高温条件下稀土尾砂干旱对4种植物生理特性的影响
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摘要
稀土矿尾砂区地表大面积裸露,土壤结构遭到破坏,保水性能差,植物生长受到严重的干旱胁迫,季节性高温干旱期甚至威胁植物的生存。以稀土矿区周边广泛分布的枫香、木荷、樟树和桉树4种植物为研究对象,对比分析高温干旱条件下4种植物幼苗的光合特征、水分利用效率(WUE)、渗透调节物质和抗氧化酶活性等生理特征及适应性,为稀土尾砂矿区生态修复中植物的筛选提供理论依据。结果显示,高温干旱(空气温度40℃,土壤含水量2%左右)条件下,4种植物的光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和叶绿素(Chl)含量显著低于高温丰水条件(空气温度40℃,土壤含水量15%左右),光合作用受到抑制,WUE升高;不管高温丰水还是干旱条件下,樟树WUE最高;干旱条件下枫香和桉树的丙二醛(MDA),木荷和桉树的可溶性糖、脯氨酸(Pro)和可溶性蛋白,以及所有植物的过氧化物酶活性(超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT))均显著高于丰水条件。隶属函数评价得出4种植物幼苗抗高温干旱的等级为樟树>桉树>木荷>枫香;灰色关联分析发现WUE和可溶性糖与抗旱性关联最大,其次为CAT、Pro和MDA,可溶性蛋白与抗旱性关联最小。其中,樟树通过提高WUE、可溶性糖和CAT来应对干旱,桉树通过可溶性糖和MDA响应干旱胁迫,木荷在Pro调节下体现抗旱性,枫香则通过可溶性蛋白来抗旱。因此,在稀土尾砂植被恢复中关于乔木树种的选择可优先考虑樟树。
Mining rare earth minerals by heap leaching processes imposes substantial threats to the soil in rare earth ore areas, including the destruction of soil structures and the exposure of the ground surface over large areas, which together leads to poor soil water retention and severe soil erosion. Consequently, the growth and survival of plants in these areas are likely to be threatened by soil water deficit, especially during the drought season when air temperatures are frequently very high. To improve the ecological environments of rare earth mill tailings, effective restoration strategies are urgently needed, of which screening for suitable plants is a critical step for ecological recovery. In this study, four woody plant species(Liquidambar formosana, Schima superba, Eucalyptus grandis, and Cinnamomum camphora) that are widely distributed around rare earth mine tailings were selected and tested for their physiological responses to drought stress at high temperatures, and the photosynthetic traits, water use efficiency(WUE), osmotic regulation substances, and antioxidant enzymes activities of seedlings were measured. All four species grown under the drought treatment(with soil water content at 2% and air temperature at 40℃) had significantly lower photosynthetic rates(Pn), stomatal conductance(Gs), and transpiration rates(Tr) than those in the wet treatment(with soil water content at 15% and air temperature at 40℃), but there was a higher WUE for seedlings under the drought condition compared to seedlings in the wet treatment. C. camphora had the highest WUE regardless of soil water status. Malondialdehyde(MDA) in L. formosana and E. grandis, the soluble sugar, proline(Pro), the soluble protein in S. superba and E. grandis, and the peroxidase activity of all species under the drought condition were significantly higher than those in wet. The comprehensive evaluation on physiological indexes based on the subordinate function value method indicated that the sequence for drought resistance of the four plants at high temperatures was C. camphora, E. grandis, S. superba, and L. formosana. The gray correlation analysis showed that WUE and soluble sugar had the largest correlation with drought resistance, followed by CAT, Pro and MDA. On the other hand, soluble protein was least correlated with drought resistance. Overall, C. camphora coped with drought stress at high temperatures by improving WUE and accumulating soluble sugar and CAT jointly, while E. grandis coped with drought stress by accumulating soluble sugar and MDA. For the other two species, S. superba responded to drought through the proline regulation, while L. formosana regulated their soluble proteins in resistance to drought. Therefore, C. camphora could be a priority for vegetation restoration in rare earth mill tailings.
