Research Progress on the Pathogen Control of Citrus Huanglongbing

doi:10.16420/j.issn.0513-353x.2024-0512

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (8): 2059-2080.doi: 10.16420/j.issn.0513-353x.2024-0512

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Research Progress on the Pathogen Control of Citrus Huanglongbing

XU Shugui, and ZHENG Zheng^*(), DENG Xiaoling^*()

Citrus Huanglongbing Research Laboratory， South China Agricultural University， Guangzhou 510642， China

Received:2025-04-21 Revised:2025-06-25 Online:2025-08-19 Published:2025-08-19
Contact: and ZHENG Zheng, DENG Xiaoling

Abstract

Abstract:

This review systematically summarizes the research advances in citrus Huanglongbing（HLB）management from multiple perspectives，including thermotherapy，antibiotics，antimicrobial agents，plant defense inducers，resistant cultivar cultivation，and nutritional regulation. A comparison of the efficacy of different approaches may help integrate their respective advantages，offering insights for future HLB control research and supporting the development of integrated management systems.

Key words: citrus Huanglongbing, antibiotic treatment, antimicrobial drugs, green control

XU Shugui, and ZHENG Zheng, DENG Xiaoling. Research Progress on the Pathogen Control of Citrus Huanglongbing[J]. Acta Horticulturae Sinica, 2025, 52(8): 2059-2080.

Figures/Tables 3

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抗菌级别 Antibacterial activity	抗生素 Antibiotics	嫁接180 d后Ct值 Ct value after 180 days of grafting			接穗成活率/% Scion Survival	传毒率/%
抗菌级别 Antibacterial activity	抗生素 Antibiotics	对照 Control	接穗 Scion	砧木 Rootstocks	接穗成活率/% Scion Survival	传毒率/%
高效 Highly efficient	氨苄西林、羧苄青霉素钠、青霉素钾、头孢氨苄、利福平、磺胺间二甲氧嘧啶钠 Ampicillin sodium，Carbenicillin disodium，Penicillin G potassium，Cefalexin，Rifampicin，Sulfadimethoxine sodium	25.9 ± 2.7 a	38.3 ± 1.7 a	37.0 ± 2.2 a	80.9 ± 8.0 a	16.9 ± 16.2 c
部分有效 Partly effective	氯霉素、井冈霉素、中生菌素、磺胺甲恶唑、环丝氨酸、卡那霉素、壮观霉素、潮霉素、利福昔明、盐酸环丙沙星、磺胺噻唑钠、利福霉素钠 Chloramphenicol，Validoxylamine A，Sulfamethoxazole，Cycloserine，Kanamycin sulfate，Spectinomycin dihydrochloride pentadrate，Zhongshengmycin，Hygromycin B，Rifaximin，Ciprofloxacin hydrochloride，Sulfathiazole sodium，Rifamycin sodium	26.1 ± 1.9 a	32.3 ± 1.4 b	33.6 ± 2.7 b	81.7 ± 15.5 a	39.2 ± 12.0 b
无效 No effective	硫酸丁胺卡那霉素、硫酸多黏菌素B、春雷霉素、多黏菌素E甲磺酸钠、硫酸链霉素、盐酸万古霉素、西诺沙星、硫酸双生霉素、硫酸新霉素、妥布霉素、盐酸林可霉素 Amikacin sulfate，Polymixin B sulfate，Kasugamycin hydrochloride，Colistinmethanesulfonate sodium，Streptomycin sulfate，Vancomycin hydrochloride，Cinoxacin，Gentamicin sulfate，Neomycin hydrate trisulfate，Tobramycin，Lincomycinhydrocloride	25.1 ± 1.9 a	27.2 ± 3.0 c	26.4 ± 2.4 c	89.8 ± 9.4 a	80.5 ± 9.8 a

