氮添加与干旱共同作用下的油松幼苗渗透调节机制

doi:10.11707/j.1001-7488.LYKX20240362

摘要/Abstract

摘要：

目的: 探究氮添加对黄土高原地区典型树种油松渗透调节物质含量的影响，揭示其通过维持水力功能缓解干旱胁迫的机制，为该地区油松人工林的经营与管理提供理论依据。方法: 以2年生油松幼苗为研究对象，设置4个干旱胁迫处理（对照CK、轻度胁迫RE₂₅、中度胁迫RE₅₀和重度胁迫RE₇₅）及3个氮添加水平（N₀、N₃、N₆质量浓度分别为0、3、6 g·m^–2a^–1），分析氮添加与干旱胁迫共同作用下油松幼苗的生理生长特性、渗透调节物质含量和水力功能变化及其之间的相关关系。结果: 1）在CK处理下，N₃和N₆水平的总生物量相比N₀水平显著增加3.69%和10.56%（P<0.05），净光合速率显著增加22.16%和79.61%（P<0.05）；在N₀水平下，RE₇₅处理与CK处理相比根冠比显著增加54.9%（P<0.05）。2）高水平施氮显著增加CK、轻度和中度干旱胁迫下新枝的脯氨酸含量；施氮显著增加各干旱处理的非结构性碳水化合物（NSC）含量，对钾离子含量没有显著影响。施氮对油松幼苗新枝的渗透调节作用影响大于干旱。3）相关分析表明，施氮条件下，可溶性糖和NSC总量与净光合速率、生物量、导水率、水分利用效率均呈显著正相关，表明施氮通过提高地上净光合速率增加可溶性糖含量和NSC形成，促进生物量积累，通过提升可溶性糖含量进行渗透调节作用以维持水力功能，进而提升油松水分利用效率。4）冗余分析表明，氮添加与干旱胁迫共同作用下对水力功能具有显著影响的渗透调节物质有脯氨酸（P<0.05）和可溶性糖（P<0.05），解释率分别为38.5%和12.9%。结论: 新枝是干旱胁迫影响下最脆弱的器官，相较3个施氮质量浓度，6 g·m^–2a^–1是缓解轻度干旱胁迫的最优氮添加水平。

关键词: 干旱胁迫, 氮添加, 水力功能, 油松, 渗透调节

Abstract:

Objective: This study explored the effect of nitrogen addition on the content of osmoregulatory substances in a typical tree species, Pinus tabuliformis, in the Loess Plateau region, for revealing the mechanism by which nitrogen addition alleviates drought stress by maintaining hydraulic function, and thereby exploring the response of forest ecosystems in the region to global change. This research aimed to provide a scientific basis for managing P. tabuliformis plantations in the Loess Plateau region. Method: Two-year-old P. tabuliformis seedlings were used as the research object, and four levels of drought stress treatments (control CK, mild stress RE₂₅, moderate stress RE₅₀, and severe stress RE₇₅) and three nitrogen addition levels (N₀, N₃, N₆ with concentrations of 0, 3, 6 g·m^–2a^–1, respectively) were set up to analyze the changes in physiological growth characteristics, osmoregulatory substances, and hydraulic function of the seedlings under the interactive effects of nitrogen addition and drought stress, along with their interrelationships. Result: 1） Under CK treatment, the total biomass at N₃ and N₆ levels significantly increased by 3.69% and 10.56% (P<0.05) compared to the N0 level, and the net photosynthetic rate increased by 22.16% and 79.61% (P<0.05). At the N₀ level, the root-to-shoot ratio under RE₇₅ treatment significantly increased by 54.9% (P<0.05) compared to the CK group. 2） Correlation analysis proved that high-concentration nitrogen application significantly increased the proline content in new branches under CK treatment, mild and moderate drought stress; nitrogen application significantly increased the content of non-structural carbohydrates under each drought treatment but had no significant effect on the content of potassium ions. The effect of nitrogen application on osmotic regulation of new branches of P. tabuliformis seedlings was greater than that of drought. 3） Under nitrogen application conditions, the total amount of soluble sugars and non-structural carbohydrates showed a significant positive correlation with net photosynthetic rate, biomass, hydraulic conductivity, and water use efficiency, indicating that nitrogen application increased the formation of soluble substances and non-structural carbohydrates by enhancing the net photosynthetic rate above ground, promoting biomass accumulation, and maintaining hydraulic function through osmotic regulation by soluble sugars, thereby improving the water use efficiency of P. tabuliformis. 4） Redundancy analysis showed that proline (P<0.05) and soluble sugar (P<0.05) had significant effects on hydraulic function under the coupled effects of nitrogen addition and drought stress, with an explanation rate of 38.5% and 12.9%, respectively. Conclusion: New branches are the most vulnerable organs under drought stress. Among nitrogen addition levels, the concentration of 6 g·m^?2a^?1 is the optimal nitrogen addition level for alleviating mild drought stress in P. tabuliformis seedlings.

