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氮和磷是植物生长的主要限制性营养物质[1]。近年来,人为氮排放导致大气氮沉降迅速增加,预计到2050年,全球氮沉积速率将达到2 × 1014 g·a−1,这将会导致土壤氮/磷比例失衡,进而影响生态系统养分循环[2]。氮沉降量的增加提高了土壤氮有效性,同时通过影响土壤pH值、土壤有机碳、阳离子交换能力等[3-4]来改变土壤的磷有效性。然而,氮沉降对土壤磷动态的影响仍存在很大争议,因为影响磷迁移的因素较多,且磷在土壤中存在无机磷和有机磷两种形态,每一种形态又可分为易分解态磷、中等易分解态磷和难分解态磷[5]。有研究表明,氮添加导致土壤pH值下降,从而活化土壤中的铝和铁离子,造成更多磷被土壤吸附[6],如易分解态磷转化为更难被生物利用的中等易分解态磷或难分解态磷。相反,易分解态磷也可以从其它磷组分的转化中得到补充[7]。土壤易分解态无机磷是土壤中最有效的磷形态,可被植物和微生物直接吸收[8-9]。亚热带地区土壤高度风化,大部分磷被铝和铁氧化物固定,磷有效性低[7]。因此,了解亚热带地区氮沉降对土壤磷形态的影响,对理解磷有效性如何维持植物生长有重要意义。
土壤微生物对磷的吸收和释放对土壤磷的再分配起着至关重要的作用,尤其是有机磷的积累[10]。微生物能产生有机酸、酸性磷酸酶等物质,将中等易分解态的或难分解态的磷转化为易分解态磷[11]。氮沉降可能改变土壤微生物群落结构,如导致真菌/细菌比值变化[12]。此外,一些研究发现,菌根真菌能够促进植物吸收利用可溶性无机磷,是提高植物磷吸收的重要因素[13]。因此,进一步明确氮沉降对土壤微生物生物量及微生物群落结构的影响,对理解土壤磷转化的微生物过程具有重要意义。
黄山松(Pinus taiwanensis Hayata)是中国亚热带森林的代表树种。福建省戴云山国家自然保护区的黄山松林占地64 km²,是中国大陆南部面积最大、保存最完好的黄山松基地[14]。通过预实验,发现本研究区黄山松林土壤总磷含量仅为0.05~0.15 g·kg−1,远低于同研究区毛竹(Phyllostachys edulis(Carriere) J. Houzeau)林(约0.88 g·kg−1)[15]和罗浮栲(Castanopsis faberi Hance)为主的阔叶林(约0.51 g·kg−1)[16],表明黄山松林地土壤“贫磷”状况较明显。本研究旨在探究氮沉降背景下黄山松林土壤磷组分的变化,并进一步探究磷组分变化的驱动因素,为黄山松林如何适应未来氮沉降持续加剧情况提供科学依据。
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短期氮添加未显著影响土壤pH值、TC、TN、C/N值(表1)。与CK相比,DOC在HN处理下显著减少(p < 0.05),0~10 cm土层LN和HN处理分别是CK的75.3%和63.5%,10~20 cm土层LN和HN处理分别是CK的72.1%和44.7%。NH4+-N含量在LN处理下显著减少(p < 0.05),2个土层LN处理分别是CK的69.0%和53.4%。0~10 cm土层的NO3−-N含量在HN处理下显著增加了101.0%(p < 0.05)。
表 1 氮添加对土壤理化性质的影响
Table 1. Effect of nitrogen addition on soil physicochemical properties(mean,n = 4)
土层
Soil depth/cm处理
TreatmentpH值 全碳
TC/(g·kg−1)全氮
TN/(g·kg−1)碳/氮
C/N可溶性有机碳
DOC/(mg·kg−1)铵态氮
NH4 + -N/(mg·kg−1)硝态氮
NO3−-N /(mg·kg−1)0~10 CK 4.32 Aa 55.73 Aa 4.06 Aa 14.98 Aa 325.75 Aa 74.28 Aa 4.80 Ab LN 4.32 Aa 55.99 Aa 3.75 Aa 15.01 Aa 245.37 Aab 51.23 Ab 5.69 Aab HN 4.35 Ba 47.02 Aa 3.12 Aa 15.12 Aa 206.87 Ab 62.77 Aab 9.65 Aa 10~20 CK 4.57 Aa 26.25 Ba 1.81 Ba 14.55 Aa 63.92 Ba 30.31 Ba 6.21 Aa LN 4.60 Aa 27.27 Ba 1.62 Ba 14.82 Aa 46.07 Bab 16.19 Bb 5.49 Aa HN 4.58 Aa 30.98 Ba 2.20 Ba 14.11 Aa 28.60 Bb 29.60 Ba 6.45 Ba 注:不同大写字母表示相同氮添加下不同土层间差异显著(p<0.05),不同小写字母表示相同土层不同氮添加间差异显著(p<0.05)。下同。 Notes:Different capital letters indicate significant differences between different soil layers in the same nitrogen treatment(p<0.05); Different lowercase letters indicate significant differences between different nitrogen treatments in the same soil layer(p<0.05).The same below. -
与CK相比,氮添加显著增加0~10 cm土层的MBN含量(p < 0.05),LN、HN处理分别比CK增加了108.0%和49.6%(表2);但在10~20 cm土层,与CK相比,HN处理的MBP含量显著降低(p < 0.05)。对于微生物生物量比,与 CK 相比,HN 处理仅显著影响 10~20 cm 土层的MBC/MBP 和 MBN/MBP(p<0.05),分别比CK增加了353.9%和429.0%(p<0.05)。
