Association between polymorphism of vasoactive intestinal peptide type Ⅰ receptor gene and carcass traits in meat quails

doi:10.3969/j.issn.1004-1524.2022.11.07

Acta Agriculturae Zhejiangensis ›› 2022, Vol. 34 ›› Issue (11): 2379-2385.DOI: 10.3969/j.issn.1004-1524.2022.11.07

• Animal Science • Previous Articles Next Articles

Association between polymorphism of vasoactive intestinal peptide type Ⅰ receptor gene and carcass traits in meat quails

WANG Xinle¹(), BAI Junyan¹^,²^,^*(), LI Jingyun¹, LEI Ying¹^,², LI Gan¹^,², PANG Youzhi¹^,², DONG Zhihao¹, CHEN Yu¹, FAN Hongdeng¹, WANG Longwei¹, CHEN Mengke¹, ZENG Fanlin¹, AN Xiaokai¹, BAI Yongzhen¹

1. College of Animal Science and Technology, Henan University of Science and Technology, Luoyang 471023, Henan, China
2. Luoyang Key Laboratory of Animal Genetics and Breeding, Luoyang 471023, Henan, China

Received:2021-08-06 Online:2022-11-25 Published:2022-11-29
Contact: BAI Junyan

Abstract

Abstract:

In order to investigate the relationship between the polymorphism of type 1 vasoactive intestinal peptide receptor (VIPR-1) gene and carcass traits of meat quail, the polymorphism of VIPR-1 gene was detected by PCR-RFLP method in two populations, including French giant meat quail and Savimaito meat quail, and the correlation between the polymorphism of VIPR-1 gene and carcass traits of meat quail was analyzed. The results showed that GG, GT and TT genotypes were detected at the BsrDⅠ locus of exon 4-5 of VIPR-1 gene in French giant meat quail and Savimaito meat quail, the GG, GT and TT genotype frequencies of French giant meat quail and Savimaito meat quail were 0.408, 0.490, 0.102; 0.222, 0.444, 0.334, respectively. AA, AG and GG genotypes were detected at the HpyCH4Ⅳ locus of exon 6-7 of VIPR-1 gene in French giant meat quail and Savimaito meat quail, the AA, AG and GG genotype frequencies of French giant meat quail and Savimaito meat quail were 0.020, 0.020, 0.960; 0.056,0.111,0.833, respectively. The results of association analysis showed that the BsrDⅠ locus in exon 4-5 of VIPR-1 gene was significantly associated with the liver weight of French giant meat quail (P<0.05). The HpyCH4Ⅳ locus in exon 6-7 of VIPR-1 gene was significantly associated with body weight, slaughter weight and liver weight of French giant meat quail (P<0.05), and with the body weight, slaughter weight, eviscerated weight, heart weight, chest muscle weight, leg muscle weight, liver rate and chest muscle rate of Savimaito meat quail (P<0.05). The VIPR-1 gene can be used as a candidate gene for marker-assisted selection of carcass traits in meat quail.

Key words: meat quail, vasoactive intestinal peptide receptor-1 (VIPR-1) gene, polymorphism, carcass trait, association analysis

CLC Number:

S837

WANG Xinle, BAI Junyan, LI Jingyun, LEI Ying, LI Gan, PANG Youzhi, DONG Zhihao, CHEN Yu, FAN Hongdeng, WANG Longwei, CHEN Mengke, ZENG Fanlin, AN Xiaokai, BAI Yongzhen. Association between polymorphism of vasoactive intestinal peptide type Ⅰ receptor gene and carcass traits in meat quails[J]. Acta Agriculturae Zhejiangensis, 2022, 34(11): 2379-2385.