抗菌级别 Antibacterial activity	抗生素 Antibiotics	嫁接180 d后Ct值 Ct value after 180 days of grafting			接穗成活率/% Scion Survival	传毒率/%
抗菌级别 Antibacterial activity	抗生素 Antibiotics	对照 Control	接穗 Scion	砧木 Rootstocks	接穗成活率/% Scion Survival	传毒率/%
高效 Highly efficient	氨苄西林、羧苄青霉素钠、青霉素钾、头孢氨苄、利福平、磺胺间二甲氧嘧啶钠 Ampicillin sodium，Carbenicillin disodium，Penicillin G potassium，Cefalexin，Rifampicin，Sulfadimethoxine sodium	25.9 ± 2.7 a	38.3 ± 1.7 a	37.0 ± 2.2 a	80.9 ± 8.0 a	16.9 ± 16.2 c
部分有效 Partly effective	氯霉素、井冈霉素、中生菌素、磺胺甲恶唑、环丝氨酸、卡那霉素、壮观霉素、潮霉素、利福昔明、盐酸环丙沙星、磺胺噻唑钠、利福霉素钠 Chloramphenicol，Validoxylamine A，Sulfamethoxazole，Cycloserine，Kanamycin sulfate，Spectinomycin dihydrochloride pentadrate，Zhongshengmycin，Hygromycin B，Rifaximin，Ciprofloxacin hydrochloride，Sulfathiazole sodium，Rifamycin sodium	26.1 ± 1.9 a	32.3 ± 1.4 b	33.6 ± 2.7 b	81.7 ± 15.5 a	39.2 ± 12.0 b
无效 No effective	硫酸丁胺卡那霉素、硫酸多黏菌素B、春雷霉素、多黏菌素E甲磺酸钠、硫酸链霉素、盐酸万古霉素、西诺沙星、硫酸双生霉素、硫酸新霉素、妥布霉素、盐酸林可霉素 Amikacin sulfate，Polymixin B sulfate，Kasugamycin hydrochloride，Colistinmethanesulfonate sodium，Streptomycin sulfate，Vancomycin hydrochloride，Cinoxacin，Gentamicin sulfate，Neomycin hydrate trisulfate，Tobramycin，Lincomycinhydrocloride	25.1 ± 1.9 a	27.2 ± 3.0 c	26.4 ± 2.4 c	89.8 ± 9.4 a	80.5 ± 9.8 a

