NaCl胁迫对两种番茄气孔特征、气体交换参数和生物量的影响
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摘要
为探讨番茄叶片气孔特征、气体交换参数和生物量对盐胁迫的响应机理,以赛棚和阿拉姆番茄为试材,通过向水培营养液中添加NaCl(0.1 mol·L~(-1)),在人工气候箱条件下进行为期90 d的NaCl盐胁迫处理.结果表明:NaCl胁迫导致赛棚番茄叶片的气孔密度、气孔宽度、气孔面积和气孔面积指数显著降低,降幅分别为32%、45%、25%、49%,但没有改变阿拉姆番茄叶片的气孔特征参数.同时,NaCl胁迫还导致赛棚和阿拉姆叶片气孔规则分布的空间尺度分别减少30%和43%,且赛棚品种的单个气孔最小邻域距离在盐胁迫下增加20%.另外,赛棚和阿拉姆叶片的净光合速率(P_n)、气孔导度(g_s)和蒸腾速率(T_r)在盐胁迫下均显著下降,通过气孔限制值分析发现,盐胁迫导致赛棚番茄叶片光合速率下降主要是由气孔限制因素引起的,而阿拉姆叶片则以非气孔限制因素为主导作用.盐胁迫还导致赛棚和阿拉姆番茄生物量显著降低,且地下生物量的下降幅度大于地上生物量.综合分析表明,阿拉姆的抗盐能力高于赛棚.
To understand the mechanism underlying responses of stomatal traits,gas exchange parameters,and biomass of tomato plants to salt stress,two tomato cultivars( Shed and Alam) were treated by salt stress by adding NaCl( 0.1 mol·L~(-1)) to nutrition medium in environmental growth chambers for 90 days. Our results showed that salt stress substantially decreased the stomatal density,stomatal width,stomatal area,and stomatal area index of Shed by 32%,45%,25%,and49%,respectively. The stomatal traits of Alam were not affected by NaCl treatment. The spatial scales of the regular stomatal distribution pattern of Shed and Alam were significantly decreased by30% and 43%,and the nearest neighbor distance Lhat( d) of shed cultivar was increased by 20%under salt stress. NaCl stress resulted in marginal declines in the net photosynthetic rates( P_n),stomatal conductance( g_s),transpiration rates( T_r) of both cultivars. The decrease of the photosynthetic rate of Shed under salt stress resulted from stomatal limitation,whereas the Pnof Alam was subjected to non-stomatal constraints. NaCl stress substantially decreased the seedling biomass of both cultivars,and the decline of belowground biomass was higher than that of aboveground biomass. Overall,our results suggested that the Alam cultivar is more salt-tolerant than Shed.
To understand the mechanism underlying responses of stomatal traits,gas exchange parameters,and biomass of tomato plants to salt stress,two tomato cultivars( Shed and Alam) were treated by salt stress by adding NaCl( 0.1 mol·L~(-1)) to nutrition medium in environmental growth chambers for 90 days. Our results showed that salt stress substantially decreased the stomatal density,stomatal width,stomatal area,and stomatal area index of Shed by 32%,45%,25%,and49%,respectively. The stomatal traits of Alam were not affected by NaCl treatment. The spatial scales of the regular stomatal distribution pattern of Shed and Alam were significantly decreased by30% and 43%,and the nearest neighbor distance Lhat( d) of shed cultivar was increased by 20%under salt stress. NaCl stress resulted in marginal declines in the net photosynthetic rates( P_n),stomatal conductance( g_s),transpiration rates( T_r) of both cultivars. The decrease of the photosynthetic rate of Shed under salt stress resulted from stomatal limitation,whereas the Pnof Alam was subjected to non-stomatal constraints. NaCl stress substantially decreased the seedling biomass of both cultivars,and the decline of belowground biomass was higher than that of aboveground biomass. Overall,our results suggested that the Alam cultivar is more salt-tolerant than Shed.