Figures/Tables 7

References 20

[1]	KANSAKU N, SHIMADA K, OHKUBO T, et al. Molecular cloning of chicken vasoactive intestinal polypeptide receptor complementary DNA, tissue distribution and chromosomal localization[J]. Biology of Reproduction, 2001, 64(5): 1575-1581. PMID
[2]	CARDOSO J C R, VIEIRA F A, GOMES A S, et al. PACAP, VIP and their receptors in the Metazoa: Insights about the origin and evolution of the ligand-receptor pair[J]. Peptides, 2007, 28(9): 1902-1919. PMID
[3]	李雨顺. 绿壳蛋鸡和芦花鸡产蛋性能测定及其相关基因(ER-α和VIPR-1)多态性研究[D]. 雅安: 四川农业大学, 2015.
	LI Y S. The laying performance of green shell layer and barred rock layer were measured and the research of relavent (ER and VIP-1) gene polymorphism[D]. Ya’an: Sichuan Agricultural University, 2015. (in Chinese with English abstract)
[4]	朱志明, 陈红萍, 杨武, 等. 河田鸡VIPR-1基因外显子2HpaⅡ位点多态性与产蛋性状的关联分析[J]. 畜牧与兽医, 2015, 47(47): 46-48.
	ZHU Z M, CHEN H P, YANG W, et al. Association analysis of polymorphism of VIPR-1 gene exon-2 Hpa Ⅱ with laying traits in Hetian chickens[J]. Animal Husbandry & Veterinary Medicine, 2015, 47(3): 46-48. (in Chinese)
[5]	周敏, 沈栩, 李莹, 等. 清远麻鸡VIPR-1基因C1704887T、C1715301T位点与繁殖性状的关联性分析[J]. 江西农业大学学报, 2011, 33(33): 117-122.
	ZHOU M, SHEN X, LI Y, et al. Analysis on association of C1704887T and C1715301T sites of VIPR-1 gene with reproduction traits in Qingyuan partridge chicken[J]. Acta Agriculturae Universitatis Jiangxiensis, 2011, 33(1): 117-122. (in Chinese with English abstract)
[6]	PU Y J, WU Y, XU X J, et al. Association of VIPR-1 gene polymorphisms and haplotypes with egg production in laying quails[J]. Journal of Zhejiang University-SCIENCE B, 2016, 17(8): 591-596. DOI URL
[7]	潘爱銮, 皮劲松, 杜金平, 等. 中国鹌鹑遗传研究进展[J]. 湖北农业科学, 2012, 51(51): 5263-5266.
	PAN A L, PI J S, DU J P, et al. Progress of quail genetic research in China[J]. Hubei Agricultural Sciences, 2012, 51(23): 5263-5266. (in Chinese with English abstract)
[8]	蒲跃进. 蛋用鹌鹑产蛋相关基因克隆、表达及其与性状的关联性研究[D]. 武汉: 华中农业大学, 2016.
	PU Y J. Molecular cloning and expression of genes related with laying quail reproduction and their associations with egg performances[D]. Wuhan: Huazhong Agricultural University, 2016. (in Chinese with English abstract)
[9]	辛清武, 缪中纬, 郑嫩珠, 等. 黑番鸭VIPR-1基因第12内含子SNP与就巢性状关联分析[J]. 家畜生态学报, 2013, 34(34): 16-19.
	XIN Q W, MIAO Z W, ZHENG N Z, et al. Correlation analysis between nucleotide polymorphism of the VIPR-112 intron and broodiness in black Muscovy duck[J]. Journal of Domestic Animal Ecology, 2013, 34(10): 16-19. (in Chinese with English abstract)
[10]	李德娟, 许盈盈, 代敏敏, 等. 太行鸡VIPR-1基因内含子1多态性及其与早期产蛋性状相关性研究[J]. 黑龙江畜牧兽医, 2020(9): 9-14,166.