分类 Category		靶标 Target	作用 Function	小分子化合物 Drug	试验结果 Efficacy	参考文献 Reference
转运系统 Transporter system		SecA	SecA是细菌独有的分泌系统II的组成成分，可以构成SecA-ATP酶，供给能量 SecA is the unique bacterial component of secretion system II that forms the SecA-ATPase to supply energy	4-（4-碘-2-甲基苯氧乙酰基）苯磺酰胺，2-（苯并咪唑-2-硫代）-N-[2-（环丙胺甲酰甲硫基）苯并噻唑-6-基]乙酰胺，2-（5-氨基-1,2,4-三唑-3-硫代）-1-（2,5-二氯苯基-2,4-二甲基环戊二烯基）乙酮， 4-（4-氯-2-甲基苯氧乙酰基）苯磺酰胺 4-[(4-Iodo-2-methylphenoxy)acetyl]benzenesulfonamide，2-(1H-Benzimidazol-2-ylthio)-N-[2-(cyclopropylcarbamoylmethylthio)benzothiazol-6-yl]acetamide，2-(5-Amino-1,2,4-triazol-3-ylthio)-1-(2,5-dichlorophenyl)-2,4-dimethylcyclopent-1-en-1-one， 4-(4-Chloro-2-methylphenoxyacetyl)benzenesulfonamide	采用虚拟筛选和分子对接的方法，发现这4种化合物在20 μmol · L^-1对SecA抑制率大于65%，它们能与SecA的Phe84位点形成稳定的π-π结构，使SecA无法和ATP结合而失活，达到抑菌的目的 Using virtual screening and molecular docking methods，it was found that these four compounds inhibited SecA by more than 65% at 20 μmol · L^-1，and they could form a stable π-π structure with the Phe84 site of SecA，which inactivated SecA by preventing it from combining with ATP，and thus achieved bacterial inhibition	Akula et al.,2011
		SecA		2,4-二氟苄基-4-氟苯基四氢吡唑并吡啶甲酰胺，4-氯苯基-2-乙氧基苯基二氢吡唑并吡啶甲酰胺 (2,4-Difluorobenzyl)(4-fluorophenyl)tetrahydropyrazolopyridinecarboxamide，(4-Chlorophenyl)(2-ethoxyphenyl)dihydropyrazolopyridinecarboxamide	分子对接的结果表明两种化合物可以和SecA的多个残基相互形成稳定的氢键和金属配位互作，浓度为1 mg · mL^-1的P684-2850、P684-3808对L. crescens BT-1的抑制率达99.6%、56.1% The results of molecular docking showed that they could form stable hydrogen bonds and metal-ligand interactions with several residues of SecA，and the inhibition rates of P684-2850 and P684-3808 at a concentration of 1 mg · mL^-1 against L. crescens BT-1 were 99.6% and 56.1% respectively	Zhang et al.,2021c
		ZunA1	黄龙病菌的ABC转运系统的组成部分，负责吸收外界Zn离子 ABC transporter system of Ca. Lasiaticus is responsible for the uptake of external Zn ions	ZINC15670529，ZINC92774705， ZINC06510089，ZINC79841324，ZINC69594834	从ZINC筛选出的5个候选小分子能够与CLas的ZnuA1活性位点残基形成氢键，结合自由能较低，形成稳定的复合物，阻断CLas对Zn²⁺摄取，可以进一步开发为CLas的抑制剂 The five candidate small molecules screened from the ZINC database，which can be further developed as inhibitors of CLas，can form hydrogen bonds with the active site residues of CLas ZnuA1，exhibiting low binding free energy and forming stable complexes，thereby blocking Zn²⁺ uptake by CLas	Saini et al.,2021
调节因子 Regulatory factors		LdtR	LdtR可以感受并调节细胞渗透压、影响细菌细胞壁的合成 LdtR senses and regulates cellular osmotic pressure and influences bacterial cell wall synthesis	根皮素，苯溴马隆，己烷雌酚 Phloretin，Benzbromarone，Hexestrol	根皮素、苯溴马隆、己烷雌酚可以结合CLas的LdtR蛋白形成稳定复合物，干扰细胞的渗透压平衡，利用根瘤菌和L. crescens进行体外实验也证明这3种化合物可以使病菌的细胞吸涨破裂 Phloretin，benzbromarone，and hexestrol can bind to the LdtR protein of CLas to form stable complexes that disrupt cellular osmotic balance. In vitro experiments using Rhizobium and L. crescens have demonstrated that these three compounds induce bacterial cell turgor and subsequent lysis	Pagliai et al.,2014
调节因子 Regulatory factors		PrbP	PrbP和RNA聚合酶β亚基互作调节基因表达，可能与细菌的致病力、细胞活性、抗逆性相关 PrbP and RNA polymerase β subunit interact to regulate gene expression，which may be related to bacterial pathogenicity，cellular activity，and resistance	托芬那酸 Tolfenamic acid	托芬那酸可以干扰PrbP和RNA聚合酶的互作，托芬那酸灌根或喷施感染黄龙病柑橘植株，在2个月后病株恢复长势，根系比对照更健康发达，病原含量在11个月后降低了80% ~ 95% Tolfenamic acid can interfere with the interaction between PrbP and RNA polymerase. When applied to citrus plants infected with Huanglongbing via root irrigation or foliar spraying，the treated plants showed recovered growth vigor after two months，with healthier and more developed root systems compared to the control group,and the number of CLas decreased by 80% to 95% after 11 months	Gardner et al.,2016
