NaCl在四翅滨藜适应渗透胁迫中的作用

doi:10.11686/cyxb2019394

摘要/Abstract

摘要： 泌盐盐生植物四翅滨藜是一种多年生C₄灌木,具有良好的适应盐碱和干旱环境的能力。前期研究发现,通过将过量Na⁺转运至叶表面盐囊泡、增加Na⁺对渗透势的贡献以提高植株保水能力是四翅滨藜耐盐的主要策略,且100 mmol·L^-1 NaCl可显著刺激其生长。为进一步探究NaCl是否有助于缓解渗透胁迫对四翅滨藜生长的不利影响,本研究分别用不含和含有100 mmol·L^-1 NaCl的山梨醇引起的总渗透势为-0.2和-0.5 MPa的1/2 Hoagland营养液处理5周龄四翅滨藜幼苗5 d。结果表明,在渗透胁迫下,NaCl的添加可增加叶片相对含水量、提高净光合速率以及降低叶渗透势(Ψ_s),从而促进四翅滨藜的生长。NaCl的添加还显著增加了茎叶组织及叶盐囊泡中的Na⁺积累并提高了Na⁺对叶Ψ_s的贡献,且对茎叶组织中K⁺含量影响较小。此外,渗透胁迫下,添加一定浓度NaCl显著增加了植株中游离脯氨酸和甜菜碱含量,它们除作为保护剂减轻渗透胁迫损伤外,还参与植株的渗透调节,从而进一步提高四翅滨藜对渗透胁迫的适应性。综上所述,适宜浓度的NaCl的确能够有效缓解渗透胁迫对四翅滨藜生长的不利影响。

关键词: 四翅滨藜, 盐生植物, 渗透胁迫, NaCl, 渗透调节

Abstract: Secretohalophyte Atriplex canescens is a C₄ perennial shrub with excellent resistance to salinity and drought. Our previous study showed that the transport of excessive Na⁺ into leaf salt bladders and contribution of the elevated Na⁺ concentration to leaf osmotic potential (Ψ_s) improves water status of plants and is a primary strategy in salt tolerance of A. canescens. The external application of 100 mmol·L^-1 NaCl can substantially stimulate the growth of A. canescens. To investigate whether NaCl could help to alleviate deleterious impacts of osmotic stress on the growth of A. canescens, five-week-old seedlings were subjected to two osmotic stress levels (-0.2 and -0.5 MPa) induced by D-sorbitol in the presence or absence of an additional 100 mmol·L^-1 NaCl. It was found that under osmotic stress, the addition of NaCl obviously improved the growth of A. canescens. Leaf relative water content was increased, net photosynthetic rate was enhanced and significantly more negative Ψ_s was induced. Furthermore, exogenous NaCl significantly increased Na⁺ concentration in stem, leaf tissues and leaf salt bladders as well as enhancing the Na⁺ contribution to leaf Ψ_s, with little adverse effect on K⁺ accumulation in stem and leaf tissues. In addition, the content of free proline and betaine in plants was significantly increased in the presence of NaCl under osmotic stress, and their contributions to leaf Ψ_s were significantly increased by additional NaCl under osmotic stress. This suggests that compatible solutes, in addition to functioning as protectants, are also involved in osmotic adjustment to further alleviate the deleterious impacts of osmotic stress. In summary, the appropriate concentration of NaCl does effectively alleviate the adverse impacts of osmotic stress on the growth of A. canescens.

Key words: Atriplex canescens, halophyte, osmotic stress, NaCl, osmotic adjustment

郭欢, 潘雅清, 包爱科. NaCl在四翅滨藜适应渗透胁迫中的作用[J]. 草业学报, 2020, 29(7): 112-121.

GUO Huan, Pan Ya-qing, BAO Ai-ke. Effect of NaCl on the adaption of Atriplex canescens under osmotic stress[J]. Acta Prataculturae Sinica, 2020, 29(7): 112-121.