引文
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[12] Li S-C(李善春). Study on the Physiological Characteristic of Five Dwarf Ornamental Bamboos under Na Cl Stress. Master Thesis. Nanjing:Nanjing Forestry University,2005(in Chinese)
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[14] Pinela J,Barros L,Carvalho AM,et al. Nutritional composition and antioxidant activity of four tomato(Lycopersicones culentum L.)farmer’s varieties in northeastern Portugal home gardens. Food and Chemical Toxicology,2012,50:829-834
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[20] Wang J,Duan B,Zhang Y. Effects of experimental warming on growth,biomass allocation,and needle chemistry of Abies faxoniana in even-aged monospecific stands.Plant Ecology,2012,213:47-55
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[25] Zhang X-X(张晓晓),Yin X-L(殷小琳),Li H-L(李红丽),et al. Effect of salt stress on the biomass and photosynthetic characteristics of Ulmus pumila L.strains. Acta Ecologica Sinica(生态学报),2017,37(21):1-8(in Chinese)
[26] Robredo A,Pérez-López U,Hector Sainz de la Maza,et al. Elevated CO2alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis.Environmental and Experimental Botany,2007,59:252-263
[27] Wang B-X(王碧霞),Li Y-X(黎云祥),Ding C-B(丁春邦). Physiological and biochemical responses of Humulus scandens seedlings to Na Cl stress. Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2017,37(2):321-329(in Chinese)
[28] Yue J-M(岳健敏),Zhang J-C(张金池),You Y-H(尤焱煌),et al. Effects of brassionstreoids on photosynthesis and ultrastructure of chloroplasts in Robinia pseudoacacis seedling under salt stress. Journal of Northwest A&F University(Natural Science)(西北农林科技大学学报:自然科学版),2017,45(10):56-66(in Chinese)
[29] Feng T(冯涛),Liu J-F(刘纪凤). Application of chlorophyll fluorescence analysis in plant photoinhibition research. Modern Agricultural Sciences and Technology(现代农业科技),2013(6):148-155(in Chinese)
[30] Zhang S-R(张守仁). Discussion on chlorophyll fluorescence kinetics parameters and their significance. Chinese Bulletin of Botany(植物学报),1999,16(4):444-448(in Chinese)
[31] He Y-H(何炎红),Guo L-S(郭连生),Tian Y-L(田有亮). Photosynthetic rates and chlorophyll fluorescence of Nitraria tangutorum at different leaf water potentials.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2005,25(11):2226-2233(in Chinese)
[32] Shao G-C(邵光成),Guo R-Q(郭瑞琪),Liu N(刘娜). Effects of different irrigation patterns on chlorophyll fluorescence parameters of hot pepper in Southern China.Transactions of the Chinese Society of Agricultural Engineering(农业工程学报),2011,27(9):226-230(in Chinese)
[33] Cui X-G(崔兴国). Effects of salt stress on the chlorophyll content and photosynthetic properties of Trifolum repnes Linn. Heilongjiang Agricultural Sciences(黑龙江农业科学),2011(4):78-79(in Chinese)
[34] Gu Y-F(谷艳芳),Ding S-Y(丁圣彦),Li T-T(李婷婷),et al. Effects of saline stress on dry matter partitioning and ecophysiological characteristics of winter wheat seedlings. Acta Ecologica Sinica(生态学报),2009,29(2):840-845(in Chinese)
[35] Chapin III FS, Bloom AJ, Field CB, et al. Plant response to multiple environmental factors. Biological Science,1987,37:49-57
[36] Osone Y,Tateno M. Applicability and limitation of optimal biomass allocation models:A test of two species from fertile and infertile habitats. Annals of Botany,2005,95:1211-1220
[37] Mauchamp A,Mésleard F. Salt tolerance in Phragmites australis populations from coastal mediterranean marshes. Aquatic Botany,2001,70:39-52