核苷酸合成 Nucleotide synthesis		IMPDH	两者均为鸟嘌呤核苷酸合成的关键蛋白，肌苷-5’-单磷酸脱氢酶（IMPDH）氧化肌苷-5’-单磷酸为5’-单磷酸黄嘌呤（XMP），鸟苷-5’-单磷酸合成酶（GMPS）再将其转化为鸟苷-5’-单磷酸（GMP） Both of them are key proteins for guanine nucleotide synthesis，Inosine-5’-monophosphate dehydrogenase (IMPDH) oxidizes inosine-5’-monophosphate to 5’-monophosphate xanthine (XMP)，which is converted to guanosine-5’-monophosphate (GMP) by guanosine-5’-monophosphate synthase (GMPS)	布罗波尔，双硫仑 Bronopol，Disulfiram	分子对接结果表明两种化合物可以靶向结合CLas IMPDHΔ98-201，抑制常数（Kᵢ）分别为234 nmol · L^-1和616 nmol · L^-1，抑制IMPDH可阻断GMP合成，从而干扰CLas的增殖 Molecular docking results demonstrated that both compounds can target and bind to CLas IMPDHΔ98-201，with inhibition constants (Kᵢ) of 234 nmol · L^-1 and 616 nmol · L^-1，respectively. By inhibiting IMPDH，these compounds block GMP biosynthesis，thereby disrupting CLas proliferation	Nan et al.,2020
		GMPS		叶酸，AZD1152 Folic acid，AZD1152	综合考虑结合力、溶解性、毒性等指标选出叶酸和化合物AZD1152进行GMPS酶活性抑制试验，不仅有良好的抑制效果，且分子对接结果表明这两种化合物可以和GMPS形成稳定的复合物使其失活 Folic acid and AZD1152 were selected for the GMPS inhibition test by considering the binding force，solubility，toxicity and other indexes，which showed not only good effect，but also stable complexes could be form from these two compounds with GMPS to inactivate it by the molecular docking results	Nan et al.,2021
		YbeY	YbeY具有内切酶活性，参与核糖体RNA的加工，影响细菌共生关系的建立和致病力 YbeY has endonuclease activity and is involved in the processing of ribosomal RNA，which affects the establishment of bacterial symbiosis and pathogenicity	木犀草素 Luteolin	木犀草素是一种黄酮类化合物，在柑橘中含量较少，但是体外试验表明它可以有效抑制YbeY的酶活性，有效降低病菌量，柑橘的黄酮类物质如黄酮、黄酮醇都对YbeY有良好的抑制作用，效果和木犀草素相似 Luteolin is a kind of flavonoids，the content in citrus is relatively small. But in vitro experiments show that it can effectively inhibit the YbeY and reduce the amount of pathogens. Other citrus flavonoids such as flavonoids，flavonols also could inhibit YbeY，and the effects is similar to that of luteolin	Zuo et al.,2019b
其他 Others	CLaBCP		CLaBCP是细菌的过氧化物还原酶，主要作用是维持细胞活性氧、自由基等过氧化物的稳态，参与DNA结合、细胞增殖和凋亡、信号转导等 CLaBCP is a peroxide reductase of bacteria，which mainly plays a role in maintaining the homeostasis of peroxides such as cellular reactive oxygen species and free radicals，and is involved in DNA binding，cell proliferation and apoptosis，signal transduction，etc	Conoidin-A，雷公藤红素 Conoidin-A，celastrol	Conoidin-A和雷公藤红素是两种过氧化物还原酶抑制剂，靶向结合CLas的CLaBCP，形成构象稳定的复合物，在200 μmol · L^-1的Conoidin-A和雷公藤红素分别将CLaBCP的活性降低至215.8 μmol · L^-1·mg^-1·min^-1和216 μmol · L^-1·mg^-1·min^-1，对CLaBCP的抑制效果显著，具有抗CLas的潜力 Conoidin-A and celastrol are two peroxide reductase inhibitors，targeting CLaBCP that binds to CLas to form a conformationally stable complex，and 200 μmol · L^-1 Conoidin-A and celastrol reduced the activity of CLaBCP to 215.8 μmol · L^-1·mg^-1·min^-1 and 216 μmol · L^-1·mg^-1·min^-1，respectively，indicating significant inhibition of CLaBCP and having the potential to fight CLas	Gupta et al.,2023
	Esbp		Esbp是一种胞外结合蛋白，参与细菌铁离子的转运过程 Esbp is an extracellular binding protein involved in the transport of iron ions	ZINC03143779，ZINC05491830，ZINC19210425，ZINC08750867， ZINC14671545	通过虚拟筛选和模拟对接选出了这5种分子，可以靶向结合Esbp，形成的复合物具有多个稳定氢键结构，从而破坏Esbp活性影响CLas对铁离子的吸收，起到抑菌的效果 The five molecules were selected by virtual screening and simulated docking，which showed good binding ability to Esbp，and the complexes were structurally stabilized by multiple hydrogen bonds. Esbp inactivity could affect the uptake of iron ions of CLas，which would have antibacterial effects	Saini et al.,2019
	细胞膜（脂多糖） Cell mem- brane（lipopolysacchari- des）		革兰氏阴性菌外膜含脂多糖，脂多糖既可以增强细菌的抗逆性，也可以成为细菌的毒力因子 The outer membrane of Gram-negative bacteria contains lipopolysaccharides，which can either enhance resistance or act as virulence factors	抗菌肽（AMP） Antimicrobial peptides(AMP)	抗菌肽SAMP可以在细胞膜形成孔隙结构导致胞质泄露和细胞裂解，向感染黄龙病的大叶橙、甜橙、柠檬分别注射10 μmol · L^-1 SAMP，两个月后病菌量相较对照大幅降低，半年内病株恢复长势，新叶没有黄龙病症状 Antimicrobial peptides SAMP can form a pore structure in the cell membrane leading to cytoplasmic leakage and cell lysis. 10 μmol · L^-1 SAMP was injected into infected big-leaf oranges，sweet oranges，and lemons respectively. The amount of the CLas was greatly reduced compared with the control after two months，and the diseased plants regained their growth in half a year，and there were no symptoms in the new leaves	Huang et al.,2021

