Research Advances in the Molecular Mechanisms of Plant Response to Saline-alkali Stress

doi:10.13560/j.cnki.biotech.bull.1985.2024-0410

Abstract

Abstract:

Soil salinization is one of a major environmental factor constraining global agricultural development. Breeding saline-alkali tolerant crops is a fundamental strategy to address soil salinization. However, currently there are challenges such as limited saline-alkali-tolerant genes and germplasm resources. Exploring new genes for saline-alkali tolerance and elucidating the molecular mechanisms of plant responses to saline-alkali stress are crucial for developing saline-alkali-tolerant crops. Saline stress includes neutral salt stress and alkaline salt stress. Neutral salt stress induces osmotic stress, and excessive accumulation of Na⁺ and Cl^- may lead to ion toxicity, causing oxidative stress and a series of secondary stresses. Compared to neutral salt stress, alkaline salt stress also induces high pH stress. This review outlines the impact of saline-alkali stress on plant growth and development, as well as the molecular mechanisms underlying plant adaptation to saline-alkali stress and summarizes the significant research progress in plant saline-alkali stress responses over the past decade. It covers plant perception and signal transduction of saline-alkali stress and the molecular mechanisms of plant responses to osmotic stress, ion toxicity, oxidative stress, bicarbonate and carbonate stress, and high pH stress caused by neutral and alkaline salt stress. On this basis, the application of saline-alkali-tolerant genes in crop breeding is also discussed, and key scientific issues that require further research to improve plant saline-alkali tolerance are proposed. The aim of this work is to deepen the understanding of the molecular mechanisms of plant adaptation to saline-alkali stress, providing a theoretical basis for breeding high-yielding and saline-alkali-tolerant crops, and improving the development and utilization rate of salt-affected land.

Key words: saline-alkali stress, ion toxicity, osmotic stress, high pH stress, Na⁺ transporters, plasma membrane H⁺-ATPase

FENG Kai-yue, ZHAO Xin-yan, LI Zi-yan, QIU Jiang-ming, CAO Yi-bo. Research Advances in the Molecular Mechanisms of Plant Response to Saline-alkali Stress[J]. Biotechnology Bulletin, 2024, 40(10): 122-138.

Figures/Tables 4

Fig. 1 Signaling pathways for homeostasis adaptation in plants under neutral salt stress Under salt stress, OSCA1 and GIPC sense changes in external osmotic pressure and Na+ concentration, respectively, mediating Ca2+ influx. SOS3 binds to Ca2+ and interacts with SOS2, activating SOS1 located on the plasma membrane to promote Na+ transport to the extracellular cell. SCaBP8 and CBL8 perform similar functions. SOS2 interacts with AtANN4 to inhibit its mediation of Ca2+ influx. The 14-3-3λ/κ protein and kinase PKS5 inhibit the activity of SOS2. J3, PKS5, and 14-3-3ω are involved in regulating the activity of AHA2. The phosphorylation of SCaBP8 releases its inhibition of AKT1, promoting K+ uptake under salt stress. NHXs, V H+-ATPase, CLCs, and ZmMATE29 located on the vacuolar membrane are involved in the compartmentalization of Na+ and Cl-, respectively. The 14-3-3 proteins activate the K+ efflux channel GORK through both CPK21-dependent and independent pathway, while AtPP2CA inhibits the activity of GORK

Fig. 2 Molecular mechanism of ABA signaling mediating stomatal closure under neutral salt stress Under normal conditions, PP2C binds to SnRK2s protein and inhibits its activity; TOR phosphorylates PYR/PYL/RCAR and suppresses PYR/PYL/RCAR binding to ABA. Under osmotic stress, PP2C forms a complex with PYR/PYL/RCAR to release SnRK2s protein and activate the transcription factor AREB/ABF and the anion channel SLAC1/QUAC1. ABF4 in Arabidopsis thaliana L. activates the expressions of FYVE1 and promote PYR/PYL degradation. Phosphorylation of RaptorB by SnRK2s promotes the dissociation of the TOR complex. SnRK2s can also inhibit the activity of KAT1, decrease cellular turgor pressure, and promote stomatal closure

Fig. 3 Ion transporters mediating the transport and compartmentalization of Na+, K+, and Cl- under neutral salt stress Na+ enters the root epidermal cells through NSCCs and high-affinity K+ transporters such as OSHKT2;1, while SOS1 mediates Na+ efflux. GORK and NSCCs mediate K+ efflux under salt stress. HAK5 and AKT1 mediate K+ influx. In root cortex cells, NPF2.5 is responsible for Cl- efflux, while ZmMATE29 and CLCs transports Cl- into the vacuole. NHX might be involved in the regulation of Na+ compartmentalization into the vacuole. Salt stress induces the expressions of ZmESBL increased, promoting CS development and inhibiting Na+ loading into the xylem via the apoplastic pathway. The CIF-GSO1/SGN3-SGN1 pathway can also enhance the function of the apoplastic barrier. In xylem parenchyma cells, Na+ loading into the xylem is mediated by OsHKT2;1 and SOS1. HKT1, ZmHAK4, and OsHAK12 transport Na+ from xylem vessels to parenchyma cells. SKOR mediates K+ loading into the xylem; while ZmHKT2 removes K+ from the xylem. NPF2.4 and SLAH1/3 mediate the loading and unloading of Cl- in the root xylem

Fig. 4 Molecular mechanism of regulating plasma membrane H+-ATPase activity in plant response to alkaline salt stress Small peptides（Peps）and receptors（PEPRs）sense changes in extracellular pH and activate downstream signaling pathways. Under alkaline salt stress, the cytosolic Ca2+ concentration increases, and SCaBP3 senses the Ca2+ signal and relieves the inhibition of AHA2. Moreover, J3 can inhibit the inhibition of AHA2 by PKS5. Ca2+ binds to ZmNSA1 and induces its degradation through the 26S proteasome pathway, upregulating the expression levels of MHA2 and MHA4. The MPK cascade pathway regulates the transcriptional level and protein activity of PM H+-ATPase under salt stress. Under alkali stress, TaCCD1 and TaSAUR215 interact to inhibit the dephosphorylation of TaPP2C.D1/8 on TaHA2 and enhance the activity of TaHA2

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