表 2 氮添加对土壤微生物生物量碳、氮和磷及其计量比的影响
Table 2. Effects of nitrogen addition on soil microbial biomass carbon, nitrogen , phosphorus and their ratios(mean,n = 4)
土层
Soil depth /cm处理
Treatment微生物生物量碳
MBC/(mg·kg−1)微生物生物量氮
MBN/(mg·kg−1)微生物生物量磷
MBP/(mg·kg−1)微生物生物量碳/
微生物生物量氮
MBC/MBN微生物生物量碳/
微生物生物量磷
MBC/MBP微生物生物量氮/
微生物生物量磷
MBN/MBP0~10 CK 543.98 Aa 97.09 Ac 42.50 Aa 5.65 Aa 13.72 Aa 2.51 Aa LN 682.61 Aa 201.99 Aa 54.24 Aa 3.35 Ab 12.57 Aa 3.73 Aa HN 616.18 Aa 145.27 Ab 51.48 Aa 4.29 Aa 11.95 Ba 2.93 Ba 10~20 CK 283.83 Ba 50.31 Ba 24.51 Aa 6.89 Aa 13.66 Ab 2.10 Bb LN 360.33 Ba 61.78 Ba 18.50 BAb 6.42 Aa 18.72 Ab 3.85 Bb HN 437.41 Aa 81.91 Ba 9.19 Bb 5.52 Aa 62.00 Aa 11.11 Aa -
本研究区最主要的土壤磷组分是Residual-P和NaOH-Po(表3)。0~10 cm土层中,与CK相比,HN处理显著降低NaOH-Po含量和Residual-P的含量(p < 0.05);10~20 cm土层,HN处理对NaOH-Po和Residual-P含量无显著影响。氮添加对NaOHs-Po含量无显著影响,但显著降低0~10 cm土层的NaHCO3-Po含量(p < 0.05)。此外,除NaOHs-Po外,其余磷组分含量均是0~10 cm土层的高于10~20 cm土层。与CK相比,LN处理显著增加了0~10 cm土层的Resin-P含量(p < 0.05)和2个土层的NaHCO3-Pi含量(p < 0.05);LN处理下,0~10、10~20 cm土层的NaHCO3-Pi含量比CK分别增加了119.21%和127.06%(p < 0.05)。此外,与CK相比,氮添加对2个土层的NaOHs-Pi含量无显著影响。
表 3 氮添加对土壤磷组分的影响
Table 3. Effects of nitrogen addition on the soil phosphorus fractions(mean, n = 4)
mg·kg−1 土层
Soil depth/cm处理
Treatment易分解态磷 Labile P 中等易分解态磷 Moderate labile P 难分解态磷 Stable P Resin-P NaHCO3-Pi NaHCO3-Po NaOH-Pi NaOH-Po NaOHs-Pi NaOHs-Po Residual-P 0~10 CK 6.32 Ab 2.29 Ab 27.86 Aa 19.36 Aa 35.84 Aa 4.94 Aa 2.86 Aa 40.00 Aa LN 13.75 Aa 5.02 Aa 12.75 Ab 21.63 Aa 26.10 Aab 5.56 Aa 2.99 Aa 38.00 Aa HN 10.20 Aab 4.90 Aa 9.28 Ab 12.98 Ab 22.80 Ab 4.33 Aa 2.87 Aa 32.00 Ab 10~20 CK 1.70 Ba 1.70 Ab 6.40 Ba 8.34 Ba 21.36 Ba 2.63 Bab 4.12 Aa 34.00 Ba LN 2.63 Ba 3.86 Aa 3.56 Ba 8.81 Ba 17.07 Aa 3.30 Ba 3.45 Aa 32.00 Aa HN 2.94 Ba 1.55 Bb 4.08 Ba 7.26 Ba 17.94 Aa 1.85 Bb 3.77 Aa 33.00 Aa 将土壤磷组分进一步归类为易分解态磷、中等易分解态磷、难分解态磷(图1)。0~10 cm土层中,HN处理显著降低中等易分解态磷、难分解态磷的含量(p<0.05),且中等易分解态磷和难解态磷分别是对照的64.8%和82.0%,而氮添加对2个土层易分解态磷含量均无显著影响(图1)。
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0~10 cm土层,氮添加对土壤ACP、PD活性无显著影响(图2);10~20 cm土层,LN和HN处理的ACP活性比CK分别增加了126.0%和157.4%,而PD活性比CK分别增加了99.%和87.51%(p < 0.05)。
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0~10 cm土层,氮添加对土壤微生物群落结构无显著影响(图3);10~20 cm土层,与CK相比,LN处理显著增加G−的含量(p < 0.05),且显著减少了G+/G−的比值(p < 0.05)。此外,0~10 cm土层各土壤微生物标志物含量均高于10~20 cm土层。
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本研究中,土壤性质和微生物特性指标对易分解态磷、中等易分解态磷和难分解态磷均有较好的解释度(图4a)。易分解态磷含量主要与PD活性、NH4+-N、DOC、AMF和G-含量有关;中等易分解态磷含量主要与NH4+-N、DOC含量有关;难分解态磷主要与NH4+-N、TN含量有关。通过冗余分析发现,土壤性质和微生物特性指标共解释了所有土壤磷组分变化的72.07%(图4b)。DOC、AMF是影响土壤磷组分变化的2个最重要因素,DOC含量的变化解释了土壤磷组分变化的63.0%。
氮添加对亚热带黄山松林土壤磷组分的影响