分类 Category		靶标 Target	作用 Function	小分子化合物 Drug	试验结果 Efficacy	参考文献 Reference
转运系统 Transporter system		SecA	SecA是细菌独有的分泌系统II的组成成分，可以构成SecA-ATP酶，供给能量 SecA is the unique bacterial component of secretion system II that forms the SecA-ATPase to supply energy	4-（4-碘-2-甲基苯氧乙酰基）苯磺酰胺，2-（苯并咪唑-2-硫代）-N-[2-（环丙胺甲酰甲硫基）苯并噻唑-6-基]乙酰胺，2-（5-氨基-1,2,4-三唑-3-硫代）-1-（2,5-二氯苯基-2,4-二甲基环戊二烯基）乙酮， 4-（4-氯-2-甲基苯氧乙酰基）苯磺酰胺 4-[(4-Iodo-2-methylphenoxy)acetyl]benzenesulfonamide，2-(1H-Benzimidazol-2-ylthio)-N-[2-(cyclopropylcarbamoylmethylthio)benzothiazol-6-yl]acetamide，2-(5-Amino-1,2,4-triazol-3-ylthio)-1-(2,5-dichlorophenyl)-2,4-dimethylcyclopent-1-en-1-one， 4-(4-Chloro-2-methylphenoxyacetyl)benzenesulfonamide	采用虚拟筛选和分子对接的方法，发现这4种化合物在20 μmol · L^-1对SecA抑制率大于65%，它们能与SecA的Phe84位点形成稳定的π-π结构，使SecA无法和ATP结合而失活，达到抑菌的目的 Using virtual screening and molecular docking methods，it was found that these four compounds inhibited SecA by more than 65% at 20 μmol · L^-1，and they could form a stable π-π structure with the Phe84 site of SecA，which inactivated SecA by preventing it from combining with ATP，and thus achieved bacterial inhibition	Akula et al.,2011
		SecA		2,4-二氟苄基-4-氟苯基四氢吡唑并吡啶甲酰胺，4-氯苯基-2-乙氧基苯基二氢吡唑并吡啶甲酰胺 (2,4-Difluorobenzyl)(4-fluorophenyl)tetrahydropyrazolopyridinecarboxamide，(4-Chlorophenyl)(2-ethoxyphenyl)dihydropyrazolopyridinecarboxamide	分子对接的结果表明两种化合物可以和SecA的多个残基相互形成稳定的氢键和金属配位互作，浓度为1 mg · mL^-1的P684-2850、P684-3808对L. crescens BT-1的抑制率达99.6%、56.1% The results of molecular docking showed that they could form stable hydrogen bonds and metal-ligand interactions with several residues of SecA，and the inhibition rates of P684-2850 and P684-3808 at a concentration of 1 mg · mL^-1 against L. crescens BT-1 were 99.6% and 56.1% respectively	Zhang et al.,2021c
		ZunA1	黄龙病菌的ABC转运系统的组成部分，负责吸收外界Zn离子 ABC transporter system of Ca. Lasiaticus is responsible for the uptake of external Zn ions	ZINC15670529，ZINC92774705， ZINC06510089，ZINC79841324，ZINC69594834	从ZINC筛选出的5个候选小分子能够与CLas的ZnuA1活性位点残基形成氢键，结合自由能较低，形成稳定的复合物，阻断CLas对Zn²⁺摄取，可以进一步开发为CLas的抑制剂 The five candidate small molecules screened from the ZINC database，which can be further developed as inhibitors of CLas，can form hydrogen bonds with the active site residues of CLas ZnuA1，exhibiting low binding free energy and forming stable complexes，thereby blocking Zn²⁺ uptake by CLas	Saini et al.,2021
调节因子 Regulatory factors		LdtR	LdtR可以感受并调节细胞渗透压、影响细菌细胞壁的合成 LdtR senses and regulates cellular osmotic pressure and influences bacterial cell wall synthesis	根皮素，苯溴马隆，己烷雌酚 Phloretin，Benzbromarone，Hexestrol	根皮素、苯溴马隆、己烷雌酚可以结合CLas的LdtR蛋白形成稳定复合物，干扰细胞的渗透压平衡，利用根瘤菌和L. crescens进行体外实验也证明这3种化合物可以使病菌的细胞吸涨破裂 Phloretin，benzbromarone，and hexestrol can bind to the LdtR protein of CLas to form stable complexes that disrupt cellular osmotic balance. In vitro experiments using Rhizobium and L. crescens have demonstrated that these three compounds induce bacterial cell turgor and subsequent lysis	Pagliai et al.,2014
