-
森林凋落物是森林生态系统内由植物组分产生并归还到林地表面,作为分解者的物质和能量来源以维持森林生态系统土壤养分的有机物质的总称[1-2],是森林土壤养分的主要补给者。凋落物分解是释放矿质元素供植物营养的重要生态过程[3],是陆地生态系统中光合产物转换和矿质养分循环的重要环节[4-5],一直是森林生态和环境的研究热点。
近几十年来,由于经济发展和人口膨胀,使矿物燃料燃烧、农业氮肥生产和使用等含氮化合物的排放日益增加,大气N沉降量增加已成为全球性的生态环境问题[6]。过量的N沉降可造成森林群落组成、凋落物化学成分、土壤微生物生物量以及酶活性发生变化等一系列生态问题,影响了凋落物的分解过程[7]。目前,国内外众多模拟N沉降的试验发现,外源性N添加对凋落物的影响出现为促进[8]、抑制[9]和无影响[10]3种结果,如陈翔等[11]对大兴安岭兴安落叶松(Larix gmelinii(Rupr.) Kuzen.)林的凋落物施N肥及汪金松等[12]对暖温带油松(Pinus tabuliformis Carr.)林的凋落物施N肥,提高了凋落物的分解速率;涂利华等[13]发现,模拟N沉降显著抑制了华西雨屏区亮叶桦(Betula luminifera H. Winkl.)凋落叶的分解;Mo等[14]发现,在凋落叶分解初期,N添加对鼎湖山木荷(Schima superba Gardn. et Champ.)林凋落物的分解速率影响不大。
有研究表明,生态系统中N沉降增加会加速土壤磷循环并导致P限制[15],并随着N沉降量的持续增加,P限制性逐渐增强[16]。随着人工速生丰产林的发展,在P缺乏的南方红壤森林中普遍使用含磷化肥[17],有利于微生物的繁衍,从而促进了土壤有机质和凋落物的分解[18-19]。Qualls等[20]和Liu等[21]研究发现,P添加可以提高凋落物中的P含量,增加土壤微生物生物量,促进凋落物的分解。DeBusk等[22]在大克拉莎(Cladium jamaicense Crantz)和宽叶香蒲(Typha latifolia Linn.)的凋落物分梯度施P肥的试验中发现,凋落物分解速率与施加的P肥浓度成正相关。弓晓静等[17]发现,N、P单独添加和复合添加均促进了湿地松(Pinus elliottii)凋落叶前期的分解速率,抑制中、后期的分解速率。Kozovits等[23]对巴西中部5个树种的研究中发现,单独添加P肥对凋落物的分解速率影响不大,而施N、P复合肥后,凋落物质量比对照减少了42%。因此,笔者推测N沉降和P富集的共同作用可能会影响森林生态系统中凋落物的分解动态,其内在影响机制有待继续深入研究。
马占相思(Acacia mangium Will.)是豆科速生常绿乔木树种,有较多的枯枝落叶养分回归土壤,是我国南方山地绿化的主要树种之一。前人已对马占相思凋落物动态[24]和水源涵养功能[25]进行了研究,而外源性N和P对马占相思凋落物分解的影响尚未见报道。本研究以广东省云勇林场马占相思林凋落叶为试验材料,采用尼龙网袋分解法,通过2 a的模拟外源性N和P添加试验,探索外源性N和P对凋落叶分解的影响过程与机制,以期为马占相思的土壤养分管理提供参考。
-
云勇林场(112°40′ E,22°53′ N)位于广东省佛山市高明区内,是佛山市唯一的国有林场,森林面积1 928.73 hm2。试验地属于亚热带季风气候,气候温和,年平均气温、最高气温和最低气温分别为22、34.5、3.5℃,无霜期长达360 d。试验地雨量充沛,年降水量平均2 000 mm,集中在4—8月,年平均相对湿度80%。地势属丘陵地带,土壤为花岗岩发育的酸性赤红壤(pH < 5),土层深厚而肥沃。试验地概况见表 1。
表 1 马占相思林概况
Table 1. General characteristics of the experimental stand
林分Stand 坡向Aspect 坡度Slope/(°) 平均胸径Mean diameter in breast height/cm 平均树高Mean tree height /m 冠幅Crown diameter/m 郁闭度Canopy density 主要林下植被Main undergrowth 马占相思Acacia mangium 30° SW 35 12.8 13.6 3 0.85 九节、鸭脚木、梅叶冬青、漫山秀竹、山芝麻、鬼灯笼、海金沙、黑面神、野牡丹、菝葜、白花酸果藤、芒萁、乌毛蕨、铁线蕨 注:九节Psychotria rubra(Lour.) Poir.、鸭脚木Schefflera octophylla(Lour.) Harms、梅叶冬青Ilex asprella(Hook. et Arn.) Champ. ex Benth.、漫山秀竹Microstegium vagans(Nees ex Steud.) A. Camus、山芝麻Heliicteres angustifolia Linn.、鬼灯笼Clerodendrum fortunatum L.、海金沙Lygodium japonicum(Thunb.) Sw.、黑面神Breynia fruticosa(Linn.) Hook. f.、野牡丹Melastoma candidum D. Don、菝葜Smilax china L.、白花酸果藤Embelia ribes Burm. f.、芒萁Dicranopteris dichotoma (Thunb.) Berhn.、乌毛蕨Blechnum orientale L.、铁线蕨Adiantum capillus-veneris L.。 -
图 1表明:不同处理的马占相思凋落叶残留量在前6个月迅速下降后平稳减少;处理6个月时,对照、施N、P和N+P的凋落叶分别分解了40%、65%、74%和61%;处理24个月后,凋落叶残留量的顺序为对照(0.90 g)>施N(0.82 g)>施N+P(0.52 g)>施P(0.08 g)。总体来看,施N、P和N+P处理对马占相思凋落叶分解均有一定的促进作用。