Key words: drought stress, nitrogen addition, hydraulic conductivity, Pinus tabuliformis, osmotic regulation

中图分类号:

曾岩,冒吉荣,陈相霖,徐馨妤,梁静,刘莹. 氮添加与干旱共同作用下的油松幼苗渗透调节机制[J]. 林业科学, 2025, 61(8): 70-79.

Yan Zeng,Jirong Mao,Xianglin Chen,Xinyu Xu,Jing Liang,Ying Liu. Osmotic Regulation Mechanism of Pinus tabuliformis Seedlings under the Joint Effects of Nitrogen Addition and Drought Stress[J]. Scientia Silvae Sinicae, 2025, 61(8): 70-79.

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参考文献 0

	陈建勋, 王晓峰. 2002. 植物生理学实验指导. 广州: 华南理工大学出版社.
	Chen J X, Wang X F. 2002. Experimental instruction of plant physiology. Guangzhou: South China University of Technology Press. ［in Chinese］
	何如梦, 王百田, 于显威, 等. 晋西黄土区油松林的生长释放与生长抑制. 应用与环境生物学报, 2018, 24 (6): 1204- 1210.
	He R M, Wang B T, Yu X W, et al. Growth release and growth inhibition of Pinus tabuliformis forest in the Loess Plateau of western Shanxi Province, China. Chinese Journal of Applied and Environmental Biology, 2018, 24 (6): 1204- 1210.
	洪琮浩, 洪　震, 雷小华, 等. 氮添加对长序榆C、N、P养分含量及非结构性碳水化合物含量的影响. 林业科学, 2020, 56 (6): 186- 192.
	Hong C H, Hong Z, Lei X H, et al. Effects of nitrogen addition on contents of C, N and P nutrient and non-structural carbohydrate in Ulmus elongata. Scientia Silvae Sinicae, 2020, 56 (6): 186- 192.
	李合生. 2000. 植物生理生化实验原理和技术. 北京: 高等教育出版社.
	Li H S. 2000. Principles and techniques of plant physiological and biochemical experiment. Beijing: Higher Education Press. ［in Chinese］
	李　娜. 2014. 落叶松幼苗对干旱胁迫及氮添加的生理生态响应. 哈尔滨: 东北林业大学.
	Li N. 2014. Physiological and ecological responses of Larix gmelinii seedlings under soil drought stress and different nitrogen levels. Harbin: Northeast Forestry University. ［in Chinese］
	苏　炜, 陈　平, 吴婷, 等. 氮添加与干季延长对降香黄檀幼苗非结构性碳水化合物、养分与生物量的影响. 植物生态学报, 2023, 47 (8): 1094- 1104. doi: 10.17521/cjpe.2022.0473
	Su W, Chen P, Wu T, et al. Effects of nitrogen addition and extended dry season on non-structural carbohydrates, nutrients and biomass of Dalbergia odorifera seedlings. Chinese Journal of Plant Ecology, 2023, 47 (8): 1094- 1104. doi: 10.17521/cjpe.2022.0473
	张聪惠. 2022. 黄土高原典型草原植被群落和土壤微生物养分利用特征对降水和氮沉降的响应. 兰州: 兰州大学.
	Zhang C H. 2022. Responses of vegetation communities and soil microbial nutrient characteristics to precipitation and nitrogen deposition in typical steppe on the Loess Plateau. Lanzhou: Lanzhou University. ［in Chinese］
	张　宏, 曾　雄, 王爱莲, 等. 不同施氮量对棉花产量、养分吸收及氮素利用的影响. 新疆农业科学, 2021, 58 (9): 1656- 1664.
	Zhang H, Zeng X, Wang A L, et al. Effects of different nitrogen application rates on yield, nutrient uptake and nitrogen utilization of cotton in southern Xinjiang. Xinjiang Agricultural Sciences, 2021, 58 (9): 1656- 1664.
	Chen X, Zhao P, Ouyang L, et al. Whole-plant water hydraulic integrity to predict drought-induced Eucalyptus urophylla mortality under drought stress. Forest Ecology and Management, 2020, 468, 118179. doi: 10.1016/j.foreco.2020.118179
	Du D S, Jiao L, Wu X, et al. 2024. Drought determines the growth stability of different dominant conifer species in central Asia. Global and Planetary Change, 234: 104370.
	Dziedek C, von Oheimb G, Calvo L, et al. Does excess nitrogen supply increase the drought sensitivity of European beech (Fagus sylvatica L. ) seedlings?Plant Ecology, 2016, 217 (4): 393- 405.
	Eastman B A, Adams M B, Brzostek E R, et al. Altered plant carbon partitioning enhanced forest ecosystem carbon storage after 25 years of nitrogen additions. New Phytologist, 2021, 230 (4): 1435- 1448. doi: 10.1111/nph.17256
	Egilla J N, Davies F T, Boutton T W. Drought stress influences leaf water content, photosynthesis, and water-use efficiency of Hibiscus rosa-sinensis at three potassium concentrations. Photosynthetica, 2005, 43 (1): 135- 140. doi: 10.1007/s11099-005-5140-2