调节因子 Regulatory factors		PrbP	PrbP和RNA聚合酶β亚基互作调节基因表达，可能与细菌的致病力、细胞活性、抗逆性相关 PrbP and RNA polymerase β subunit interact to regulate gene expression，which may be related to bacterial pathogenicity，cellular activity，and resistance	托芬那酸 Tolfenamic acid	托芬那酸可以干扰PrbP和RNA聚合酶的互作，托芬那酸灌根或喷施感染黄龙病柑橘植株，在2个月后病株恢复长势，根系比对照更健康发达，病原含量在11个月后降低了80% ~ 95% Tolfenamic acid can interfere with the interaction between PrbP and RNA polymerase. When applied to citrus plants infected with Huanglongbing via root irrigation or foliar spraying，the treated plants showed recovered growth vigor after two months，with healthier and more developed root systems compared to the control group,and the number of CLas decreased by 80% to 95% after 11 months	Gardner et al.,2016
核苷酸合成 Nucleotide synthesis		IMPDH	两者均为鸟嘌呤核苷酸合成的关键蛋白，肌苷-5’-单磷酸脱氢酶（IMPDH）氧化肌苷-5’-单磷酸为5’-单磷酸黄嘌呤（XMP），鸟苷-5’-单磷酸合成酶（GMPS）再将其转化为鸟苷-5’-单磷酸（GMP） Both of them are key proteins for guanine nucleotide synthesis，Inosine-5’-monophosphate dehydrogenase (IMPDH) oxidizes inosine-5’-monophosphate to 5’-monophosphate xanthine (XMP)，which is converted to guanosine-5’-monophosphate (GMP) by guanosine-5’-monophosphate synthase (GMPS)	布罗波尔，双硫仑 Bronopol，Disulfiram	分子对接结果表明两种化合物可以靶向结合CLas IMPDHΔ98-201，抑制常数（Kᵢ）分别为234 nmol · L^-1和616 nmol · L^-1，抑制IMPDH可阻断GMP合成，从而干扰CLas的增殖 Molecular docking results demonstrated that both compounds can target and bind to CLas IMPDHΔ98-201，with inhibition constants (Kᵢ) of 234 nmol · L^-1 and 616 nmol · L^-1，respectively. By inhibiting IMPDH，these compounds block GMP biosynthesis，thereby disrupting CLas proliferation	Nan et al.,2020
		GMPS		叶酸，AZD1152 Folic acid，AZD1152	综合考虑结合力、溶解性、毒性等指标选出叶酸和化合物AZD1152进行GMPS酶活性抑制试验，不仅有良好的抑制效果，且分子对接结果表明这两种化合物可以和GMPS形成稳定的复合物使其失活 Folic acid and AZD1152 were selected for the GMPS inhibition test by considering the binding force，solubility，toxicity and other indexes，which showed not only good effect，but also stable complexes could be form from these two compounds with GMPS to inactivate it by the molecular docking results	Nan et al.,2021
		YbeY	YbeY具有内切酶活性，参与核糖体RNA的加工，影响细菌共生关系的建立和致病力 YbeY has endonuclease activity and is involved in the processing of ribosomal RNA，which affects the establishment of bacterial symbiosis and pathogenicity	木犀草素 Luteolin	木犀草素是一种黄酮类化合物，在柑橘中含量较少，但是体外试验表明它可以有效抑制YbeY的酶活性，有效降低病菌量，柑橘的黄酮类物质如黄酮、黄酮醇都对YbeY有良好的抑制作用，效果和木犀草素相似 Luteolin is a kind of flavonoids，the content in citrus is relatively small. But in vitro experiments show that it can effectively inhibit the YbeY and reduce the amount of pathogens. Other citrus flavonoids such as flavonoids，flavonols also could inhibit YbeY，and the effects is similar to that of luteolin	Zuo et al.,2019b