[2] Yang J-S(杨劲松). Development and prospect of the research on salt-affected soils in China. Acta Pedologica Sinica(土壤学报),2008,45(5):835-837(in Chinese)
[3] Jaleel CA,Lakshmanan GMA,Gomathinyagam M,et al.Triadimefon induced salt stress tolerance in Withania somnifera and its relationship to antioxidant defense system. South African Journal of Botany,2008,74:126-132
[4] Wang D-M(王东明),Jia Y(贾媛),Cui J-Z(崔继哲). Advances in research on effects of salt stress on plant and adaptive mechanism of the plant to salinity.Chinese Agricultural Science Bulletin(中国农业通讯),2009,25(4):124-128(in Chinese)
[5] Lawlor DW,Cornic G. Photosynthetic carbon assimilation and associated metabolism in relation to water deficits in higher plants. Plant, Cell and Environment,2002,25:275-294
[6] Li H-B(李海波),Chen W-F(陈温福),Li Q-Y(李全英). Responses of rice leaf photosynthetic parameters to light intensity under Na Cl stress. Chinese Journal of Applied Ecology(应用生态学报),2006,17(9):1588-1592(in Chinese)
[7] Peng F-R(彭方仁),Zhu Z-X(朱振贤),Tan P-P(谭鹏鹏),et al. Effect of Na Cl stress on chlorophyll fluorescence parameter in leaf of five tree species seedling. Journal of Plant Resources and Environment(植物资源与环境学报),2010,19(3):42-47(in Chinese)
[8] Wani AS,Ahmad A,Hayat S,et al. Salt-induced modulation in growth,photosynthesis and antioxidant system in two varieties of Brassica juncea. Saudi Journal of Biological Sciences,2013,20:183-193
[9] Shawkat M,Nasir M,Chen QJ,et al. Effect of salt stress on photosynthetic gas exchange and chlorophyll fluorescence parameters in Alhagi pseudalhagi. Agricultural Science&Technology,2017,18:411-416,423
[10] Korn RW. Evidence in dicots for stomatal patterning by inhibition. International Journal of Plant Science,1993,154:367-377
[11] Xu M. The optimal atmospheric CO2concentration for the growth of winter wheat(Triticum aestivum). Journal of Plant Physiology,2015,184:89-97
[12] Li S-C(李善春). Study on the Physiological Characteristic of Five Dwarf Ornamental Bamboos under Na Cl Stress. Master Thesis. Nanjing:Nanjing Forestry University,2005(in Chinese)
[13] Yang S(杨升). Study on Physiological Mechanism for Different Salt Tolerance of Elaeagnus angustifolia Seedings from Difference Provenances under Na Cl Stress.PhD Thesis. Beijing:Chinese Academy of Fore-stry,2014(in Chinese)
[14] Pinela J,Barros L,Carvalho AM,et al. Nutritional composition and antioxidant activity of four tomato(Lycopersicones culentum L.)farmer’s varieties in northeastern Portugal home gardens. Food and Chemical Toxicology,2012,50:829-834
[15] Zhang J-T(张纪涛),Xu M(徐猛),Han K(韩坤),et al. Effect of salt stress on plant nutrition and physiology of tomato seedlings. Acta Agriculturae Borealioccidentalis Sinica(西北农业学报),2011,20(2):128-133(in Chinese)
[16] Zheng YP,Xu M,Hou RX,et al. Effects of experimental warming on stomatal traits in leaves of maize(Zea may L.). Ecology and Evolution,2013,3:3095-3111
[17] Mokhtar R,Mohamed VA,Arbi G,et al. Effect of Na Cl on the growth and the ionic balance K+/Na+of two populations of Lotus criticus. Lotus Newsletter,2006,36:34-53
[18] Casson S,Gray JE. Influence of environmental factors on stomatal development. New Phytologist,2008,178:9-23
[19] Ren A-X(任安祥),Wang Y-M(王羽梅). Effects of salt stress on stomatal differentiation and movements of amaranth(Amaranthus tricolor L.)leaves. Acta Horticulturae Sinica(园艺学报),2010,37(3):479-484(in Chinese)
[20] Wang J,Duan B,Zhang Y. Effects of experimental warming on growth,biomass allocation,and needle chemistry of Abies faxoniana in even-aged monospecific stands.Plant Ecology,2012,213:47-55