Mining rare earth minerals by heap leaching processes imposes substantial threats to the soil in rare earth ore areas, including the destruction of soil structures and the exposure of the ground surface over large areas, which together leads to poor soil water retention and severe soil erosion. Consequently, the growth and survival of plants in these areas are likely to be threatened by soil water deficit, especially during the drought season when air temperatures are frequently very high. To improve the ecological environments of rare earth mill tailings, effective restoration strategies are urgently needed, of which screening for suitable plants is a critical step for ecological recovery. In this study, four woody plant species(Liquidambar formosana, Schima superba, Eucalyptus grandis, and Cinnamomum camphora) that are widely distributed around rare earth mine tailings were selected and tested for their physiological responses to drought stress at high temperatures, and the photosynthetic traits, water use efficiency(WUE), osmotic regulation substances, and antioxidant enzymes activities of seedlings were measured. All four species grown under the drought treatment(with soil water content at 2% and air temperature at 40℃) had significantly lower photosynthetic rates(Pn), stomatal conductance(Gs), and transpiration rates(Tr) than those in the wet treatment(with soil water content at 15% and air temperature at 40℃), but there was a higher WUE for seedlings under the drought condition compared to seedlings in the wet treatment. C. camphora had the highest WUE regardless of soil water status. Malondialdehyde(MDA) in L. formosana and E. grandis, the soluble sugar, proline(Pro), the soluble protein in S. superba and E. grandis, and the peroxidase activity of all species under the drought condition were significantly higher than those in wet. The comprehensive evaluation on physiological indexes based on the subordinate function value method indicated that the sequence for drought resistance of the four plants at high temperatures was C. camphora, E. grandis, S. superba, and L. formosana. The gray correlation analysis showed that WUE and soluble sugar had the largest correlation with drought resistance, followed by CAT, Pro and MDA. On the other hand, soluble protein was least correlated with drought resistance. Overall, C. camphora coped with drought stress at high temperatures by improving WUE and accumulating soluble sugar and CAT jointly, while E. grandis coped with drought stress by accumulating soluble sugar and MDA. For the other two species, S. superba responded to drought through the proline regulation, while L. formosana regulated their soluble proteins in resistance to drought. Therefore, C. camphora could be a priority for vegetation restoration in rare earth mill tailings.
引文
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[6] 王友生,吴鹏飞,侯晓龙,岳辉,彭绍云,马祥庆.稀土矿废弃地不同植被恢复模式对土壤肥力的影响.生态环境学报,2015,24(11):1831- 1836.
[7] 裴斌,张光灿,张淑勇,吴芹,徐志强,徐萍.土壤干旱胁迫对沙棘叶片光合作用和抗氧化酶活性的影响.生态学报,2013,33(5):1386- 1396.
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[10] Sobrado M A.Relation of water transport to leaf gas exchange properties in three mangrove species.Trees,2000,14(5):258- 262.
[11] Gao Y,Markkanen T,Aurela M,Mammarella I,Thum T,Tsuruta A,Yang H Y,Aalto T.Response of water use efficiency to summer drought in boreal Scots pine forests in Finland.Biogeosciences Discussions,2016,doi:10.5194/bg- 2016- 198.
[12] 白灵娜,王占林,贺康宁,樊光辉,张得芳,谢守忠.青海4种抗旱树种水分利用效率对水分和光照的响应.中国农学通报,2016,32(16):12- 17.
[13] Chazdon R L,Pearcy R W,Lee D W,Fetcher N.Photosynthetic responses of tropical forest plants to contrasting light environments//Mulkey S S,Chazdon R L,Smith A P,eds.Tropical Forest Plant Ecophysiology.Boston,MA:Springer,1996:5- 55.
[14] 李子英,丛日春,杨庆山,周健.盐碱胁迫对柳树幼苗生长和渗透调节物质含量的影响.生态学报,2017,37(24):8511- 8517.
[15] 周瑞莲,解卫海,侯玉平,王艳芳,黄清荣.东北沙地7种植物高温时段的生理适应对策.林业科学,2014,50(6):74- 81.
[16] 郭安琪,周瑞莲,宋玉,马会雷.刈割后黑麦草生理保护作用对其补偿性生长的影响.生态学报,2018,38(10):3495- 3503.
[17] 李合生.植物生理生化实验原理和技术.北京:高等教育出版社,2000.
[18] Liu Y B,Xiao J F,Ju W M,Zhou Y L,Wang S Q,Wu X C.Water use efficiency of China′s terrestrial ecosystems and responses to drought.Scientific Reports,2015,5:13799.
[19] 杜明凤,丁贵杰,赵熙州.不同家系马尾松对持续干旱的响应及抗旱性.林业科学,2017,53(6):21- 29.
[20] 初庆刚,胡正海.中国樟科植物叶中油细胞和粘液细胞的比较解剖研究.植物分类学报,1999,37(6):529- 540.