其他 Others	CLaBCP		CLaBCP是细菌的过氧化物还原酶，主要作用是维持细胞活性氧、自由基等过氧化物的稳态，参与DNA结合、细胞增殖和凋亡、信号转导等 CLaBCP is a peroxide reductase of bacteria，which mainly plays a role in maintaining the homeostasis of peroxides such as cellular reactive oxygen species and free radicals，and is involved in DNA binding，cell proliferation and apoptosis，signal transduction，etc	Conoidin-A，雷公藤红素 Conoidin-A，celastrol	Conoidin-A和雷公藤红素是两种过氧化物还原酶抑制剂，靶向结合CLas的CLaBCP，形成构象稳定的复合物，在200 μmol · L^-1的Conoidin-A和雷公藤红素分别将CLaBCP的活性降低至215.8 μmol · L^-1·mg^-1·min^-1和216 μmol · L^-1·mg^-1·min^-1，对CLaBCP的抑制效果显著，具有抗CLas的潜力 Conoidin-A and celastrol are two peroxide reductase inhibitors，targeting CLaBCP that binds to CLas to form a conformationally stable complex，and 200 μmol · L^-1 Conoidin-A and celastrol reduced the activity of CLaBCP to 215.8 μmol · L^-1·mg^-1·min^-1 and 216 μmol · L^-1·mg^-1·min^-1，respectively，indicating significant inhibition of CLaBCP and having the potential to fight CLas	Gupta et al.,2023
	Esbp		Esbp是一种胞外结合蛋白，参与细菌铁离子的转运过程 Esbp is an extracellular binding protein involved in the transport of iron ions	ZINC03143779，ZINC05491830，ZINC19210425，ZINC08750867， ZINC14671545	通过虚拟筛选和模拟对接选出了这5种分子，可以靶向结合Esbp，形成的复合物具有多个稳定氢键结构，从而破坏Esbp活性影响CLas对铁离子的吸收，起到抑菌的效果 The five molecules were selected by virtual screening and simulated docking，which showed good binding ability to Esbp，and the complexes were structurally stabilized by multiple hydrogen bonds. Esbp inactivity could affect the uptake of iron ions of CLas，which would have antibacterial effects	Saini et al.,2019
	细胞膜（脂多糖） Cell mem- brane（lipopolysacchari- des）		革兰氏阴性菌外膜含脂多糖，脂多糖既可以增强细菌的抗逆性，也可以成为细菌的毒力因子 The outer membrane of Gram-negative bacteria contains lipopolysaccharides，which can either enhance resistance or act as virulence factors	抗菌肽（AMP） Antimicrobial peptides(AMP)	抗菌肽SAMP可以在细胞膜形成孔隙结构导致胞质泄露和细胞裂解，向感染黄龙病的大叶橙、甜橙、柠檬分别注射10 μmol · L^-1 SAMP，两个月后病菌量相较对照大幅降低，半年内病株恢复长势，新叶没有黄龙病症状 Antimicrobial peptides SAMP can form a pore structure in the cell membrane leading to cytoplasmic leakage and cell lysis. 10 μmol · L^-1 SAMP was injected into infected big-leaf oranges，sweet oranges，and lemons respectively. The amount of the CLas was greatly reduced compared with the control after two months，and the diseased plants regained their growth in half a year，and there were no symptoms in the new leaves	Huang et al.,2021