	LI D J, XU Y Y, DAI M M, et al. Research on intron 1 polymorphism of VIPR-1 gene and its association with early laying traits in Taihang chicken[J]. Heilongjiang Animal Science and Veterinary Medicine, 2020(9): 9-14,166. (in Chinese with English abstract)
[11]	ZHOU M, LEI M, RAO Y, et al. Polymorphisms of vasoactive intestinal peptide receptor-1 gene and their genetic effects on broodiness in chickens[J]. Poultry Science, 2008, 87(5): 893-903. DOI PMID
[12]	ZHAO L F, YU Z P, ZHAO B Q. Mechanism of VIPR1 gene regulating human lung adenocarcinoma H1299 cells[J]. Medical Oncology, 2019, 36(11): 1-7. DOI URL
[13]	VUCKOVIC A, RONDELET B, BRION J P, et al. Expression of vasoactive intestinal peptide and related receptors in overcirculation-induced pulmonary hypertension in piglets[J]. Pediatric Research, 2009, 66(4): 395-399. DOI PMID
[14]	XU H P, ZENG H, LUO C L, et al. Genetic effects of polymorphisms in candidate genes and the QTL region on chicken age at first egg[J]. BMC Genetics, 2011, 12: 33. DOI PMID
[15]	周敏, 梁菲菲, 饶友生, 等. VIPR-1基因12个多态位点与鸡早期产蛋性状的相关性[J]. 畜牧兽医学报, 2008, 39(39): 1147-1152.
	ZHOU M, LIANG F F, RAO Y S, et al. Association of twelve polymorphisms of the VIPR-1 gene with chicken early egg production traits[J]. Chinese Journal of Animal and Veterinary Sciences, 2008, 39(9): 1147-1152. (in Chinese with English abstract)
[16]	周敏, 刘满清, 徐海平, 等. 血管活性肠肽Ⅰ型受体基因多态位点与鸡冠高度和体重的相关性[J]. 中国家禽, 2009, 31(31): 17-20.
	ZHOU M, LIU M Q, XU H P, et al. Association of polymorphisms of VIPR-1 gene with chicken comb height and body weight[J]. China Poultry, 2009, 31(20): 17-20. (in Chinese with English abstract)
[17]	洪军, 王克华, 李东锋, 等. VIPR-1基因SNP位点与如皋黄鸡产蛋性能和蛋品质的关联分析[J]. 中国畜牧杂志, 2011, 47(47): 1-5.
	HONG J, WANG K H, LI D F, et al. Association of SNP of VIPR-1 gene with egg production and egg quality traits in Rugao-yellow chicken[J]. Chinese Journal of Animal Science, 2011, 47(23): 1-5. (in Chinese with English abstract)
[18]	周敏, 何丹林, 刘满清, 等. 清远麻鸡VIPR-1基因多态性及与生长性状的关联分析[J]. 养禽与禽病防治, 2009(7): 2-4.
	ZHOU M, HE D L, LIU M Q, et al. Polymorphism of VIPR-1 gene and its association with growth traits in Qingyuan partridge chickens[J]. Poultry Husbandry and Disease Control, 2009(7): 2-4. (in Chinese)
[19]	周敏, 谢袖娟, 和俊, 等. VIPR-1基因5'调控区多态位点与鸡开产性状的相关性[C]// 安全优质的家禽生产: 第十五次全国家禽学术讨论会论文集. 广州, 2011: 219-223.
[20]	董智豪, 陈宇, 黄高想, 等. 蛋用鹌鹑的VIPR-1基因的多态性与早期生长性状的关联分析[J]. 浙江农业学报, 2021, 33(8):1393-1401. DOI
	DONG Z H, CHEN Y, HUANG G X, et al. Association analysis of VIPR-1 gene polymorphism and early growth traits in egg quail[J]. Acta Agriculturae Zhejiangensis, 2021, 33(8) : 1393-1401. (in Chinese with English abstract)