Effects of Nitrogen Addition on the Soil Phoshorus Fractions in Subtropical Pinus taiwanensis Forests
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摘要:
目的 探究氮沉降背景下黄山松林土壤磷组分的变化,并进一步探究磷组分变化的驱动因素,为黄山松林如何适应未来氮沉降持续加剧情况提供科学依据。 方法 对福建省戴云山黄山松林进行短期氮添加试验,测定土壤磷组分、微生物生物量、酸性磷酸单脂酶(ACP)、磷酸二酯酶(PD)和磷脂脂肪酸,探究氮沉降对亚热带土壤磷组分、土壤微生物群落的影响和驱动土壤磷转化的关键因素。 结果 与对照相比,高氮添加显著降低0~10 cm土层中等易分解态磷和难分解态磷含量(p<0.05),对易分解态磷的含量、微生物生物量磷、ACP、PD活性和土壤微生物群落组成无显著影响。总体上,10~20 cm土层磷组分变化趋势与0~10 cm土层一致,但变化不显著。高氮添加显著降低10~20 cm土层微生物生物量磷含量,显著提高ACP、PD活性和微生物生物量氮/微生物生物量磷。此外,低氮添加显著降低10~20 cm土层革兰氏阳性菌/革兰氏阴性菌的比值(p<0.05)。冗余分析表明,可溶性有机碳和丛枝菌根真菌是影响土壤磷组分变化的关键因素。 结论 短期低氮添加下通过磷组分转化(如中等易分解态磷的矿化)维持了土壤磷有效性。这些结果有助于理解短期氮沉降下贫磷生态系统土壤磷有效性和生产力的维持机制。 Abstract:Objective To investigate the changes in soil phosphorus (P) fractions in Pinus taiwanensis forest under nitrogen (N) deposition, and further explore the driving factors of P fractions changes for providing scientific basis for how Pinus taiwanensis forest adapts to the continuous aggravation of nitrogen deposition in the future. Method Soil P fractions, microbial biomass, acid phosphomonoesterase activity (ACP), phosphodiesterase enzyme activity (Phosphodiesterase enzyme, PD), and soil phospholipid fatty acids were detected in a short-term N-addition experiment in a Pinus taiwanensis forest on Daiyun Mountain, Fujian Province, China. Result There were significant effects of high N addition on the content of moderately labile P and stable P at 0−10 cm depth, whereas insignificant effects on the content of labile P, microbial biomass, ACP, PD, and community composition. In addition, low nitrogen supplementation significantly decreased the ratio of gram-positive bacteria to gram-negative bacteria in 10−20 cm soil layer (p < 0.05). In general, the variation trend of P fractions in the 10−20 cm soil layer was consistent with that in the 0−10 cm soil layer, but the change was not significant. However, N addition significantly decreased the content of microbial biomass P, and significantly increased ACP, PD activity and microbial biomass N/microbial biomass P. In addition, low N addition significantly decreased the ratio of gram-positive bacteria to gram-negative bacteria in 10−20 cm depth (p < 0.05). Redundancy analysis showed that soluble organic carbon and arbuscular mycorrhizal fungi key factors affecting changes in soil P fractions. Conclusion Short-term N addition maintains the soil P availability by promoting P transformation (e.g. the mineralization of moderate labile P), which is helpful for understanding the maintenance mechanism of soil P availability and productivity in P-poor ecosystems under short-term nitrogen deposition. -