[21] Apple ME,Olszyk DM,Ormrod DP,et al. Morphology and stomatal function of Douglas fir needles exposed to climate change:Elevated CO2and temperature. International Journal of Plant Science,2000,161:127-132
[22] Han C,Liu Q,Yang Y. Short-term effects of experimental warming and enhanced ultraviolet-B radiation on photosynthesis and antioxidant defense of Picea asperata seedlings. Plant Growth&Regulation,2009,58:153-162
[23] Hugh JE. Stomatal and non-stomatal restrictions to carbon assimilation in soybean(Glycine max)lines differing in water use efficiency. Environmental and Experimental Botany,2002,48:237-246
[24] Farquhar GD,Sharkey TD. Stomatal conductance and photosynthesis. Annual Review of Plant Physiology,1982,33:317-345
[25] Zhang X-X(张晓晓),Yin X-L(殷小琳),Li H-L(李红丽),et al. Effect of salt stress on the biomass and photosynthetic characteristics of Ulmus pumila L.strains. Acta Ecologica Sinica(生态学报),2017,37(21):1-8(in Chinese)
[26] Robredo A,Pérez-López U,Hector Sainz de la Maza,et al. Elevated CO2alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis.Environmental and Experimental Botany,2007,59:252-263
[27] Wang B-X(王碧霞),Li Y-X(黎云祥),Ding C-B(丁春邦). Physiological and biochemical responses of Humulus scandens seedlings to Na Cl stress. Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2017,37(2):321-329(in Chinese)
[28] Yue J-M(岳健敏),Zhang J-C(张金池),You Y-H(尤焱煌),et al. Effects of brassionstreoids on photosynthesis and ultrastructure of chloroplasts in Robinia pseudoacacis seedling under salt stress. Journal of Northwest A&F University(Natural Science)(西北农林科技大学学报:自然科学版),2017,45(10):56-66(in Chinese)
[29] Feng T(冯涛),Liu J-F(刘纪凤). Application of chlorophyll fluorescence analysis in plant photoinhibition research. Modern Agricultural Sciences and Technology(现代农业科技),2013(6):148-155(in Chinese)
[30] Zhang S-R(张守仁). Discussion on chlorophyll fluorescence kinetics parameters and their significance. Chinese Bulletin of Botany(植物学报),1999,16(4):444-448(in Chinese)
[31] He Y-H(何炎红),Guo L-S(郭连生),Tian Y-L(田有亮). Photosynthetic rates and chlorophyll fluorescence of Nitraria tangutorum at different leaf water potentials.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2005,25(11):2226-2233(in Chinese)
[32] Shao G-C(邵光成),Guo R-Q(郭瑞琪),Liu N(刘娜). Effects of different irrigation patterns on chlorophyll fluorescence parameters of hot pepper in Southern China.Transactions of the Chinese Society of Agricultural Engineering(农业工程学报),2011,27(9):226-230(in Chinese)
[33] Cui X-G(崔兴国). Effects of salt stress on the chlorophyll content and photosynthetic properties of Trifolum repnes Linn. Heilongjiang Agricultural Sciences(黑龙江农业科学),2011(4):78-79(in Chinese)
[34] Gu Y-F(谷艳芳),Ding S-Y(丁圣彦),Li T-T(李婷婷),et al. Effects of saline stress on dry matter partitioning and ecophysiological characteristics of winter wheat seedlings. Acta Ecologica Sinica(生态学报),2009,29(2):840-845(in Chinese)
[35] Chapin III FS, Bloom AJ, Field CB, et al. Plant response to multiple environmental factors. Biological Science,1987,37:49-57
[36] Osone Y,Tateno M. Applicability and limitation of optimal biomass allocation models:A test of two species from fertile and infertile habitats. Annals of Botany,2005,95:1211-1220
[37] Mauchamp A,Mésleard F. Salt tolerance in Phragmites australis populations from coastal mediterranean marshes. Aquatic Botany,2001,70:39-52