-
图 2表明,马占相思林地对照的凋落叶N含量整体呈小幅波动,施P的呈降-升后呈小幅波动,施N和N+P的呈升-降后呈小幅波动;处理24个月时,凋落叶N含量表现为施N(24.54 g·kg-1)>对照(22.73 g·kg-1)>施N+P(22.31 g·kg-1)>施P(21.48 g·kg-1)。试验末期,对照、施N和施N+P处理的凋落叶N含量均大于凋落叶初始的N含量(P < 0.05)。
图 2 马占相思林地凋落叶中的N含量变化
Figure 2. Change of N concentration during leaf litter decomposition in the A. mangium woodland
图 3表明,马占相思林地对照和施N的凋落叶P含量呈小幅波动,施P的小幅波动后大幅上升,施N+P的呈波动性上升;处理24个月时,凋落叶的P含量表现为施P(1.47 g·kg-1)>施N+P(1.05 g·kg-1)>施N(0.51 g·kg-1)>对照(0.47 g·kg-1)。试验末期,对照和施N处理的凋落叶P含量显著小于其初始的P含量,施P和施N+P处理的凋落叶P含量显著大于对照和施N处理(P < 0.05)。
图 3 马占相思林地凋落叶中的P含量变化
Figure 3. Change of P concentration during leaf litter decomposition in the A. mangium woodland
图 4表明:马占相思林地4种处理的凋落叶的K含量均大幅波动;对照、施N、施P和N+P处理的凋落叶K含量分别在分解第12个月、第9个月、第15个月和第6个月时达到最大值;处理24个月时,凋落叶的K含量表现为对照(3.22 g·kg-1)>施P(2.63 g·kg-1)>施N+P(2.22 g·kg-1)>施N(1.95 g·kg-1)。试验末期,各处理凋落叶的K含量均大于凋落叶初始的K含量(P < 0.05)。
-
表 2表明,施N、P和N+P处理的马占相思林地土壤的有机质、全N、有效P含量均显著大于对照(P < 0.05),施N和N+P处理的林地土壤有效N含量显著大于对照(P < 0.05),而施N处理的却减少了土壤中的全P、全K和有效K含量。表 3表明,施N、P和N+P处理均显著提高了马占相思林地土壤的酶活性。
表 2 马占相思林地土壤化学性质
Table 2. Soil chemical properties of A. mangium woodland
处理Treatment 有机质Organic matter/(g·kg-1) 全氮Total nitrogen/(g·kg-1) 全磷Total phosphorus/ (g·kg-1) 全钾Total potassium/(g·kg-1) 有效氮Available nitrogen/(mg·kg-1) 有效磷Available phosphorus/(mg·kg-1) 有效钾Available potassium/(mg·kg-1) CK 15.17±0.28c 0.80±0.02c 0.17±0.01ab 34.45±0.64a 77.83±0.89b 0.59±0.05d 67.34±0.58c 施N N treatment 18.19±0.25b 0.87±0.01b 0.15±0.00b 33.85±0.30a 114.81±3.39a 0.70±0.03c 63.78±0.36d 施P N treatment 19.59±0.36a 0.88±0.01b 0.19±0.01a 34.08±0.33a 109.14±0.90ab 2.34±0.05b 77.49±0.44b 施N+P N+P treatment 19.99±0.16a 1.06±0.04a 0.18±0.01a 33.29±0.08a 119.28±1.36a 2.67±0.02a 84.19±0.78a 表 3 马占相思林地土壤酶活性
Table 3. Soil enzyme activities of A. mangium woodland
处理Treatment 脲酶Urease NH4+-N/(mg·kg-1) 磷酸酶Acid phosphatase P5O2/(mg·kg-1) 过氧化氢酶CatalaseKMnO4/(mL·g-1) 对照CK 156.03±3.35c 261.97±1.58d 1.08±0.02c 施N N treatment 183.93±2.25b 273.33±2.62c 1.29±0.02b 施P N treatment 208.87±1.72a 292.90±1.58b 1.32±0.01b 施N+P N+P treatment 179.63±2.25b 308.80±1.57a 1.53±0.02a
模拟外源性氮磷对马占相思凋落叶分解及土壤生化特性的影响
Effects of Nitrogen and Phosphorus Additions on Leaf Litter Decomposition and Soil Biochemical Characteristics in an Acacia mangium Plantation
-
摘要:
