
Plant Salt-exclusion Mechanism: A Review
LI Xia, LIU Chuanxin, XU Bin, DONG Rongshu, HUAN Hengfu, HUANG Chunqiong, YAN Linling, WANG Wenqiang, YANG Hubiao, YU Daogeng, WANG Zhiyong, LIU Yiming
Plant Salt-exclusion Mechanism: A Review
This paper aims to provide new ideas for exploring the internal mechanism of plant salt exclusion, screening of salt-exclusion plants and breeding of salt-exclusion crops. The authors review how plants cope with salt stress through salt-exclusion pathways such as tissue structure adaptation, signaling pathway and transporter gene regulation, clarify the key roles of casparian strip of endodermis, suberization of endodermis and exodermis of root, pericycle and xylem parenchyma in plant salt exclusion, and elucidate the species differences in tissue structure in plant salt exclusion. SOS pathway, NHX, HAK and HKT and other transporters play important regulatory roles in the process of plant salt exclusion. SOS pathway, NHX and HAK are ubiquitous in various types of root cells, mainly responsible for Na+ exclusion, transport and vacuole compartmentalization; HKT genes are mainly expressed in xylem parenchyma tissues, and it may play role of salt compartmentalization in xylem parenchyma.
salt-resistant plants / salt exclusion / salt avoidance / pericycle / tissue barriers {{custom_keyword}} /
表1 土壤样品的微生物多样性指数 |
Sample | Sobs | Shannon | Simpson | Ace | Chao1 | Coverage/% |
---|---|---|---|---|---|---|
对照 | 1921.0±136.7a | 6.2±0.1a | 0.007±0.002a | 2242.7±127.3a | 2249.3±118.3a | 98±0.00a |
处理 | 1646.3±61.78b | 5.8±0.12b | 0.012±0.003a | 1906.1±66.1b | 1923.7±92.2b | 98±0.00a |
表2 土壤细菌群落功能丰度表 |
一级功能 | 二级功能 | C | 百分比/% | T | 百分比/% | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Metabolism | Amino Acid Metabolism | 2854091 | 10.47 | 2331140 | 10.18 | |||||||||
Metabolism | Carbohydrate Metabolism | 2670399 | 9.80 | 2206482 | 9.64 | |||||||||
Environmental Information Processing | Membrane Transport | 2604045 | 9.56 | 2383110 | 10.41 | |||||||||
Genetic Information Processing | Replication and Repair | 2021780 | 7.42 | 1708886 | 7.46 | |||||||||
Metabolism | Energy Metabolism | 1804503 | 6.62 | 1491229 | 6.51 | |||||||||
Unclassified | Poorly Characterized | 1398465 | 5.13 | 1172485 | 5.12 | |||||||||
Genetic Information Processing | Translation | 1301802 | 4.78 | 1122984 | 4.90 | |||||||||
Metabolism | Metabolism of Cofactors and Vitamins | 1245312 | 4.57 | 1025830 | 4.48 | |||||||||
Unclassified | Cellular Processes and Signaling | 1034875 | 3.80 | 871649 | 3.81 | |||||||||
Metabolism | Lipid Metabolism | 984298 | 3.61 | 804376 | 3.51 | |||||||||
Metabolism | Nucleotide Metabolism | 925572 | 3.40 | 774079 | 3.38 | |||||||||
Cellular Processes | Cell Motility | 911201 | 3.34 | 863550 | 3.77 | |||||||||
Metabolism | Xenobiotics Biodegradation and Metabolism | 778081 | 2.86 | 593932 | 2.59 | |||||||||
Unclassified | Genetic Information Processing | 735318 | 2.70 | 615411 | 2.69 | |||||||||
Genetic Information Processing | Folding, Sorting and Degradation | 708558 | 2.60 | 616715 | 2.69 | |||||||||
Unclassified | Metabolism | 700475 | 2.57 | 570276 | 2.49 | |||||||||
Metabolism | Glycan Biosynthesis and Metabolism | 627282 | 2.30 | 536108 | 2.34 | |||||||||
Environmental Information Processing | Signal Transduction | 617938 | 2.27 | 531786 | 2.32 | |||||||||
Genetic Information Processing | Transcription | 605085 | 2.22 | 496863 | 2.17 | |||||||||
一级功能 | 二级功能 | C | 百分比/% | T | 百分比/% | |||||||||
Metabolism | Metabolism of Terpenoids and Polyketides | 580423 | 2.13 | 467502 | 2.04 | |||||||||
Metabolism | Enzyme Families | 552628 | 2.03 | 462769 | 2.02 | |||||||||
Metabolism | Metabolism of Other Amino Acids | 499610 | 1.83 | 390798 | 1.71 | |||||||||
Metabolism | Biosynthesis of Other Secondary Metabolites | 279783 | 1.03 | 227262 | 0.99 | |||||||||
Cellular Processes | Cell Growth and Death | 162926 | 0.60 | 131388 | 0.57 | |||||||||
Human Diseases | Infectious Diseases | 116452 | 0.43 | 90288 | 0.39 | |||||||||
Organismal Systems | Endocrine System | 109991 | 0.40 | 90806 | 0.40 | |||||||||
Cellular Processes | Transport and Catabolism | 89898 | 0.33 | 70245 | 0.31 | |||||||||
