植物盐腺泌盐及发育研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2023-0581

摘要/Abstract

摘要：

盐腺是泌盐盐生植物抵御盐胁迫的重要表皮结构，泌盐盐生植物可以通过盐腺将体内多余的盐离子排出体外，从而避免盐胁迫。盐腺作为泌盐盐生植物实现高效抗盐的重要结构，在逆境生理、发育和进化等领域都引起了关注和讨论，集中在盐腺的超微结构、生理功能、泌盐机制以及发育模式等不同层面已有广泛的研究报道。本文综述了盐腺结构、分泌机制、盐腺发育的研究进展，总结了盐腺泌盐的可能途径以及盐腺发育的调控方式和关键基因，对未来盐腺泌盐和发育的研究提出了相关见解，讨论了盐腺这一独特形态学结构对于植物耐盐性的作用，并对提高植物耐盐性、培育耐盐品种提出了理论依据和建议，有利于深入解析植物耐盐适应演化、培育抗盐作物和高效利用盐碱地。

关键词: 盐腺, 盐腺泌盐, 盐腺发育, 抗盐性

Abstract:

Salt gland is one of the important epidermal structures of recreto-halophytes in resistance to salt stress, which secrete excessive salt ions out of the body to prevent the plant from being damaged by salt stress. As an important structure to achieve high salt resistance in halophytes, salt glands deserve extensive attention and discussion in the fields of stress physiology, development and evolutionary biology. A large number of studies have been reported in the ultrastructure, physiological function, mechanism of salt secretion and developmental pattern of salt glands. In this paper, we reviewed the research progress of salt gland structure, secretion mechanism and salt gland development, summarized the feasible pathways of salt gland secretion and the regulation mode and key genes of salt gland development, proposed relevant views for future research on salt gland secretion and development, and discussed the role of this unique morphological structure of salt gland on plant salt tolerance. We also proposed the theoretical basis and suggestions for improving plant salt tolerance and breeding salt-tolerant plant varieties. The related studies will be beneficial for in-depth analysis of plant salt tolerance adaptation evolution, cultivation of salt resistant crops, and efficient utilization of saline land.

Key words: salt gland, salt gland excretion, salt gland development, salt resistance

马秋雨, 袁芳. 植物盐腺泌盐及发育研究进展[J]. 生物技术通报, 2023, 39(11): 74-85.

MA Qiu-yu, YUAN Fang. Research Progress in Salt Gland Secretion and Development in Plants[J]. Biotechnology Bulletin, 2023, 39(11): 74-85.