参考文献

[1] Xiong L, Zhu J K. Molecular and genetic aspects of plant responses to osmotic stress. Plant, Cell and Environment, 2002, 25(2): 131-139.
[2] Yue L J, Cui Y N, Yuan K, et al. The osmotic adjustment in Pugionium cornutum subjected to salt stress. Plant Physiology Journal, 2016, 52(4): 569-574.
岳利军, 崔彦农, 袁坤, 等. NaCl胁迫下沙芥的渗透调节作用. 植物生理学报, 2016, 52(4): 569-574.
[3] Vurukonda S S K P, Vardharajula S, Shrivastava M, et al. Enhancement of drought stress tolerance in crops by plant growth promoting rhizobacteria. Microbiological Research, 2016, 184: 13-24.
[4] Martínez J P, Kinet J M, Bajji M, et al. NaCl alleviates polyethylene glycol-induced water stress in the halophyte species Atriplex halimus L. Journal of Experimental Botany, 2005, 56(419): 2421-2431.
[5] Carmo-Silva A E, Gore M A, Andrade-Sanchez P, et al. Decreased CO₂ availability and inactivation of rubisco limit photosynthesis in cotton plants under heat and drought stress in the field. Environmental and Experimental Botany, 2012, 83: 1-11.
[6] Shabala S, Bose J, Hedrich R. Salt bladders: Do they matter? Trends in Plant Science, 2014, 19(11): 687-691.
[7] Oh D H, Barkla B J, Vera-Estrella R, et al. Cell type-specific responses to salinity-the epidermal bladder cell transcriptome of Mesembryanthemum crystallinum. New Phytologist, 2015, 207(3): 627-644.
[8] Nemat Alla M M, Khedr A H A, Serag M M, et al. Regulation of metabolomics in Atriplex halimus growth under salt and drought stress. Plant Growth Regulation, 2012, 67: 281-304.
[9] Wang Z X, Zhang S Z. Research progress of drought resistance and drought-resistant improvement in crop. Crops, 2014, 2: 26-31.
王尊欣, 张树珍. 作物抗旱性及抗旱育种研究进展. 作物杂志, 2014, 2: 26-31.
[10] Hao G Y, Lucero M E, Sanderson S C, et al. Polyploidy enhances the occupation of heterogeneous environments through hydraulic related trade-offs in Atriplex canescens (Chenopodiaceae). New Phytologist, 2013, 197(3): 970-978.
[11] Kong D S. Morphological characteristics and eco-physiological adaptability of Atriplex canescens: A review. Chinese Journal of Ecology, 2013, 32(1): 210-216.
孔东升. 四翅滨藜形态特征及生理生态适应性. 生态学杂志, 2013, 32(1): 210-216.
[12] Glenn E, Pfister R, Brown J J, et al. Na and K accumulation and salt tolerance of Atriplex canescens (Chenopodiaceae) genotypes. American Journal of Botany, 1996, 83(8): 997-1005.
[13] Benzarti M, Ben Rejeb K, Debez A, et al. Environmental and economical opportunities for the valorisation of the genus Atriplex: New insights//Hakeem K, Ahmad P, Ozturk M. Crop improvement: New approaches and modern techniques. New York, USA: Springer, 2013: 441-457.
[14] Kiani-Pouya A, Roessner U, Jayasinghe N S, et al. Epidermal bladder cells confer salinity stress tolerance in the halophyte quinoa and Atriplex species. Plant, Cell and Environment, 2017, 40: 1900-1915.
[15] Pan Y Q, Guo H, Wang S M, et al. The photosynthesis, Na⁺/K⁺ homeostasis and osmotic adjustment of Atriplex canescens in response to salinity. Frontiers in Plant Science, 2016, (7): 848.
[16] Ma Q, Yue L J, Zhang J L, et al. Sodium chloride improves photosynthesis and water status in the succulent xerophyte Zygophyllum xanthoxylum. Tree Physiology, 2012, 32(1): 4-13.
[17] Tsutsumi K, Yamada N, Chaum S, et al. Differential accumulation of glycinebetaine and choline monooxygenase in bladder hairs and lamina leaves of Atriplex gmelini under high salinity. Journal of Plant Physiology, 2015, 176: 101-107.
[18] Wang S, Wan C, Wang Y, et al. The characteristics of Na⁺, K⁺ and free proline distribution in several drought-resistant plants of the Alxa Desert, China. Journal of Arid Environments, 2004, 56(3): 525-539.