Human Diseases | Neurodegenerative Diseases | 86995 | 0.32 | 68964 | 0.30 | |||||||||
Environmental Information Processing | Signaling Molecules and Interaction | 46054 | 0.17 | 34673 | 0.15 | |||||||||
Organismal Systems | Environmental Adaptation | 40935 | 0.15 | 35208 | 0.15 | |||||||||
Human Diseases | Cancers | 40283 | 0.15 | 26719 | 0.12 | |||||||||
Human Diseases | Metabolic Diseases | 23895 | 0.09 | 18772 | 0.08 | |||||||||
Organismal Systems | Nervous System | 22358 | 0.08 | 17221 | 0.08 | |||||||||
Organismal Systems | Immune System | 15163 | 0.06 | 12411 | 0.05 | |||||||||
Organismal Systems | Digestive System | 13605 | 0.05 | 8702 | 0.04 | |||||||||
Organismal Systems | Circulatory System | 11550 | 0.04 | 10397 | 0.05 | |||||||||
Human Diseases | Immune System Diseases | 10862 | 0.04 | 7727 | 0.03 | |||||||||
Organismal Systems | Excretory System | 10817 | 0.04 | 8228 | 0.04 | |||||||||
Human Diseases | Cardiovascular Diseases | 4608 | 0.02 | 1683 | 0.01 | |||||||||
Cellular Processes | Cell Communication | 54 | 0.00 | 36 | 0.00 | |||||||||
Organismal Systems | Sensory System | 18 | 0.00 | 12 | 0.00 |
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Intracellular Na+/H+ antiporters (NHXs) play important roles in cellular pH and Na+ and K+ homeostasis in all eukaryotes. Based on sequence similarity, the six intracellular Arabidopsis thaliana members are divided into two groups. Unlike the vacuolar NHX1-4, NHX5 and NHX6 are believed to be endosomal; however, little data exist to support either their function or localization. Using reverse genetics, we show that whereas single knockouts nhx5 or nhx6 did not differ from the wild type, the double knockout nhx5 nhx6 showed reduced growth, with smaller and fewer cells and increased sensitivity to salinity. Reduced growth of nhx5 nhx6 was due to slowed cell expansion. Transcriptome analysis indicated that nhx5, nhx6, and the wild type had similar gene expression profiles, whereas transcripts related to vesicular trafficking and abiotic stress were enriched in nhx5 nhx6. We show that unlike other intracellular NHX proteins, NHX5 and NHX6 are associated with punctate, motile cytosolic vesicles, sensitive to Brefeldin A, that colocalize to known Golgi and trans-Golgi network markers. We provide data to show that vacuolar trafficking is affected in nhx5 nhx6. Possible involvements of NHX5 and NHX6 in maintaining organelle pH and ion homeostasis with implications in endosomal sorting and cellular stress responses are discussed.
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[87] |
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[88] |
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[89] |
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[90] |
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[91] |
The Arabidopsis high-affinity K+ transporter (AtHKT1;1) plays roles in salt tolerance by unloading Na+ from the root xylem to the xylem parenchyma cells and/or uploading Na+ from the shoot/leaf xylem to the xylem parenchyma cells. To use this promoter for the molecular breeding of salt-tolerant plants, I evaluated the expression profile of the AtHKT1;1 promoter in detail. Approximately 1.1 kbp of sequence upstream from the start codon of AtHKT1;1 was polymerase chain reaction (PCR)-amplified, fused to the β-glucuronidase (GUS) gene, and introduced into Arabidopsis. The resultant transformants were evaluated under nonstressed and salt-stress conditions at the seedling and reproductive stages. Histochemical analysis showed that GUS activity was detected in vascular bundle tissue in roots, hypocotyls, petioles, leaves, and petals, and in root tips. GUS enzyme activity in shoots tended to be higher than that in roots at both stages. After treatment with 50 mM NaCl for 24 h, GUS transcription levels and GUS enzyme activity were enhanced in transgenic lines. These results indicate that the AtHKT1;1 promoter isolated in this study could be useful in expressing transgenes specifically in vascular tissue and root tips, and in a mild salt-stress-responsive manner. The data provide novel insights into the functions of AtHKT1;1.