目的 研究外源性氮和磷对马占相思凋落叶的分解速率、分解过程中N、P、K含量和土壤生化特性的影响, 以便为森林土壤养分管理提供参考。 方法 以广东省云勇林场马占相思林下凋落叶为试验材料, 采用尼龙网袋分解法, 设置对照(CK)、施N(10 g·m-2)、施P(5 g·m-2)、施N+P(N 10 g·m-2+P 5 g·m-2)4种处理, 每隔3个月取样1次, 并测定凋落叶残留量和N、P、K含量。 结果 表明:施N、P和N+P处理对马占相思凋落叶的分解均为促进作用。各处理马占相思凋落叶的N含量在分解过程中大致保持稳定, 施P和N+P处理的凋落叶P含量在分解过程中总体呈波动性上升, 而各处理的凋落叶K含量变化规律不明显。施N、P和N+P处理提高了马占相思林土壤的有机质和全N含量, 促进脲酶、磷酸酶及过氧化氢酶的活性。 结论 施N、P和N+P处理促进了马占相思凋落叶的分解, 有利于马占相思林的养分循环。 Abstract:Objective To study the effects of nitrogen and phosphorus additions on leaf litter decomposition, nutrient dynamics and soil biochemical characteristics during the decomposition process in a Acacia mangium plantation in order to understand the mechanism of influence of nitrogen and phosphorus on leaf litter decomposition of A. mangium and forest soil nutrient management. Method Leaf litter decomposition of the A. mangium plantation in Yunyong Forest Farm was investigated using litter bag method. N and P additions were designed in four treatments:the control, N addition (N 10 g·m-2), P addition (P 5g·m-2), and N+P addition (N 10g·m-2+P 5g·m-2). It was sampled in a three-month interval and then the remaining leaf litter and N, P, K contents were analyzed. Result The N, P and N+P additions improved leaf litter decomposition. The N content of leaf litter with the four treatments remained stable. The P content of leaf litter with P and N+P additions tended to increase and were greater than their initial value during litter decomposition process. The K content of leaf litter with the four treatments changed irregularly. The N, P and N+P additions significantly increased soil organic matter and total N contents and improved activities of soil urease, acid phosphatase and catalase. Conclusion N, P and N+P additions accelerated leaf litter decomposition of the A. mangium, which improved the nutrient cycling of A. mangium plantation. -
表 1 马占相思林概况
Table 1. General characteristics of the experimental stand
林分Stand 坡向Aspect 坡度Slope/(°) 平均胸径Mean diameter in breast height/cm 平均树高Mean tree height /m 冠幅Crown diameter/m 郁闭度Canopy density 主要林下植被Main undergrowth 马占相思Acacia mangium 30° SW 35 12.8 13.6 3 0.85 九节、鸭脚木、梅叶冬青、漫山秀竹、山芝麻、鬼灯笼、海金沙、黑面神、野牡丹、菝葜、白花酸果藤、芒萁、乌毛蕨、铁线蕨 注:九节Psychotria rubra(Lour.) Poir.、鸭脚木Schefflera octophylla(Lour.) Harms、梅叶冬青Ilex asprella(Hook. et Arn.) Champ. ex Benth.、漫山秀竹Microstegium vagans(Nees ex Steud.) A. Camus、山芝麻Heliicteres angustifolia Linn.、鬼灯笼Clerodendrum fortunatum L.、海金沙Lygodium japonicum(Thunb.) Sw.、黑面神Breynia fruticosa(Linn.) Hook. f.、野牡丹Melastoma candidum D. Don、菝葜Smilax china L.、白花酸果藤Embelia ribes Burm. f.、芒萁Dicranopteris dichotoma (Thunb.) Berhn.、乌毛蕨Blechnum orientale L.、铁线蕨Adiantum capillus-veneris L.。 表 2 马占相思林地土壤化学性质
Table 2. Soil chemical properties of A. mangium woodland