[19] Guerrier G. Fluxes of Na⁺, K⁺ and Cl^-, and osmotic adjustment in Lycopersicon pimpinellifolium and L. esculentum during short- and long-term exposures to NaCl. Physiologia Plantarum, 1996, 97(3): 583-591.
[20] Wang Q Z, Liu Q, Gao Y N, et al. Review on the mechanisms of the response to salinity-alkalinity stress in plants. Acta Ecologica Sinica, 2017, 37(16): 5565-5577.
王佺珍, 刘倩, 高娅妮, 等. 植物对盐碱胁迫的响应机制研究进展. 生态学报, 2017, 37(16): 5565-5577.
[21] Shabala S, Mackay A. Ion transport in halophytes. Advances in Botanical Research, 2011, 57: 151-199.
[22] Cui Y N, Xia Z R, Ma Q, et al. The synergistic effects of sodium and potassium on the xerophyte Apocynum venetum in response to drought stress. Plant Physiology and Biochemistry, 2019, 135: 489-498.
[23] He F L, Bao A K, Wang S M, et al. NaCl stimulates growth and alleviates drought stress the salt-secreting xerophyte Reaumuria soongorica. Environmental and Experimental Botany, 2019, 162: 433-443.
[24] Jupa R, Plichta R, Paschová Z, et al. Mechanisms underlying the long-term survival of the monocot Dracaena marginata under drought conditions. Tree Physiology, 2017, 37(9): 1182-1197.
[25] Giorio P, Sorrentino G, D'Andria R. Stomatal behaviour, leaf water status and photosynthetic response in field-grown olive trees under water deficit. Environmental and Experimental Botany, 1999, 42(2): 95-104.
[26] Hedrich R, Shabala S. Stomata in a saline world. Current Opinion in Plant Biology, 2018, 46: 87-95.
[27] Ache P, Bauer H, Kollist H, et al. Stomatal action directly feeds back on leaf turgor: New insights into the regulation of the plant water status from non-invasive pressure probe measurements. The Plant Journal, 2010, 62(6): 1072-1082.
[28] Kronzucker H J, Coskun D, Schulze L M, et al. Sodium as nutrient and toxicant. Plant and Soil, 2013, 369: 1-23.
[29] Guo H, Zhang L, Cui Y N, et al. Identification of candidate genes related to salt tolerance of the secretohalophyte Atriplex canescens by transcriptomic analysis. BMC Plant Biology, 2019, 19: 213.
[30] Flowers T J, Munns R, Colmer T D. Sodium chloride toxicity and the cellular basis of salt tolerance in halophytes. Annals of Botany, 2015, 115(3): 419-431.
[31] Böhm J, Messerer M, Müller H M, et al. Understanding the molecular basis of salt sequestration in epidermal bladder cells of Chenopodium quinoa. Current Biology, 2018, 28: 1-11.
[32] Zhang L, Guo H, Bao A K. The unique salt-secreting structures of halophytes: Salt bladders. Plant Physiology Journal, 2019, 55(3): 232-240.
张乐, 郭欢, 包爱科. 盐生植物的独特泌盐结构—盐囊泡. 植物生理学报, 2019, 55(3): 232-240.
[33] Benito B, Haro R, Amtmann A, et al. The twins K⁺ and Na⁺ in plants. Journal of Plant Physiology, 2014, 171(9): 723-731.
[34] Martineau E, Domec J C, Bosc A, et al. The effects of potassium nutrition on water use in field-grown maize (Zea mays L.). Environmental and Experimental Botany, 2017, 134: 62-71.
[35] Hessini K, Martínez J P, Gandour M, et al. Effect of water stress on growth, osmotic adjustment, cell wall elasticity and water-use efficiency in Spartina alterniflora. Environmental and Experimental Botany, 2009, 67(2): 312-319.
[36] Ming D F, Pei Z F, Naeem M S, et al. Silicon alleviates PEG-induced water-deficit stress in upland rice seedlings by enhancing osmotic adjustment. Journal of Agronomy and Crop Science, 2012, 198: 14-26.
[37] Wang B, Chen J, Chen L, et al. Combined drought and heat stress in Camellia oleifera cultivars: Leaf characteristics, soluble sugar and protein contents, and rubisco gene expression. Trees, 2015, 29(5): 1483-1492.