图/表 4

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Family	Genus	Species	Reference
Acanthaceae	Acanthus	A. ebracteatus, A. ilicifolius	[11]
Acanthaceae	Avicennia	A. germinans, A. marina, A. officinalis, A. schaueriana, A. bicolor	[12]
Aizoaceae	Sesuvium	S. sesuvioides, S. portulacastrum	[13]
Aizoaceae	Mesembryanthemum	M. nodiflorum, M. crystallinum	[14]
Amaranthaceae	Atriplex	A. farinosa, A. centralasiatica, A. confertifolia, A. hortensis, A. hymenelytra, A. laciniata, A. micrantha, A. oestophora, A. patens, A. portulacoides, A. rosea, A. prostrata, A. prostrata, A. tatarica, A. verrucifera, A. littoralis, A. nummularia, A. cristata, A. canescens, A. halimus, A. semibaccata, A. vesicaria, A. spongiosa, A. amnicola, A. lindleyi, A. turcomanica	[15]
Amaranthaceae	Chenopodium	C. album	[16]
Amaranthaceae	Bienertia	B. sinuspersici	[17]
Amaranthaceae	Oxybasis	O. glauca	[18]
Amaranthaceae	Chenopodiastrum	C. murale	[19]
Apiaceae	Glehnia	G. littoralGis	[20]
Asteraceae	Pseudobaccharis	P. spartioides	[21]
Convolvulaceae	Cressa	C. cretica	[22]
Convolvulaceae	Calystegia	C. soldanella	[23]
Fabaceae	Glycyrrhiza	G. uralensis	[24]
Frankeniaceae	Frankenia	F. juniperoides, F. pauciflora, F. corymbosa, F. salina, F. hirsuta, F. laevis, F. pulverulenta, F. thymifolia	[21]
Malvaceae	Malva	M. arborea	[25]
Oleaceae	Phillyrea	P. latifolia	[26]
Plumbaginaceae	Limonium	L. otolepis, L. suffruticosum, L. reniforme, L. santapolense, L. girardianum, L. platyphyllum, L. bellidifolium, L. binervosum, L. nudum, L. aureum, L. axillare, L. iranicum, L. lobatum, L. oleifolium, L. virgatum, L. pectinatum, L. perezii, L. sinense, L. sinuatum, L. stocksii, L. franchetii, L. bicolor, L. caspium, L. pruinosum, L. gmelinii, L. vulgare, L. brasiliense, L. californicum, L. narbonense, L. delicatulum, L. ovalifolium, L. ovalifolium	[27]
Plumbaginaceae	Goniolimon	G. tataricum	[28]
Plumbaginaceae	Plumbago	P. auriculata	[29]
Plumbaginaceae	Aegialitis	A. annulata, A. rotundifolia	[30]
Plumbaginaceae	Armeria	A. maritima	[31]
Plumbaginaceae	Limoniastrum	L. monopetalum, L. guyonianum	[32]
Plumbaginaceae	Ceratostigma	C. plumbaginoides	[33]
Poaceae	Distichlis	D. humilis, D. spicata	[34]
Poaceae	Oryza	O. coarctata	[35]
Poaceae	Odyssea	O. paucinervis	[36]
Poaceae	Spinifex	S. hirsutus	[37]
Poaceae	Sporobolus	S. foliosus, S. montevidensis, S. xtownsendii, S. helvolus, S. virginicus, S. airoides, S. spicatus, S. michauxianus, S. ioclados, S. alterniflorus, S. anglicus, S. pungens, S. maritimus	[38]
Poaceae	Leptochloa	L. crinita	[39]
Poaceae	Zoysia	Z. macrantha, Z. matrella, Z. macrostachya, Z. japonica, Z. matrella	[37]
Poaceae	Chloris	C. gayana	[37]
Poaceae	Cynodon	C. dactylon	[37]
Poaceae	Dactyloctenium	D. aegyptium	[40]
Poaceae	Diplachne	D. fusca	[37]
Poaceae	Eleusine	E. indica	[37]
Poaceae	Panicum	P. virgatum	[37]
Poaceae	Pappophorum	P. philippianum	[41]
Poaceae	Setaria	S. viridis	[42]
Poaceae	Aeluropus	A. lagopoides, A. littoralis	[43]
22Poaceae	Imperata	I. cylindrica	[37]
Poaceae	Cenchrus	C. clandestinum	[37]
Poaceae	Bouteloua	B. dactyloides	[44]
Primulaceae	Lysimachia	L. maritima	[45]
Primulaceae	Aegiceras	A. floridum, A. corniculatum	[11]
Scrophulariaceae	Myoporum	M. bontioides	[46]
Tamaricaceae	Reaumuria	R. alternifolia, R. hirtella, R. trigyna	[47]
Tamaricaceae	Tamarix	T. amplexicaulis, T. arborea, T. karelini, T. gansuensisi, T. smyrnensis, T. passerinoides, T. laxa, T. hispida, T. elongata, T. austromongolica, T. arceuthoides, T. nilotica, T. africana, T. usneoides, T. chinensis, T. ramosissima, T. aphylla, T. hohenackeri, T. gallica	[48]
Verbenaceae	Phyla	P. nodiflora	[48]