处理Treatment 有机质Organic matter/(g·kg-1) 全氮Total nitrogen/(g·kg-1) 全磷Total phosphorus/ (g·kg-1) 全钾Total potassium/(g·kg-1) 有效氮Available nitrogen/(mg·kg-1) 有效磷Available phosphorus/(mg·kg-1) 有效钾Available potassium/(mg·kg-1) CK 15.17±0.28c 0.80±0.02c 0.17±0.01ab 34.45±0.64a 77.83±0.89b 0.59±0.05d 67.34±0.58c 施N N treatment 18.19±0.25b 0.87±0.01b 0.15±0.00b 33.85±0.30a 114.81±3.39a 0.70±0.03c 63.78±0.36d 施P N treatment 19.59±0.36a 0.88±0.01b 0.19±0.01a 34.08±0.33a 109.14±0.90ab 2.34±0.05b 77.49±0.44b 施N+P N+P treatment 19.99±0.16a 1.06±0.04a 0.18±0.01a 33.29±0.08a 119.28±1.36a 2.67±0.02a 84.19±0.78a 表 3 马占相思林地土壤酶活性
Table 3. Soil enzyme activities of A. mangium woodland
处理Treatment 脲酶Urease NH4+-N/(mg·kg-1) 磷酸酶Acid phosphatase P5O2/(mg·kg-1) 过氧化氢酶CatalaseKMnO4/(mL·g-1) 对照CK 156.03±3.35c 261.97±1.58d 1.08±0.02c 施N N treatment 183.93±2.25b 273.33±2.62c 1.29±0.02b 施P N treatment 208.87±1.72a 292.90±1.58b 1.32±0.01b 施N+P N+P treatment 179.63±2.25b 308.80±1.57a 1.53±0.02a -
[1] 王相娥, 薛立, 谢腾芳.凋落物分解研究综述[J].土壤通报, 2009, 40(6):1473-1478. [2] Maisto G, De M A, Meola A, et al. Nutrient dynamics in litter mixtures of four Mediterranean maquis species decomposing in situ[J]. Soil Biology and Biochemistry, 2011, 43(3):520-530. doi: 10.1016/j.soilbio.2010.11.017 [3] 陆耀东, 薛立, 曹鹤, 等.去除地面枯落物对加勒比松林土壤特性的影响[J].生态学报, 2008, 28(7):3205-3211. [4] Wang Q K, Wang S L, Huang Y. Comparisons of litterfall, litter decomposition and nutrient return in a monoculture Cunninghamia lanceolata and a mixed stand in southern China[J]. Forest Ecology and Management, 2008, 255(3-4):1210-1218. doi: 10.1016/j.foreco.2007.10.026 [5] Xu X N, Hirata E, Enoki T, et al. Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance[J]. Plant Ecology, 2004, 173(2):161-170. doi: 10.1023/B:VEGE.0000029319.05980.70 [6] Bai Y F, Wu J G, Clark C M, et al. Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning:evidence from Inner Mongolia Grasslands[J]. Global Change Biology, 2010, 16(1):358-372. doi: 10.1111/(ISSN)1365-2486 [7] 卢广超, 邵怡若, 薛立.氮沉降对凋落物分解的影响研究进展[J].世界林业研究, 2014, 27(1):35-42. [8] Luca B, Alexandre B, Jonathan H, et al. High nitrogen deposition alters the decomposition of bog plant litter and reduces carbon accumulation[J]. Global Change Biology, 2012, 18(3):1163-1172. doi: 10.1111/j.1365-2486.2011.02585.x [9] 郭春兰, 方向民, 李佩擎, 等.毛竹原状和粉状叶片分解特征对施氮和温度的响应[J].林业科学研究, 2016, 29(5):719-725. doi: 10.3969/j.issn.1001-1498.2016.05.014 [10] Knops J H, Naeem S, Reich P M. The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition[J]. Global Change Biology, 2007, 13(9):1960-1971. doi: 10.1111/gcb.2007.13.issue-9 [11] 陈翔, 周梅, 魏江生, 等.