptors in butterflies. PLOS Genetics, 2013, 9(7): e1003620. doi: 10.1371/journal.pgen.1003620
[20]	YOU M, YUE Z, HE W, YANG X, YANG G, XIE M, ZHAN D, BAXTER S W, VASSEUR L, GURR G M, et al. A heterozygous moth genome provides insights into herbivory and detoxification. Nature Genetics, 2013, 45(2): 220-225. doi: 10.1038/ng.2524 pmid: 23313953
[21]	LI L L, XU J W, YAO W C, YANG H H, DEWER Y, ZHANG F, ZHU X Y, ZHANG Y N. Chemosensory genes in the head of Spodoptera litura larvae. Bulletin of Entomological Research, 2021, 111(4): 454-463. doi: 10.1017/S0007485321000109
[22]	DU L X, ZHAO X C, LIANG X Z, GAO X W, LIU Y, WANG G R. Identification of candidate chemosensory genes in Mythimna separata by transcriptomic analysis. BMC Genomics, 2018, 19: 518. doi: 10.1186/s12864-018-4898-0
[23]	ZHANG Y N, QIAN J L, XU J W, ZHU X Y, LI M Y, XU X X, LIU C X, XUE T, SUN L. Identification of chemosensory genes based on the transcriptomic analysis of six different chemosensory organs in Spodoptera exigua. Frontiers in Physiology, 2018, 9: 432. doi: 10.3389/fphys.2018.00432
[24]	罗智心, 姚静, 张茹, 罗光华, 方继朝. 二化螟二化螟36 d标准化饲养技术与管理体系. 江苏农业科学, 2021, 49(24): 100-109.
	LUO Z X, YAO J, ZHANG R, LUO G H, FANG J C. A 36-days standardized mass rearing technique and management system for rice stem borer, Chilo suppressalis (Lepidoptera: Crambidae), Jiangsu Agricultural Sciences, 2021, 49(24): 100-109. (in Chinese)
[25]	HE F J, LU M X, DU Y Z. Selection and evaluation of reference genes for qRT-PCR analysis of different developmental stages in Chilo suppressalis. Journal of Asia-Pacific Entomology, 2021, 24(4): 1228-1234. doi: 10.1016/j.aspen.2021.08.008
[26]	YANG K, GONG X L, LI G C, HUANG L Q, NING C, WANG C Z. A gustatory receptor tuned to the steroid plant hormone brassinolide in Plutella xylostella (Lepidoptera: Plutellidae). eLife, 2020, 9: e64114. doi: 10.7554/eLife.64114
[27]	ABDEL-LATIEF M. A family of chemoreceptors in Tribolium castaneum (Tenebrionidae: Coleoptera). PLoS ONE, 2007, 2(12): e1319. doi: 10.1371/journal.pone.0001319
[28]	ROBERTSON H M, WANNER K W. The chemoreceptor superfamily in the honey bee, Apis mellifera: Expansion of the odorant, but not gustatory, receptor family. Genome Research, 2006, 16(11): 1395-1403. doi: 10.1101/gr.5057506
[29]	JONES W D, CAYIRLIOGLU P, KADOW I G, VOSSHALL L B. Two chemosensory receptors together mediate carbon dioxide detection in Drosophila. Nature, 2007, 445(7123): 86-90. doi: 10.1038/nature05466
[30]	SATO K, TANAKA K, TOUHARA K. Sugar-regulated cation channel formed by an insect gustatory receptor. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(28): 11680-11685.
[31]	MANG D, SHU M, ENDO H, YOSHIZAWA Y, NAGATA S, KIKUTA S, SATO R. Expression of a sugar clade gustatory receptor, BmGr6, in the oral sensory organs, midgut, and central nervous system of larvae of the silkworm Bombyx mori. Insect Biochemistry and Molecular Biology, 2016, 70: 85-98. doi: 10.1016/j.ibmb.2015.12.008
[32]	LIU X L, SUN S J, HOU W, ZHANG J, YAN Q, DONG S L. Functional characterization of two spliced variants of fructose gustatory receptor in the diamondback moth, Plutella xylostella. Pesticide Biochemistry and Physiology, 2020, 164: 7-13.