	Gersony J T, Hochberg U, Rockwell F E, et al. Leaf carbon export and nonstructural carbohydrates in relation to diurnal water dynamics in mature oak trees. Plant Physiology, 2020, 183 (4): 1612- 1621. doi: 10.1104/pp.20.00426
	Jaleel C A, Manivannan P, Sankar B, et al. Induction of drought stress tolerance by ketoconazole in Catharanthus roseus is mediated by enhanced antioxidant potentials and secondary metabolite accumulation. Colloids and Surfaces B: Biointerfaces, 2007, 60 (2): 201- 206. doi: 10.1016/j.colsurfb.2007.06.010
	Kishor P B K, Sangam S, Amrutha R N, et al. 2005. Regulation of proline biosynthesis, degradation, uptake and transport in higher plants:its implications in plant growth and abiotic stress tolerance. Current Science, 88(3): 424–438.
	Li W B, Hartmann H, Adams H D, et al. The sweet side of global change-dynamic responses of non-structural carbohydrates to drought, elevated CO₂, and nitrogen fertilization in tree species. Tree Physiology, 2018, 38 (11): 1706- 1723.
	Mahmood T, Abdullah M, Ahmar S, et al. Incredible role of osmotic adjustment in grain yield sustainability under water scarcity conditions in wheat (Triticum aestivum L). Plants, 2020, 9 (9): 1208. doi: 10.3390/plants9091208
	Ozturk M, Turkyilmaz Unal B, García-Caparrós P, et al. Osmoregulation and its actions during the drought stress in plants. Physiologia Plantarum, 2021, 172 (2): 1321- 1335. doi: 10.1111/ppl.13297
	Peng Y H, Chen K L, Wang G L, et al. Nitrogen addition regulates the growth of Pinus tabuliformis by changing distribution patterns of endogenous hormones in different organs. New Forests, 2023, 54 (5): 853- 865. doi: 10.1007/s11056-022-09947-5
	Savi T, Casolo V, Dal Borgo A, et al. Drought-induced dieback of Pinus nigra: a tale of hydraulic failure and carbon starvation. Conservation Physiology, 2019, 7 (1): coz012.
	Shi H L, Ma W J, Song J Y, et al. Physiological and transcriptional responses of Catalpa bungei to drought stress under sufficient-and deficient-nitrogen conditions. Tree Physiology, 2017, 37 (11): 1457- 1468. doi: 10.1093/treephys/tpx090
	Sigala J A, Uscola M, Oliet J A, et al. Drought tolerance and acclimation in Pinus ponderosa seedlings: the influence of nitrogen form. Tree Physiology, 2020, 40 (9): 1165- 1177. doi: 10.1093/treephys/tpaa052
	Sperry J S, Donnelly J R, Tyree M T. 1988. A method for measuring hydraulic conductivity and embolism in xylem. Plant, Cell and Environment, 11(1): 35–40.
	Sperry J S, Love D M. What plant hydraulics can tell us about responses to climate-change droughts. New Phytologist, 2015, 207 (1): 14- 27. doi: 10.1111/nph.13354
	Tomasella M, Petrussa E, Petruzzellis F, et al. The possible role of non-structural carbohydrates in the regulation of tree hydraulics. International Journal of Molecular Sciences, 2020, 21 (1): 144.
	Wang R Z, Yun L L, Mao Y X, et al. Nitrogen deposition alters drought-induced changes in biomass and nonstructural carbohydrates allocation patterns of Quercus mongolica seedlings. Scientia Horticulturae, 2024, 325, 112573. doi: 10.1016/j.scienta.2023.112573
	Wang X, Wu G Y, Li D Y, et al. Moderate nitrogen deposition alleviates drought stress of Bretschneidera sinensis. Forests, 2023, 14 (1): 137. doi: 10.3390/f14010137
	Zhang D, Jing H, Wang G L. Responses of non-structural carbohydrates content in leaves of different plant species in Pinus tabuliformis plantation to nitrogen addition. The Journal of Applied Ecology, 2019, 30 (2): 489- 495.
	Zuo K Y, Fan L L, Guo Z W, et al. Aboveground biomass component plasticity and allocation variations of bamboo (Pleioblastus amarus) of different regions. Forests, 2024, 15 (1): 43.