[21] 王倩,关雪莲,胡增辉,卢存福,冷平生.3种景天植物叶片结构特征与抗寒性的关系.应用与环境生物学报,2013,19(2):280- 285.
[22] 周钰佩,刘慧霞,于成,林丽果,林选栋.硅对不同盐生境下高羊茅生物量及渗透调节物质含量的影响.生态学报,2017,37(16):5514- 5521.
[23] 叶龙华,黄香兰,薛立.干旱对树木叶片性状及抗旱生理的影响.世界林业研究,2014,27(1):29- 34.
[24] 张婷,曹扬,陈云明,刘国彬.生长季末期干旱胁迫对刺槐幼苗非结构性碳水化合物的影响.水土保持学报,2016,30(5):297- 304.
[25] 赵雪,张秀珍,牟洪香,孙晓,贺红月,杨可伟,单媛媛,李春友.干旱胁迫对不同种源文冠果幼苗水分生理特性及渗透调节物质的影响.东北林业大学学报,2017,45(6):17- 21.
[26] 马剑,刘贤德,孟好军,赵维俊,王荣新,王艺林,任小凤.水分胁迫对文冠果幼苗生长及生理特性的影响.干旱区资源与环境,2018,32(1):128- 132.
[27] 张露婷,吴江,梅丽,吴家胜.喜树种源耐盐能力评价及耐盐指标筛选.林业科学,2011,47(11):66- 72.
[28] 王卓敏,郑欣颖,薛立.樟树幼苗对干旱胁迫和种植密度的生理响应.生态学杂志,2017,36(6):1495- 1502.
[29] 宋爱琴,陈圣宾,李振基,汪秀芳.水分胁迫对生态恢复重要树种木荷与白楸幼苗的影响.厦门大学学报:自然科学版,2006,45(S1):109- 113.
[30] 段洪浪,吴建平,刘文飞,廖迎春,张海娜,樊后保.干旱胁迫下树木的碳水过程以及干旱死亡机理.林业科学,2015,51(11):113- 120.
[31] Fonti P,Tabakova M A,Kirdyanov A V,Bryukhanova M V,von Arx G.Variability of ray anatomy of Larix gmelinii along a forest productivity gradient in Siberia.Trees,2015,29(4):1165- 1175.
[32] Imahori Y,Takemura M,Bai J H.Chilling-induced oxidative stress and antioxidant responses in mume (Prunus mume) fruit during low temperature storage.Postharvest Biology and Technology,2008,49(1):54- 60.
[2] Liang T,Li K X,Wang L Q.State of rare earth elements in different environmental components in mining areas of China.Environmental Monitoring and Assessment,2014,186(3):1499- 1513.
[3] 刘文深,刘畅,王志威,滕文凯,汤叶涛,仇荣亮.离子型稀土矿尾砂地植被恢复障碍因子研究.土壤学报,2015,52(4):879- 887.
[4] 李小青,陈昌春,池明茹.几种植物在稀土尾砂基质上的抗旱力比较.中国科技信息,2012(7):96- 97.
[5] 钟敏,郭礼荣,贺强,何世林,曾庆远,徐祥,蓝军,朱钦娟.中草药在赣南稀土尾砂治理中的适应性研究.中国水土保持,2017(11):57- 60.
[6] 王友生,吴鹏飞,侯晓龙,岳辉,彭绍云,马祥庆.稀土矿废弃地不同植被恢复模式对土壤肥力的影响.生态环境学报,2015,24(11):1831- 1836.
[7] 裴斌,张光灿,张淑勇,吴芹,徐志强,徐萍.土壤干旱胁迫对沙棘叶片光合作用和抗氧化酶活性的影响.生态学报,2013,33(5):1386- 1396.
[8] Schierenbeck K A,Marshall J D.Seasonal and diurnal patterns of photosynthetic gas exchange for Lonicera sempervirens and L.Japonica (Caprifoliaceae).American Journal of Botany,1993,80(11):1292- 1299.
[9] Yang Y T,Guan H D,Batelaan O,McVicar T R,Long D,Piao S L,Liang W,Liu B,Jin Z,Simmons C T.Contrasting responses of water use efficiency to drought across global terrestrial ecosystems.Scientific Reports,2016,6:23284.
[10] Sobrado M A.Relation of water transport to leaf gas exchange properties in three mangrove species.Trees,2000,14(5):258- 262.