柑橘种质/品种 Citrus variety	耐病能力 Disease resistance	参考文献 Reference
十里香 Murraya paniculata（L.）Jack.	感病后没有显著病症，韧皮部没有胼胝质和淀粉堆积，生物胁迫相关基因和通路的表达上调 The infected plants have no significant disease symptoms，no callus，starch accumulation in the phloem，and up-regulation of the expression of biostress-related genes and pathways in citrus after susceptibility to disease	Liu et al.,2023
粗柠檬 Citrus jambhiri	感病后应激反应显著上调，叶中脉的物质转运活性并未受到显著影响 Stress responses were significantly up-regulated in C. jambhiri，and substance transport activity in leaf midveins was not significantly affected	Fan et al.,2012
澳洲手指酸橙 Citrus australasica	病果园内澳洲指橙始终未表现出黄龙病症状，嫁接到阳性砧木后再检测病菌浓度依然很低，具有特定的氧化还原控制系统，感病后还能显著上调囊蛋白和PR1蛋白表达 C. australasica never showed symptom in the diseased orchard，and the pathogen concentration remained low when tested again after grafting onto positive rootstocks. They have a specific redox control system that can significantly up-regulate vesicle protein and PR1 protein expression even after disease susceptibility	Weber et al.,2022
野生柑橘品种 Citrus ichangensis‘2586’	黄龙病菌在C. ichangensis中的繁殖速度较慢；与胼胝质堆积、淀粉合成以及光合作用相关的基因在植株感病后仍维持稳态，避免了韧皮部堵塞和养分不足从而表现出耐病 CLas reproduces slowly in C. ichangensis. Genes related to callus accumulation，starch synthesis，and photosynthesis maintain homeostasis even after disease susceptibility，avoiding phloem clogging and nutrient deficiencies. Thus，it showed disease tolerance	Wu et al.,2020
晚熟温州蜜柑 Citrus unshiu Marc.	自嫁接到阳性病株后8个月内没有检测到黄龙病菌，叶片颜色和健康对照相似，均为正常的绿色，表现出显著的抗性 C. unshiu did not detect the disease for 8 months since grafting onto the positively diseased plants，and the leaf color was similar to that of healthy plants. This variety showed significant resistance	刘登全等,2014
澳指檬 Microcitrus australasica （F.Muell.）Swing.	澳指檬嫁接至感病砂糖橘后多次对黄龙病菌进行检测，结果均为阴性（Ct值 ≥ 40），植株生长正常，叶片无斑驳、黄化等症状 M. australasica was negative (Ct value ≥ 40) for several tests after grafting onto susceptible sugar orange，with normal plant growth and leaf disease	武晓晓等,2023
胡柚、佛手、金柑 Citrus maxima（Burm）Merr. Citrus medica‘Fingered’ Citrus japonica Thunb.	感病温州蜜柑枝条嫁接到3个品种，1年内未出现黄化叶片，感病率为0（未进行qPCR检测）Susceptible mandarin branches grafted to these three varieties did not show yellowing leaves within 1 year，and the susceptibility rate was 0 (not tested by qPCR)	蒋自珍等,2006

Research Progress on the Pathogen Control of Citrus Huanglongbing

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[3]	LI Zhenxi, PAN Ruixuan, XU Meirong, ZHENG Zheng, DENG Xiaoling. Development of Duplex Real-time PCR Assay of‘Candidatus Liberibacter asiatics’ [J]. Acta Horticulturae Sinica, 2023, 50(1): 188-196.
[4]	WEN Qingli，XIE Zhu，WU Liu，HE Yongrui，CHEN Shanchun，and ZOU Xiuping. Clone and Expression Analysis of the Citrus Phloem Protein 2 Gene CsPP2B15 Responding to Huanglongbing Infection in Citrus [J]. ACTA HORTICULTURAE SINICA, 2018, 45(12): 2347-2357.
[5]	HU Yan，WANG Xue-feng，and ZHOU Chang-yong . Recent Advances in Interactions Among‘Candidatus Liberibacter asiaticus’，Insect Vector and Plant Host [J]. ACTA HORTICULTURAE SINICA, 2016, 43(9): 1688-1698.