[33]	LIU X L, YAN Q, YANG Y L, HOU W, MIAO C L, PENG Y C, DONG S L. A gustatory receptor GR8 tunes specifically to D-fructose in the common cutworm Spodoptera litura. Insects, 2019, 10(9): 272. doi: 10.3390/insects10090272
[34]	JIANG X J, NING C, GUO H, JIA Y Y, HUANG L Q, QU M J, WANG C Z. A gustatory receptor tuned to D-fructose in antennal sensilla chaetica of Helicoverpa armigera. Insect Biochemistry and Molecular Biology, 2015, 60: 39-46. doi: 10.1016/j.ibmb.2015.03.002
[35]	MIYAMOTO T, SLONE J, SONG X, AMREIN H. A fructose receptor functions as a nutrient sensor in the Drosophila brain. Cell, 2012, 151(5): 1113-1125. doi: 10.1016/j.cell.2012.10.024
[36]	MANG D, SHU M, TANAKA S, NAGATA S, TAKADA T, ENDO H, KIKUTA S, TABUNOKI H, IWABUCHI K, SATO R. Expression of the fructose receptor BmGr9 and its involvement in the promotion of feeding, suggested by its co-expression with neuropeptide F1 in Bombyx mori. Insect Biochemistry and Molecular Biology, 2016, 75: 58-69. doi: 10.1016/j.ibmb.2016.06.001
[37]	KIKUTA S, ENDO H, TOMITA N, TAKADA T, MORITA C, ASAOKA K, SATO R. Characterization of a ligand-gated cation channel based on an inositol receptor in the silkworm, Bombyx mori. Insect Biochemistry and Molecular Biology, 2016, 74: 12-20.
[38]	ZHANG H J, ANDERSON A R, TROWELL S C, LUO A R, XIANG Z H, XIA Q Y. Topological and functional characterization of an insect gustatory receptor. PLoS ONE, 2011, 6(8): e24111. doi: 10.1371/journal.pone.0024111
[39]	POPESCU A, COUTON L, ALMAAS T J, ROSPARS J P, WRIGHT G A, MARION-POLL F, ANTON S. Function and central projections of gustatory receptor neurons on the antenna of the noctuid moth Spodoptera littoralis. Journal of Comparative Physiology A, 2013, 199(5): 403-416. doi: 10.1007/s00359-013-0803-0
[40]	贾岩岩. 雌性棉铃虫触角刺形感器和喙管栓锥感器的味觉感受特性[D]. 合肥: 安徽大学, 2016.
	JIA Y Y. Gustatory characteristics of sensilla chaetica in antennae and sensilla styloconica in proboscis of female Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)[D]. Hefei: Anhui University, 2016. (in Chinese)
[41]	MONTELL C. A taste of the Drosophila gustatory receptors. Current Opinion in Neurobiology, 2009, 19(4): 345-353. doi: 10.1016/j.conb.2009.07.001
[42]	AI D, DONG C, YANG B, YU C, WANG G. A fructose receptor gene influences development and feed intake in Helicoverpa armigera. Insect Science, 2022, 29(4): 993-1005. doi: 10.1111/ins.v29.4
[43]	张军飞, 杨雪梅, 孙惠, 张楠, 安传远, 吴帆, 李红亮. 基于循环伏安法的蜜蜂组织传感器对蜂蜜品质的鉴别. 中国食品学报, 2022, 22(6): 307-314.
	ZHANG J F, YANG X M, SUN H, ZHANG N, AN C Y, WU F, LI H L. Identification of honey quality with bee tissue sensor based on cyclic voltammetry. Journal of Chinese Institute of Food Science and Technology, 2022, 22(6): 307-314. (in Chinese)
[44]	AMREIN H, THORNE N. Gustatory perception and behavior in Drosophila melanogaster. Current Biology, 2005, 15(17): R673-R684. doi: 10.1016/j.cub.2005.08.021
[45]	ZHANG G, CAO S, GUO T, WANG H, QI X, REN X, NIU C. Identification and expression profiles of gustatory receptor genes in Bactrocera minax larvae (Diptera: Tephritidae): Role of BminGR59f in larval growth. Insect Science, 2022, 29(5): 1240-1250. doi: 10.1111/ins.v29.5
[46]	DEL CAMPO M L, MILES C I, SCHROEDER F C, MUELLER C, BOOKER R, RENWICK J A. Host recognition by the tobacco hornworm is mediated by a host plant compound. Nature, 2001, 411(6834): 186-189. doi: 10.1038/35075559
[47]	侯文华, 孙龙龙, 马英, 孙惠婉, 张佳佳, 白润娥, 赵新成, 汤清波. 草地贪夜蛾幼虫对四种刺激物质的味觉感受和取食选择. 昆虫学报, 2020, 63(5): 545-557.
	HOU W H, SUN L L, MA Y, SUN H W, ZHANG J J, BAI R E, ZHAO X C, TANG Q B. Gustatory perception and feeding preference of Spodoptera frugiperda (Lepidoptera: Noctuidae) larvae to four stimuli. Acta Entomologica Sinica, 2020, 63(5): 545-557. (in Chinese)
[48]	周东升, 龙九妹. 棉铃虫5龄幼虫对棉酚和印楝素的味觉适应性行为研究. 南方农业学报, 2013, 44(4): 590-593.