名称 Name	引物序列 Primer sequence(5'-3')	长度 Size/bp	退火温度 Annealing temperature/℃	位置 Position
BsrDⅠ	F:GCGTTCTATGGCACAGTTA R:AAAGCAATGTTCGGGTTCT	925	54	外显子4-5 Extron 4-5
HpyCH4Ⅳ	F:GCTGCTGGTGGAAGGGTTA R:CCGTCCAAGCAGTGATGAA	1 096	58.7	外显子6-7 Extron 6-7

名称 Name	引物序列 Primer sequence(5'-3')	长度 Size/bp	退火温度 Annealing temperature/℃	位置 Position
BsrDⅠ	F:GCGTTCTATGGCACAGTTA R:AAAGCAATGTTCGGGTTCT	925	54	外显子4-5 Extron 4-5
HpyCH4Ⅳ	F:GCTGCTGGTGGAAGGGTTA R:CCGTCCAAGCAGTGATGAA	1 096	58.7	外显子6-7 Extron 6-7

群体 Group	基因型频率 Genotype frequency			等位基因频率 Allele frequency		杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
群体 Group	GG	GT	TT	G	T	杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
莎维麦脱肉用鹌鹑	0.222	0.444	0.334	0.444	0.556	0.494	1.976	0.372	0.360(0.835)
Savimaito meat quail
法国巨型肉用鹌鹑	0.408	0.490	0.102	0.653	0.347	0.453	1.829	0.350	0.321(0.852)
French giant meat quail

群体 Group	基因型频率 Genotype frequency			等位基因频率 Allele frequency		杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
群体 Group	GG	GT	TT	G	T	杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
莎维麦脱肉用鹌鹑	0.222	0.444	0.334	0.444	0.556	0.494	1.976	0.372	0.360(0.835)
Savimaito meat quail
法国巨型肉用鹌鹑	0.408	0.490	0.102	0.653	0.347	0.453	1.829	0.350	0.321(0.852)
French giant meat quail

群体 Group	基因型频率 Genotype frequency			等位基因频率 Allele frequency		杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
群体 Group	AA	AG	GG	A	G	杂合度 Hybridization (He)	有效等位基因数 Number of effective alleles(Ne)	多态信息含量 Polymorphic information content(PIC)	哈代温伯格平衡检验 Hardy-Weinberg equilibrium[X²(P)]
莎维麦脱肉用鹌鹑	0.056	0.111	0.833	0.111	0.889	0.198	1.246	0.178	6.891(0.032)
Savimaito meat quail
法国巨型肉用鹌鹑	0.020	0.020	0.960	0.031	0.969	0.059	1.063	0.058	21.095(0.000)
French giant meat quail

Association between polymorphism of vasoactive intestinal peptide type Ⅰ receptor gene and carcass traits in meat quails

RichHTML

PDF (PC)

Knowledge

Abstract

Cite this article

share this article

Figures/Tables 7

References 20

Related Articles 15

Recommended Articles

Metrics

Comments

胴体性状 Carcass traits	莎维麦脱肉用鹌鹑Savimaito meat quail			法国巨型肉用鹌鹑French giant meat quail
胴体性状 Carcass traits	GG	GT	TT	GG	GT	TT
体重Body weight/g	138.550±5.688 a	130.450±7.016 a	140.433±7.426 a	147.405±2.694 a	144.396±3.812 a	149.340±8.743 a
屠宰重Slaughter weight/g	130.838±5.360 a	123.100±6.732 a	133.117±7.264 a	139.930±2.674 a	137.033±3.724 a	142.000±8.668 a
全净膛重Eviscerated weight/g	97.588±4.350 a	88.875±5.659 a	96.792±5.807 a	101.680±1.868 a	100.392±3.093 a	103.380±7.292 a
心重Heart weight/g	0.988±0.083 a	0.969±0.092 a	1.133±0.089 a	1.120±0.071 a	1.221±0.062 a	1.060±0.081 a
肝重Liver weight/g	3.463±0.185 a	3.600±0.138 a	3.550±0.173 a	3.760±0.139 ab	3.583±0.116 b	4.120±0.146 a
胸肌重Chest muscle weight/g	30.150±1.150 a	27.481±2.147 a	30.683±2.298 a	32.510±0.735 a	32.017±1.177a	33.060±2.975 a
腿肌重Leg muscle weight/g	6.363±0.319 a	6.131±0.381 a	6.650±0.397 a	7.440±0.294 a	7.171±0.243 a	7.760±0.480 a
屠宰率Slaughter rate/%	94.442±0.341 a	94.280±0.273 a	94.680±0.312 a	94.902±0.206 a	94.863±0.187 a	95.040±0.533 a
全净膛率Eviscerated rate/%	70.371±0.545 a	67.491±0.939 a	68.634±0.927 a	68.998±0.448 a	69.345±0.508 a	69.143±2.637 a
心比率Heart rate/%	1.011±0.071 a	1.073±0.060 a	1.213±0.142 a	1.100±0.065 a	1.225±0.061 a	1.055±0.126 a
肝比率Liver rate/%	3.579±0.196 a	4.249±0.257 a	3.757±0.184 a	3.706±0.133 a	3.640±0.158 a	4.057±0.291 a
胸肌率Chest muscle rate/%	31.022±0.759 a	30.555±0.745 a	31.402±0.753 a	31.961±0.388 a	31.783±0.447 a	31.925±1.617 a
腿肌率Leg muscle rate/%	13.027±0.198 a	13.947±0.437 a	13.857±0.440 a	14.564±0.443 a	14.372±0.359 a	15.254±1.162 a