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[92] |
HKT channels are a plant protein family involved in sodium (Na+) and potassium (K+) uptake and Na+-K+ homeostasis. Some HKTs underlie salt tolerance responses in plants, while others provide a mechanism to cope with short-term K+ shortage by allowing increased Na+ uptake under K+ starvation conditions. HKT channels present a functionally versatile family divided into two classes, mainly based on a sequence polymorphism found in the sequences underlying the selectivity filter of the first pore loop. Physiologically, most class I members function as sodium uniporters, and class II members as Na+/K+ symporters. Nevertheless, even within these two classes, there is a high functional diversity that, to date, cannot be explained at the molecular level. The high complexity is also reflected at the regulatory level. HKT expression is modulated at the level of transcription, translation, and functionality of the protein. Here, we summarize and discuss the structure and conservation of the HKT channel family from algae to angiosperms. We also outline the latest findings on gene expression and the regulation of HKT channels.
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[93] |
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[94] |
李学文, 游西龙, 王艳. 钾离子转运载体HAK/KUP/KT家族参与植物耐盐性的研究进展[J]. 植物科学学报, 2019, 37(1):101-108.
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[95] |
游西龙. 两个HAK/KUP/KT成员的钾转运功能及耐盐性分析[D]. 乌鲁木齐: 新疆大学, 2018:15-21.
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[96] |
<p>Potassium (K<sup>+</sup>) is an essential macronutrient in plants and a lack of K<sup>+</sup> significantly reduces the potential for plant growth and development. By contrast, sodium (Na<sup>+</sup>), while beneficial to some extent, at high concentrations it disturbs and inhibits various physiological processes and plant growth. Due to their chemical similarities, some functions of K<sup>+</sup> can be undertaken by Na<sup>+</sup> but K<sup>+</sup> homeostasis is severely affected by salt stress, on the other hand. Recent advances have highlighted the fascinating regulatory mechanisms of K<sup>+</sup> and Na<sup>+</sup> transport and signaling in plants. This review summarizes three major topics: (i) the transport mechanisms of K<sup>+</sup> and Na<sup>+</sup> from the soil to the shoot and to the cellular compartments; (ii) the mechanisms through which plants sense and respond to K<sup>+</sup> and Na<sup>+</sup> availability; and (iii) the components involved in maintenance of K<sup>+</sup>/Na<sup>+</sup> homeostasis in plants under salt stress.</p><p>Adams E, Shin R (2014) Transport, signaling, and homeostasis of potassium and sodium in plants. <strong>J Integr Plant Biol</strong> 56: 231–249. doi: 10.1111/jipb.12159</p>
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[97] |
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[98] |
Background and Aims In the present study, we show that development of endodermis and exodermis is sensitively regulated by water accessibility. As cadmium (Cd) is known to induce xeromorphic effects in plants, maize roots were exposed also to Cd to understand the developmental process of suberin lamella deposition in response to a local Cd source. Methods In a first experiment, maize roots were cultivated in vitro and unilaterally exposed to water-containing medium from one side and to air from the other. In a second experiment, the roots were placed between two agar medium layers with a strip of Cd-containing medium attached locally and unilaterally to the root surface. Key Results The development of suberin lamella (the second stage of exodermal and endodermal development) started asymmetrically, preferentially closer to the root tip on the side exposed to the air. In the root contact with Cd in a spatially limited area exposed to one side of the root, suberin lamella was preferentially developed in the contact region and additionally along the whole length of the root basipetally from the contact area. However, the development was unilateral and asymmetrical, facing the treated side. The same pattern occurred irrespective of the distance of Cd application from the root apex. Conclusions These developmental characteristics indicate a sensitive response of root endodermis and exodermis in the protection of vascular tissues against abiotic stresses.© The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
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[99] |
杨朝东, 张霞, 刘国锋, 等. 植物根中质外体屏障结构和生理功能研究进展[J]. 植物研究, 2013, 33(1):114-119.
综述了近10年来植物根中质外体屏障结构和功能的研究进展。质外体屏障指根中内、外皮层初生壁的凯氏带,或次生壁栓质化和木质化,以及植物体表角质层组成的保护组织,能隔绝水、离子和氧气不能自由进出植物体的屏障结构,具有保护植物体的生理功能。根中凯氏带的分子发育机理研究表明根内皮层类似哺乳动物上皮组织的保护作用。植物根中质外体保证内部各种生理代谢在稳定的内部环境中进行,是植物适应各种逆境的重要屏障结构。根中质外体屏障在植物适应干旱、洪涝灾害、离子胁迫和病虫害的侵袭等方面具有重要作用,在探索适应并修复极端生态环境的植物资源中有广阔的应用前景。
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