Key words:
- nitrogen deposition
- / phosphorus fractions
- / phosphatase activity
- / microbial community
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表 1 氮添加对土壤理化性质的影响
Table 1. Effect of nitrogen addition on soil physicochemical properties(mean,n = 4)
土层
Soil depth/cm处理
TreatmentpH值 全碳
TC/(g·kg−1)全氮
TN/(g·kg−1)碳/氮
C/N可溶性有机碳
DOC/(mg·kg−1)铵态氮
NH4 + -N/(mg·kg−1)硝态氮
NO3−-N /(mg·kg−1)0~10 CK 4.32 Aa 55.73 Aa 4.06 Aa 14.98 Aa 325.75 Aa 74.28 Aa 4.80 Ab LN 4.32 Aa 55.99 Aa 3.75 Aa 15.01 Aa 245.37 Aab 51.23 Ab 5.69 Aab HN 4.35 Ba 47.02 Aa 3.12 Aa 15.12 Aa 206.87 Ab 62.77 Aab 9.65 Aa 10~20 CK 4.57 Aa 26.25 Ba 1.81 Ba 14.55 Aa 63.92 Ba 30.31 Ba 6.21 Aa LN 4.60 Aa 27.27 Ba 1.62 Ba 14.82 Aa 46.07 Bab 16.19 Bb 5.49 Aa HN 4.58 Aa 30.98 Ba 2.20 Ba 14.11 Aa 28.60 Bb 29.60 Ba 6.45 Ba 注:不同大写字母表示相同氮添加下不同土层间差异显著(p<0.05),不同小写字母表示相同土层不同氮添加间差异显著(p<0.05)。下同。 Notes:Different capital letters indicate significant differences between different soil layers in the same nitrogen treatment(p<0.05); Different lowercase letters indicate significant differences between different nitrogen treatments in the same soil layer(p<0.05).The same below. 表 2 氮添加对土壤微生物生物量碳、氮和磷及其计量比的影响
Table 2. Effects of nitrogen addition on soil microbial biomass carbon, nitrogen , phosphorus and their ratios(mean,n = 4)
土层
Soil depth /cm处理
Treatment微生物生物量碳
MBC/(mg·kg−1)微生物生物量氮
MBN/(mg·kg−1)微生物生物量磷
MBP/(mg·kg−1)微生物生物量碳/
微生物生物量氮
MBC/MBN微生物生物量碳/
微生物生物量磷
MBC/MBP微生物生物量氮/
微生物生物量磷
MBN/MBP0~10 CK 543.98 Aa 97.09 Ac 42.50 Aa 5.65 Aa 13.72 Aa 2.51 Aa LN 682.61 Aa 201.99 Aa 54.24 Aa 3.35 Ab 12.57 Aa 3.73 Aa HN 616.18 Aa 145.27 Ab 51.48 Aa 4.29 Aa 11.95 Ba 2.93 Ba 10~20 CK 283.83 Ba 50.31 Ba 24.51 Aa 6.89 Aa 13.66 Ab 2.10 Bb LN 360.33 Ba 61.78 Ba 18.50 BAb 6.42 Aa 18.72 Ab 3.85 Bb HN 437.41 Aa 81.91 Ba 9.19 Bb 5.52 Aa 62.00 Aa 11.11 Aa 表 3 氮添加对土壤磷组分的影响
Table 3. Effects of nitrogen addition on the soil phosphorus fractions(mean, n = 4)
mg·kg−1 土层
Soil depth/cm处理
Treatment易分解态磷 Labile P 中等易分解态磷 Moderate labile P 难分解态磷 Stable P Resin-P NaHCO3-Pi NaHCO3-Po NaOH-Pi NaOH-Po NaOHs-Pi NaOHs-Po Residual-P 0~10 CK 6.32 Ab 2.29 Ab 27.86 Aa 19.36 Aa 35.84 Aa 4.94 Aa 2.86 Aa 40.00 Aa LN 13.75 Aa 5.02 Aa 12.75 Ab 21.63 Aa 26.10 Aab 5.56 Aa 2.99 Aa 38.00 Aa HN 10.20 Aab 4.90 Aa 9.28 Ab 12.98 Ab 22.80 Ab 4.33 Aa 2.87 Aa 32.00 Ab 10~20 CK 1.70 Ba 1.70 Ab 6.40 Ba 8.34 Ba 21.36 Ba 2.63 Bab 4.12 Aa 34.00 Ba LN 2.63 Ba 3.86 Aa 3.56 Ba 8.81 Ba 17.07 Aa 3.30 Ba 3.45 Aa 32.00 Aa HN 2.94 Ba 1.55 Bb 4.08 Ba 7.26 Ba 17.94 Aa 1.85 Bb 3.77 Aa 33.00 Aa -
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