Family	Genus	Species	Reference
Acanthaceae	Acanthus	A. ebracteatus, A. ilicifolius	[11]
Acanthaceae	Avicennia	A. germinans, A. marina, A. officinalis, A. schaueriana, A. bicolor	[12]
Aizoaceae	Sesuvium	S. sesuvioides, S. portulacastrum	[13]
Aizoaceae	Mesembryanthemum	M. nodiflorum, M. crystallinum	[14]
Amaranthaceae	Atriplex	A. farinosa, A. centralasiatica, A. confertifolia, A. hortensis, A. hymenelytra, A. laciniata, A. micrantha, A. oestophora, A. patens, A. portulacoides, A. rosea, A. prostrata, A. prostrata, A. tatarica, A. verrucifera, A. littoralis, A. nummularia, A. cristata, A. canescens, A. halimus, A. semibaccata, A. vesicaria, A. spongiosa, A. amnicola, A. lindleyi, A. turcomanica	[15]
Amaranthaceae	Chenopodium	C. album	[16]
Amaranthaceae	Bienertia	B. sinuspersici	[17]
Amaranthaceae	Oxybasis	O. glauca	[18]
Amaranthaceae	Chenopodiastrum	C. murale	[19]
Apiaceae	Glehnia	G. littoralGis	[20]
Asteraceae	Pseudobaccharis	P. spartioides	[21]
Convolvulaceae	Cressa	C. cretica	[22]
Convolvulaceae	Calystegia	C. soldanella	[23]
Fabaceae	Glycyrrhiza	G. uralensis	[24]
Frankeniaceae	Frankenia	F. juniperoides, F. pauciflora, F. corymbosa, F. salina, F. hirsuta, F. laevis, F. pulverulenta, F. thymifolia	[21]
Malvaceae	Malva	M. arborea	[25]
Oleaceae	Phillyrea	P. latifolia	[26]
Plumbaginaceae	Limonium	L. otolepis, L. suffruticosum, L. reniforme, L. santapolense, L. girardianum, L. platyphyllum, L. bellidifolium, L. binervosum, L. nudum, L. aureum, L. axillare, L. iranicum, L. lobatum, L. oleifolium, L. virgatum, L. pectinatum, L. perezii, L. sinense, L. sinuatum, L. stocksii, L. franchetii, L. bicolor, L. caspium, L. pruinosum, L. gmelinii, L. vulgare, L. brasiliense, L. californicum, L. narbonense, L. delicatulum, L. ovalifolium, L. ovalifolium	[27]
Plumbaginaceae	Goniolimon	G. tataricum	[28]
Plumbaginaceae	Plumbago	P. auriculata	[29]
Plumbaginaceae	Aegialitis	A. annulata, A. rotundifolia	[30]
Plumbaginaceae	Armeria	A. maritima	[31]
Plumbaginaceae	Limoniastrum	L. monopetalum, L. guyonianum	[32]
Plumbaginaceae	Ceratostigma	C. plumbaginoides	[33]
Poaceae	Distichlis	D. humilis, D. spicata	[34]
Poaceae	Oryza	O. coarctata	[35]
Poaceae	Odyssea	O. paucinervis	[36]
Poaceae	Spinifex	S. hirsutus	[37]
Poaceae	Sporobolus	S. foliosus, S. montevidensis, S. xtownsendii, S. helvolus, S. virginicus, S. airoides, S. spicatus, S. michauxianus, S. ioclados, S. alterniflorus, S. anglicus, S. pungens, S. maritimus	[38]
Poaceae	Leptochloa	L. crinita	[39]
Poaceae	Zoysia	Z. macrantha, Z. matrella, Z. macrostachya, Z. japonica, Z. matrella	[37]
Poaceae	Chloris	C. gayana	[37]
Poaceae	Cynodon	C. dactylon	[37]
Poaceae	Dactyloctenium	D. aegyptium	[40]
Poaceae	Diplachne	D. fusca	[37]
Poaceae	Eleusine	E. indica	[37]
Poaceae	Panicum	P. virgatum	[37]
Poaceae	Pappophorum	P. philippianum	[41]
Poaceae	Setaria	S. viridis	[42]
Poaceae	Aeluropus	A. lagopoides, A. littoralis	[43]
22Poaceae	Imperata	I. cylindrica	[37]
Poaceae	Cenchrus	C. clandestinum	[37]
Poaceae	Bouteloua	B. dactyloides	[44]
Primulaceae	Lysimachia	L. maritima	[45]
Primulaceae	Aegiceras	A. floridum, A. corniculatum	[11]
Scrophulariaceae	Myoporum	M. bontioides	[46]
Tamaricaceae	Reaumuria	R. alternifolia, R. hirtella, R. trigyna	[47]
Tamaricaceae	Tamarix	T. amplexicaulis, T. arborea, T. karelini, T. gansuensisi, T. smyrnensis, T. passerinoides, T. laxa, T. hispida, T. elongata, T. austromongolica, T. arceuthoides, T. nilotica, T. africana, T. usneoides, T. chinensis, T. ramosissima, T. aphylla, T. hohenackeri, T. gallica	[48]
Verbenaceae	Phyla	P. nodiflora	[48]

基因 Gene	蛋白质 Protein	功能 Function	参考文献 Reference
NHX1	Na⁺/H⁺逆向转运蛋白	将Na⁺泵出细胞外或储存在液泡	[75]
NHX7	Na⁺/H⁺逆向转运蛋白	将Na⁺泵出细胞外或储存在液泡	[76]
AtCCC、NKCC	Na⁺:K⁺:Cl^-共转运体	参与Cl^-的长距离运输	[77-78]
AlHKT2;1	高亲和性钾转运蛋白	将Na⁺泵入细胞	[79]

基因 Gene	蛋白质 Protein	功能 Function	参考文献 Reference
NHX1	Na⁺/H⁺逆向转运蛋白	将Na⁺泵出细胞外或储存在液泡	[75]
NHX7	Na⁺/H⁺逆向转运蛋白	将Na⁺泵出细胞外或储存在液泡	[76]
AtCCC、NKCC	Na⁺:K⁺:Cl^-共转运体	参与Cl^-的长距离运输	[77-78]
AlHKT2;1	高亲和性钾转运蛋白	将Na⁺泵入细胞	[79]

转录因子 Transcription factor	基因 Gene	功能 Function	参考文献 Reference
bHLH	ENHANCER OF GLABRA3（EGL3）, LbHLH	盐腺发育的负调节因子	[88,92]
MYB	TRIPTYCHON（TRY）	盐腺发育的负调节因子	[88,93]
MYB	CAPRICE（CPC）	盐腺发育的负调节因子	[88,94]
WD40 repeat-like superfamily	TRANSPARENT TESTA GLABRA1（TTG1）	盐腺发育的负调节因子	[88]
ARM repeat superfamily	SENSITIVE TO ABA AND DROUGHT2（SAD2）	盐腺发育的正调节因子	[88,95]
NAC	LbNAC4	盐腺发育的正调节因子	[56]