模拟氮沉降对兴安落叶松林凋落物分解的影响[J].生态环境学报, 2013, 22(9):1496-1503. doi: 10.3969/j.issn.1674-5906.2013.09.007 [12] 汪金松, 王晨, 赵秀海, 等.模拟氮沉降对油松林单一及混合叶凋落物分解的影响[J].北京林业大学学报, 2015, 37(10):14-21. [13] 涂利华, 胡红玲, 胡庭兴, 等.华西雨屏区亮叶桦凋落叶分解对模拟氮沉降的响应[J].植物生态学报, 2012, 36(2):99-108. [14] Mo J M, Brown S, Xue J H, et al. Response of Litter Decomposition to Simulated N Deposition in Disturbed, Rehabilitated and Mature Forests in Subtropical China[J]. Plant and Soil, 2006, 282(1-2):135-151. doi: 10.1007/s11104-005-5446-7 [15] Marklein A R, Houlton B Z. Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems[J]. New Phytologist, 2012, 193(3):696-704. doi: 10.1111/j.1469-8137.2011.03967.x [16] Peuelas J, Sardans J, Rivas-ubach A, et al. The human-induced imbalance between C, N and P in Earth's life system[J]. Global Change Biology, 2012, 18(1):3-6. doi: 10.1111/j.1365-2486.2011.02568.x [17] 弓晓静, 余明泉, 胡小飞, 等.氮磷添加对红壤区城郊湿地松林凋落叶分解的影响[J].生态学杂志, 2010, 29(12):2327-2333. [18] Liu Z F, Fu B J, Zheng X X, et al. Plant biomass, soil water content and soil N:P ratio regulating soil microbial functional diversity in a temperate steppe:A regional scale study[J]. Soil Biology & Biochemistry, 2010, 42(3):445-450. [19] 蔡金桓, 王卓敏, 薛立, 等.外源性氮和磷对藜蒴林凋落叶分解的影响[J].中南林业科技大学学报, 2017, 36(7):105-111. [20] Qualls R G, Richardson C J. Phosphorus enrichment affects litter decomposition, immobilization, and soil microbial phosphorus in wetland mesocosms[J]. Soil Science Society of America Journal, 2000, 64(2):799-808. doi: 10.2136/sssaj2000.642799x [21] Liu L, Gundersen P, Zhang T, et al. Effects of phosphorus addition on soil microbial biomass and community composition in three forest types in tropical China[J]. Soil Biology and Biochemistry, 2012, 44(1):31-38. doi: 10.1016/j.soilbio.2011.08.017 [22] DeBusk W F, Reddy K R. Turnover of detrital organic carbon in a nutrient impacted Everglades marsh[J]. Soil Science Society of America, 1998, 62(5):1460-1468. doi: 10.2136/sssaj1998.03615995006200050045x [23] Kozovits A R, Bustamante M M C, Garofalo C R, et al. Nutrient resorption and patterns of litter production and decomposition in a Neotropical Savanna[J]. Functional Ecology, 2007, 21(6):1034-1043. doi: 10.1111/fec.2007.21.issue-6 [24] 邹碧, 李志安, 丁永祯, 等.南亚热带4种人工林凋落物动态特征[J].生态学报, 2006, 26(3):715-721. doi: 10.3321/j.issn:1000-0933.2006.03.011 [25] 薛立; 何跃君; 屈明, 等.华南典型人工林凋落物的持水特性[J].植物生态学报, 2005, 29(3):415-421. doi: 10.3321/j.issn:1005-264X.2005.03.011 [26] 中国土壤学会.