胴体性状 Carcass traits	莎维麦脱肉用鹌鹑Savimaito meat quail			法国巨型肉用鹌鹑French giant meat quail
胴体性状 Carcass traits	AA	AG	GG	AA	AG	GG
体重Body weight/g	163.350±4.750 a	103.600±6.989 c	137.990±4.185 b	143.900±2.002 b	169.700±2.302 a	145.674±2.346 b
屠宰重Slaughter weight/g	155.550±6.650 a	97.400±6.768 c	130.433±4.030 b	134.600±2.061 b	163.600±2.261 a	138.281±2.294 b
全净膛重Eviscerated weight/g	114.850±7.250 a	67.100±5.006 c	95.537±3.317 b	98.900±1.011 a	120.900±1.811 a	100.853±1.839 a
心重Heart weight/g	1.200±0.000 a	0.675±0.085 c	1.063±0.059 b	1.100±0.043 a	1.400±0.043 a	1.160±0.044 a
肝重Liver weight/g	3.700±0.800 a	3.450±0.343 a	3.557±0.095 a	3.700±0.084 b	5.100±0.084 a	3.681±0.082 b
胸肌重Chest muscle weight/g	38.900±2.800 a	19.000±2.107 c	29.843±1.209 b	33.200±0.670 a	36.600±0.700 a	32.215±0.724 a
腿肌重Leg muscle weight/g	7.950±0.450 a	5.050±0.119 c	6.423±0.241 b	7.700±0.185 a	6.900±0.175 a	7.343±0.182 a
屠宰率Slaughter rate/%	95.187±1.303 a	93.970±0.371 a	94.464±0.190 a	93.537±0.142 a	96.405±0.132 a	94.894±0.131 a
全净膛率Eviscerated rate/%	70.240±2.396 a	64.677±0.759 a	68.908±0.586 a	68.728±0.401 a	71.243±0.391 a	69.148±0.405 a
心比率Heart rate/%	1.049±0.066 a	1.023±0.138 a	1.121±0.066 a	1.112±0.052 a	1.158±0.042 a	1.158±0.044 a
肝比率Liver rate/%	3.279±0.904 c	5.179±0.459 a	3.814±0.126 b	3.741±0.109 a	4.218±0.099 a	3.698±0.102 a
胸肌率Chest muscle rate/%	33.851±0.301 a	28.066±1.203 b	31.131±0.451 ab	33.569±0.326 a	30.273±0.306 a	31.868±0.315 a
腿肌率Leg muscle rate/%	13.850±0.091 a	15.293±1.126 a	13.493±0.244 a	15.571±0.284 a	11.414±0.274 a	14.585±0.277 a