指标 Indicator	干旱胁迫处理 Drought stress treatments	氮添加水平 Nitrogen addition levels
指标 Indicator	干旱胁迫处理 Drought stress treatments	N₀	N₃	N₆
净光合速率 Net photosynthetic rate/(μmol?m^–2s^–1)	CK	5.64±0.13Bb	6.56±0.75Bab	10.22±0.04Aa
	RE₂₅	8.11±0.50Aab	10.05±0.07Ab	10.27±0.04Aa
	RE₅₀	5.33±0.01Bb	5.56±0.01BCb	6.77±1.06Ba
	RE₇₅	4.94±0.05Ba	5.30±0.04Ca	5.45±0.53Ba
水分利用效率 Water use efficiency/(μmol?mol^–1)	CK	8.63±0.22Bb	11.75±1.68Bb	16.28±0.07Aa
	RE₂₅	11.03±0.67Ab	15.99±0.13Aab	16.36±0.06Aa
	RE₅₀	10.88±0.01Aa	10.86±0.01Ba	11.63±1.41Ba
	RE₇₅	7.46±0.09Ca	8.06±0.06Ca	7.64±0.62Ca
叶面积 Leaf area/m²	CK	0.65±0.02Cc	0.81±0.01Cb	1.14±0.04Ca
	RE₂₅	0.75±0.03Bc	0.95±0.05Bb	1.52±0.04Aa
	RE₅₀	1.04±0.04Ab	1.09±0.03Ab	1.32±0.04Ba
	RE₇₅	0.57±0.03Cc	0.92±0.04BCb	1.21±0.03BCa
总生物量 Total biomass/g	CK	118.71±1.61Bb	123.09±3.47Ba	131.05±2.33Ba
	RE₂₅	116.82±2.88Bc	129.83±6.76Ab	156.41±2.07Aa
	RE₅₀	125.15±5.93Ab	126.01±6.99Ab	155.35±4.26Aa
	RE₇₅	90. 67±3.55Cc	115.95±1.48Bb	128.49±2.54Ca
根冠比 Root-shoot ratio/(g?g^–1)	CK	0.153±0.009Cb	0.167±0.013Ca	0.177±0.008Ba
	RE₂₅	0.160±0.005Bb	0.191±0.032Ba	0.174±0.008Bb
	RE₅₀	0.165±0.008Bb	0.171±0.006Ca	0.187±0.005Aa
	RE₇₅	0.237±0.008Aa	0.201±0.009Ab	0.233±0.019Aa

指标 Index	干旱胁迫处理 Drought stress treatment	氮添加水平 Nitrogen addition level	干旱胁迫处理×氮添加水平 Drought stress treatment × nitrogen addition level
总生物量Total biomass	65.66^**	88.93^**	13.52^**
净光合速率Net photosynthetic rate	146.60^**	43.24^**	14.78^**
水分利用效率Water use efficiency	69.98^**	29.79^**	15.65^**
导水率Hydraulic conductivity	71.07^**	232.60^**	65.70^**
可溶性糖Soluble sugars	83.06^**	154.50^**	11.35^**
淀粉Starch	5.24^**	152.30^**	5.67^**
非结构性碳水化合物Non-structural carbohydrates	42.68^**	491.00^**	6.43^**
脯氨酸Proline	5.89^**	231.50^**	20.56^**
K⁺	30.51^**	1.99^ns	5.75^**

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