	ZHOU D S, LONG J M. Taste adaptability to azadirachtin and gossypol in fifth instar larvae of Helicoverpa armigera. Journal of Southern Agriculture, 2013, 44(4): 590-593. (in Chinese)
[49]	XU W, ZHANG H J, ANDERSON A. A sugar gustatory receptor identified from the foregut of cotton bollworm Helicoverpa armigera. Journal of Chemical Ecology, 2012, 38(12): 1513-1520. doi: 10.1007/s10886-012-0221-8

引物名称 Primer name	引物序列 Primer sequence (5′-3′)	引物用途 Usage of primers
CsGR1-F	ATGTCCTATTATACACATAA	开放阅读框扩增Amplification of ORF
CsGR1-R	TTAATGTTTCCTTTTCTTCT
CsGR2-F	ATGATGATTCCTGATCATTT
CsGR2-R	TCATGAGAGGGCTGCTTGAG
CsGR3-F	ATGTTTGTGGAGAAGCCGGG
CsGR3-R	TTACTGTTCCGAAAGCTGGA
CsGR4-F	GGCTTCGCCCAAGGATCTCC
CsGR4-R	TCAAGCGACAGCGATTACCG
CsGR5-F	ATGAAAACTCAAATCCTGGG
CsGR5-R	TTACTTAATAAATTGCAGCA
CsqGR1-F	AACGCGATAGCACCTTTGGA	实时荧光定量PCR RT-qPCR
CsqGR1-R	AGAAGGTCTGGTGATCGGGA
CsqGR2-F	TGCCGTCATGCCTATCACAA
CsqGR2-R	ACGCTCGTATCCAACGAACA
CsqGR3-F	CGGGGTTCTGCTGTTACTCC
CsqGR3-R	CAGACCACACCACAACACCT
CsqGR4-F	CCAAGTTCACGGGGATACGG
CsqGR4-R	GAAATGGTGGCGGCAAGAAA
CsqGR5-F	GTCAGCAGCGTTGGTCAAAT
CsqGR5-R	GAGCGATACTGACCCTGACG
EF1-F	AAAATGGACTCGACTGAACCCC
EF1-R	TCTCCGTGCCAACCAGAAATA

基因 Gene	登录号 Accession number	开放阅读框长度 ORF length (bp)	氨基酸数量 Number of AA	跨膜结构域数量 Number of TMD	最佳比对结果BLAST best hit
基因 Gene	登录号 Accession number	开放阅读框长度 ORF length (bp)	氨基酸数量 Number of AA	跨膜结构域数量 Number of TMD	物种 Species	登录号 Accession number	E值 E-value	相似度 Similarity (%)
CsupGR1	OQ834263	1428	475	8	亚洲玉米螟O. furnacalis	XP_028179334.1	0	80.62
CsupGR2	OQ834264	1302	433	7	粉纹夜蛾T. ni	XP_026735279.1	0	86.95
CsupGR3	OQ834265	1317	438	8	棉红铃虫P. gossypiella	XP_049878938.1	3.00E-63	31.45
CsupGR4	OQ834266	1122	373	7	黄野螟H. vitessoides	UVB79175.1	7.00E-174	73.70
CsupGR5	OQ834267	1269	422	7	亚洲玉米螟O. furnacalis	XP_028178841.1	0	66.91

		CsupGR1	CsupGR2	CsupGR3	CsupGR4	CsupGR5
分子式Molecular formula		C₂₄₉₁H₃₉₀₈N₆₁₀O₆₇₆S₂₆	C₂₂₈₇H₃₅₄₇N₅₆₇O₆₁₈S₂₄	C₂₂₃₉H₃₅₉₆N₅₆₆O₆₀₈S₂₃	C₁₈₉₉H₂₉₆₈N₄₇₄O₅₁₃S₂₀	C₂₂₇₃H₃₅₅₈N₅₆₀O₅₈₉S₂₄
分子质量Molecular weight (Da)		54051.67	49643.19	48909.99	41288.52	48924.09
等电点Theoretical pI		8.75	7.51	9.28	7.09	9.31
氨基酸含量 Amino acid content	Maximum	Leu 11.2%	Leu 11.8%	Leu 15.8%	Leu 12.3%	Leu 14.7%
	Minimum	Trp 1.1%	Trp 1.4%	Trp 0.9%	Asp 0.8%	Trp 1.2%
	Nonentity	Pyl, Sec	Pyl, Sec	Pyl, Sec	Pyl, Sec	Pyl, Sec
正电荷残基Positively charged residues		40	40	43	21	36
负电荷残基Negatively charged residues		34	39	29	21	23
吸收率Absorptivity		1.205	1.370	1.041	0.884	1.294
不稳定指数Instability index		33.75	33.17	34.53	34.79	38.05
脂肪族指数Aliphatic index		109.18	110.35	120.59	114.48	119.36
疏水性系数的总平均值 Grand average of hydrophobicity coefficient		0.327	0.337	0.522	0.656	0.489