[1]	LIU Pengcheng, ZHANG Ji, QIU Ganyuan, GONG Yu, LI Xuesong, LI Wei, ZHANG Yiyu, LIU Ruoyu. Single nucleotide polymorphism screening and bioinformatics analysis of TBC1D7 gene in Guanling cattle [J]. Acta Agriculturae Zhejiangensis, 2022, 34(7): 1402-1411.
[2]	XU Jingen, JIN Erhui, WANG Chonglong, GU Youfang, LI Qinggang. Polymorphism and bioinformatics analysis of pig CAST gene [J]. Acta Agriculturae Zhejiangensis, 2022, 34(1): 17-23.
[3]	DONG Zhihao, CHEN Yu, HUANG Gaoxiang, BAI Junyan, LI Jingyun, ZHAO Shujuan, LEI Ying, WANG Xinle, HU Qihang, FAN Zhengyu. Association analysis of VIPR-1 gene polymorphism and early growth traits in egg quail [J]. Acta Agriculturae Zhejiangensis, 2021, 33(8): 1393-1401.
[4]	LIU Xiuhui, ZOU Guihua, ZHAI Guowei, LIU Heqin, ZHENG Xueqiang, CHEN Heyun. Association analysis between SSR markers and germination-related traits in sorghum germplasm resources [J]. Acta Agriculturae Zhejiangensis, 2021, 33(6): 965-973.
[5]	LIN Mengjie, WEN Huiping, XIAO Jianzhong, ZHENG Qiang. Construction of DNA fingerprint for 48 broccoli cultivars [J]. Acta Agriculturae Zhejiangensis, 2021, 33(12): 2304-2312.
[6]	TAO Peng, YUE Zhichen, ZHAO Yanting, LEI Juanli, LI Biyuan. Variation of acceptor splicing site of BrSPS1Fb of Chinese cabbage and its effect on splicing [J]. Acta Agriculturae Zhejiangensis, 2021, 33(11): 2068-2074.
[7]	LIAO Chaomei, YOU Minfang, TAN Guanghui, LI Jiezhang, ZHANG Yiyu, QIN Yuanyu, LIU Ruoyu. Effect of variation of PRKCB gene intron 14 in Sansui ducks on eggshell quality [J]. , 2020, 32(9): 1574-1580.
[8]	LIU Xinyu, TIAN Jie. Analysis of simple sequence repeats in transcriptome of garlic (Allium sativum L.) and development of molecular markers [J]. , 2020, 32(9): 1615-1625.
[9]	BAI Junyan, LU Junhao, FU Xueyan, WU Xiaohong, YANG Youbing, LEI Ying, PANG Youzhi, LU Xiaoning, GONG Huirong, HU Luxing, LIU Hongtao, FAN Hongdeng, CAO Heng, SHI Kunpeng, CHEN Mengke, MA Yongkang. Analysis of correlation between polymorphism of IGF-1R gene and body size traits in egg quail [J]. , 2020, 32(3): 398-405.
[10]	HU Qianwen, XU Yanhao, WANG Rong, ZHANG Wenying, HUA Wei, LYU Chao. Association analysis of four spike traits in barley [J]. , 2020, 32(11): 1941-1953.
[11]	BAI Junyan, CAO Heng, WANG Xu, YANG Youbing, FAN Hongdeng, FU Xueyan, SHI Kunpeng, DONG Zhihao, LU Xiaoning, LI Xinyue, HAO Weiguang, LI Ziheng, ZHENG Feiyang. Association between the polymorphism of PvuⅡ locus of GH gene and growth traits in sheep [J]. , 2019, 31(9): 1416-1422.
[12]	ZHANG Juan, MU Tong, ZHAO Ping, CHEN Jiaping, FENG Xiaofang, GUO Peng, WU Zewen, LIU Liyuan, JIANG Qiufei, GU Yaling. Polymorphism of ELOVL5 gene in Jingyuan chicken [J]. , 2019, 31(2): 200-206.
[13]	YANG Youbing, BIAN Junping, WU Yan, LOU Ran, LI Shihao. Association analysis between polymorphism in FTO gene and meat quality traits in pigs [J]. , 2019, 31(12): 1971-1978.
[14]	LU Xin, ZHOU Jinghang, YANG Chaoyun, ZHANG Menghua, YE Lianmeng, LI Shuzhen, HUANG Xixia, MA Yun, WANG Xingping, SHI Yuangang. Functional annotation of candidate genes for milk production and reproductive traits in Xinjiang Brown cattle [J]. , 2019, 31(12): 1987-1995.
[15]	BAI Junyan, FAN Hongdeng, CAO Heng, PANG Youzhi, SONG Xiaojie, JIANG Mengjuan, FU Xueyan, LU Xiaoning, YANG Shuai, LI Xinyue, HAO Weiguang, LI Ziheng, ZHENG Feiyang. Association analysis between polymorphism of ESR1 gene and egg quality in quail [J]. , 2019, 31(10): 1608-1614.