[11] Gao Y,Markkanen T,Aurela M,Mammarella I,Thum T,Tsuruta A,Yang H Y,Aalto T.Response of water use efficiency to summer drought in boreal Scots pine forests in Finland.Biogeosciences Discussions,2016,doi:10.5194/bg- 2016- 198.
[12] 白灵娜,王占林,贺康宁,樊光辉,张得芳,谢守忠.青海4种抗旱树种水分利用效率对水分和光照的响应.中国农学通报,2016,32(16):12- 17.
[13] Chazdon R L,Pearcy R W,Lee D W,Fetcher N.Photosynthetic responses of tropical forest plants to contrasting light environments//Mulkey S S,Chazdon R L,Smith A P,eds.Tropical Forest Plant Ecophysiology.Boston,MA:Springer,1996:5- 55.
[14] 李子英,丛日春,杨庆山,周健.盐碱胁迫对柳树幼苗生长和渗透调节物质含量的影响.生态学报,2017,37(24):8511- 8517.
[15] 周瑞莲,解卫海,侯玉平,王艳芳,黄清荣.东北沙地7种植物高温时段的生理适应对策.林业科学,2014,50(6):74- 81.
[16] 郭安琪,周瑞莲,宋玉,马会雷.刈割后黑麦草生理保护作用对其补偿性生长的影响.生态学报,2018,38(10):3495- 3503.
[17] 李合生.植物生理生化实验原理和技术.北京:高等教育出版社,2000.
[18] Liu Y B,Xiao J F,Ju W M,Zhou Y L,Wang S Q,Wu X C.Water use efficiency of China′s terrestrial ecosystems and responses to drought.Scientific Reports,2015,5:13799.
[19] 杜明凤,丁贵杰,赵熙州.不同家系马尾松对持续干旱的响应及抗旱性.林业科学,2017,53(6):21- 29.
[20] 初庆刚,胡正海.中国樟科植物叶中油细胞和粘液细胞的比较解剖研究.植物分类学报,1999,37(6):529- 540.
[21] 王倩,关雪莲,胡增辉,卢存福,冷平生.3种景天植物叶片结构特征与抗寒性的关系.应用与环境生物学报,2013,19(2):280- 285.
[22] 周钰佩,刘慧霞,于成,林丽果,林选栋.硅对不同盐生境下高羊茅生物量及渗透调节物质含量的影响.生态学报,2017,37(16):5514- 5521.
[23] 叶龙华,黄香兰,薛立.干旱对树木叶片性状及抗旱生理的影响.世界林业研究,2014,27(1):29- 34.
[24] 张婷,曹扬,陈云明,刘国彬.生长季末期干旱胁迫对刺槐幼苗非结构性碳水化合物的影响.水土保持学报,2016,30(5):297- 304.
[25] 赵雪,张秀珍,牟洪香,孙晓,贺红月,杨可伟,单媛媛,李春友.干旱胁迫对不同种源文冠果幼苗水分生理特性及渗透调节物质的影响.东北林业大学学报,2017,45(6):17- 21.
[26] 马剑,刘贤德,孟好军,赵维俊,王荣新,王艺林,任小凤.水分胁迫对文冠果幼苗生长及生理特性的影响.干旱区资源与环境,2018,32(1):128- 132.
[27] 张露婷,吴江,梅丽,吴家胜.喜树种源耐盐能力评价及耐盐指标筛选.林业科学,2011,47(11):66- 72.
[28] 王卓敏,郑欣颖,薛立.樟树幼苗对干旱胁迫和种植密度的生理响应.生态学杂志,2017,36(6):1495- 1502.
[29] 宋爱琴,陈圣宾,李振基,汪秀芳.水分胁迫对生态恢复重要树种木荷与白楸幼苗的影响.厦门大学学报:自然科学版,2006,45(S1):109- 113.
[30] 段洪浪,吴建平,刘文飞,廖迎春,张海娜,樊后保.干旱胁迫下树木的碳水过程以及干旱死亡机理.林业科学,2015,51(11):113- 120.
[31] Fonti P,Tabakova M A,Kirdyanov A V,Bryukhanova M V,von Arx G.Variability of ray anatomy of Larix gmelinii along a forest productivity gradient in Siberia.Trees,2015,29(4):1165- 1175.
[32] Imahori Y,Takemura M,Bai J H.Chilling-induced oxidative stress and antioxidant responses in mume (Prunus mume) fruit during low temperature storage.Postharvest Biology and Technology,2008,49(1):54- 60.
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