土壤农业化学分析方法[M].北京:中国农业科技出版社, 2000 [27] 刘彩霞, 焦如珍, 董玉红, 等.杉木林土壤微生物区系对短期模拟氮沉降的响应[J].林业科学研究, 2015, 28(2):271-276. [28] 李英滨, 李琪, 杨俊杰, 等.模拟氮沉降对温带草原凋落物质量的影响[J].生态学杂志, 2016, 35(10):2732-2737. [29] Knorr M, Frey S D, Curtis P S. Nitrogen additions and litter decomposition:a meta-analysis[J]. Ecology, 2005, 86(12):3252-3257. doi: 10.1890/05-0150 [30] Liu L, Gundersen P, Zhang T, et al. Effects of phosphorus addition on soil microbial biomass and community composition in three forest types in tropical China[J]. Soil Biology and Biochemistry, 2012, 44(1):31-38. doi: 10.1016/j.soilbio.2011.08.017 [31] Chen Y, Fan J B, Du L, et al. The application of phosphate solubilizing endophyte Pantoea dispersa triggers the microbial community in red acidic soil[J]. Applied Soil Ecolgy, 2014, 84:235-244. doi: 10.1016/j.apsoil.2014.05.014 [32] Chen Y, Sun T T, Qian H Y, et al. Nitrogen mineralization as a result of phosphorus supplementation in long-term phosphate deficient soil[J]. Applied Soil Ecolgy, 2016, 106:24-32. doi: 10.1016/j.apsoil.2016.04.019 [33] 陈昊.氮磷添加对亚热带北部常绿阔叶林落叶分解及其养分释放的影响[D].合肥: 安徽农业大学, 2013. [34] 李文亚.贝加尔针茅草原凋落物分解对氮磷添加的响应[D].沈阳: 沈阳农业大学, 2016. [35] 李茂, 徐俊, 田地, 等.氮磷添加对苦槠次生林凋落物量及其养分动态的影响[J].中国农学通报, 2016, 32(19):7-13. doi: 10.11924/j.issn.1000-6850.casb15110145 [36] Jacobson T K B, Bustamante M M C, Kozovits A R. Diversity of shrub tree layer, leaf litter decomposition and N release in a Brazilian Cerrado under N, P and N plus P additions[J]. Environmental Pollution, 2011, 159(10):2236-2242. doi: 10.1016/j.envpol.2010.10.019 [37] 吕妍, 郑泽梅, 美丽班·马木提, 等.增施氮磷肥对木荷林凋落物生产量及其养分的影响[J].应用生态学报, 2013, 24(11):3027-3034. [38] 廖利平, 高洪, 汪思龙, 等.外加氮源对杉木叶凋落物分解及土壤养分淋失的影响[J].植物生态学报, 2000, 24(1):34-39. doi: 10.3321/j.issn:1005-264X.2000.01.007 [39] 李淑兰, 陈永亮.不同落叶林林下凋落物的分解与养分归还[J].南京林业大学学报:自然科学版, 2004, 28(5):59-62. [40] Prescott C E, Kabzems R, Zabek L M. Effects of fertilization on decomposition rate of Populus tremuloides foliar litter in a boreal forest[J]. Canadian Journal of Forestry Research, 1999, 29(3):393-397. doi: 10.1139/x99-016 [41] Berg B and Ekbohm G. Litter mass-loss rates and decomposition patterns in some needle and leaf litter types. Long-term decomposition in a Scotspine forest[J]. Canadian Journal of Botany, 1991, 69(7):1449-1456. doi: 10.1139/b91-187 [42] 董喜光, 张越, 薛立, 等.火力楠林的土壤特性对外源性N和P的响应[J].中南林业科技大学学报, 2016, 35(9):104-108, 113. [43] Yang J X, Zhang T, Wu D X. Study on effect of phosphorus nutrition on drought resistance of plants[J]. Trace Elem Sci, 2003, 10(12):13-19.