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磷是森林生态系统中重要的限制营养元素,参与生物体的各种代谢,其有效性在植物净初级生产力和微生物活性中起着重要作用[1]。为研究土壤磷的有效性,一般将磷分为无机磷和有机磷两大类,利用不同的化学提取液对土壤磷进行分级[2]。植物根系结合微生物利用一系列机制增加土壤磷的有效性[3],传统的化学分级方法没有充分反映植物根系和微生物介导磷转化过程。基于此,DELUCA等[4]提出了基于生物有效性的磷分级方法(biologically based phosphorus method,BBP法),定义了4种形式的生物可利用土壤磷组分,分别为用氯化钙提取的根系拦截和扩散的可溶性无机磷(CaCl2-P),用柠檬酸盐提取的吸附在黏土颗粒上或弱结合在无机沉淀物上的活性无机磷(Citrate-P),用酶提取被酸性磷酸酶和植酸酶水解的有机磷(Enzyme-P),用盐酸提取被植物和微生物产生的具有可溶性的活性无机磷质子(HCl-P)。该方法考虑到土壤根际与微生物分泌活化的磷表征磷组分[5],有助于深入理解生态系统内驱动磷转化的生物学机制。
由于人为排放的活性氮含量增加,大气氮沉降加剧[6]。中国亚热带地区是氮沉降最严重的地区之一,在过去30 a里氮沉降量的增幅约60%[7-8]。相关研究表明:长期高强度氮输入会改变土壤微生物生物量和活性,增加土壤酸性磷酸酶活性,从而影响土壤磷循环过程[9-11]。因此,在氮沉降背景下维持有效磷的供应来满足较高森林生产力已成为重要问题。毛竹Phyllostachys edulis是东亚和东南亚地区广泛栽植的竹种[12],在中国面积为467.78万 hm2,占全国竹林面积的72.96%,是重要的森林资源,在缓解气候变化方面具有重要的潜力[7, 13-14]。毛竹是无性繁殖植物,由复杂的地下根鞭上的笋芽发育而成[15],地下根系生物量占地上生物量的30%~50%,远高于其他类型森林生态系统[7, 16]。因此,毛竹林土壤磷循环尤其是生物有效磷组分极有可能与其他森林不同。毛竹主要分布在中国亚热带地区,其氮沉降量已经达30 kg·hm−2·a−1 [17]。LI等[11]研究发现:氮添加增加了毛竹林凋落物量、磷归还量、土壤酸化、土壤微生物生物量磷和有效磷,但降低了叶片磷养分吸收率和丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)侵染率[18],而氮沉降对毛竹土壤磷组分的研究还未见报道,制约了对大气氮沉降增强背景下毛竹林生产力响应机制的认识。本研究拟在不同氮添加强度处理下,分析毛竹林土壤不同深度磷组分的含量及其影响因素,量化各组分对土壤有效磷的贡献程度,为改进毛竹林经营管理措施、提升毛竹林生产力提供科学依据。
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可重复双因素方差分析(表1)表明:除总磷外,氮添加、土壤深度及两者复合作用均显著影响了土壤理化性质(P<0.05)。由表2可知:与对照组相比,表层土壤中,氮添加显著降低了pH (P<0.05),降幅为5.2%~6.2%,显著增加了表层土壤中的全磷、有效磷和微生物生物量磷质量分数及酸性磷酸酶活性(P<0.05),增幅分别为14.3%~23.8%、62.4%~197.8%、100.1%~229.1%和47.9%~79.2%;添加低氮和高氮显著增加了表层土壤有机碳质量分数(P<0.05),增幅分别为27.2%和4.7%;添加中氮显著降低了表层土壤有机碳质量分数 (P<0.05),降幅为3.9%。在深层土壤中,氮添加显著降低了土壤pH和有机碳质量分数(P<0.05),降幅分别为6.2%~11.6%和9.3%~33.3%;显著增加了土壤全磷、有效磷和微生物生物量磷质量分数及酸性磷酸酶活性 (P<0.05),增幅分别为26.5%~40.8%、36.4%~175.5%、34.3%~59.3%、23.7%~76.3%。在对照组和氮添加处理下,深层土壤中的pH显著高于表层土壤(P<0.05),高3.5%~10.7%,而有机碳质量分数、有效磷质量分数和酸性磷酸酶活性显著低于表层土壤(P<0.05),分别低49.6%~185.3%、320.9%~819.3%、26.3%~46.8%。深层土壤全磷质量分数在低氮和中氮添加下分别显著提高了32.7%和29.2% (P<0.05),微生物生物量磷质量分数在中氮添加下显著增加了20.2% (P<0.05),在高氮添加下显著降低了23.7% (P<0.05)。
表 1 氮添加和土壤深度及其交互作用对土壤磷组分质量分数和理化性质的双因素方差分析
Table 1. Two-way ANOVA about the effects of N addition, soil depth and their interactions on the concentrations of P fraction content and soil physicochemical properties
项目 氮添加 土壤深度 氮添加×土壤深度 F η2 F η2 F η2 pH 79.76** 0.93 253.45** 0.94 12.09** 0.69 SOC 82.37** 0.94 603.42** 0.99 41.48** 0.89 TN 105.50** 0.95 213.81** 0.99 7.46* 0.58 TP 81.43** 0.94 68.09** 0.81 0.38 0.07 AP 87.08** 0.94 101.26** 0.95 82.86** 0.94 MBP 452.98** 0.98 60.66** 0.79 182.38** 0.97 ACP 512.74** 0.99 124.97** 0.94 22.77** 0.63 CaCl2-P 24.50** 0.82 8.26* 0.34 1.59 0.23 Citrate-P 118.65** 0.93 65.89** 0.92 39.42** 0.84 Enzyme-P 30.23** 0.65 8.13* 0.60 9.02* 0.63 HCl-P 574.53** 0.99 86.86** 0.94 81.68** 0.94 说明:pH指酸碱度;SOC指有机碳;TN指总氮;TP指总磷;AP指有效磷;MBP指土壤微生物生物量磷;ACP指酸性磷酸酶活性; η2指方差分析中的效应量。*P<0.05;**P<0.01 表 2 氮添加对表层和深层土壤理化性质的影响
Table 2. Soil physical and chemical properties of topsoil and subsoil under N addition treatments
土壤深度/cm 处理 pH SOC/(g·kg−1) TN/(g·kg−1) TP/(g·kg−1) AP/(mg·kg−1) MBP/(mg· kg−1) ACP/
(μmol·g−1·h−1)0~20 N0 4.20±0.07 aB 24.31±0.29 cA 2.29±0.09 aA 0.42±0.01 cA 4.63±0.09 dA 7.26±0.31 cB 0.48±0.02 dA N30 3.98±0.03 bB 30.93±0.16 aA 1.80±0.04 cA 0.52±0.01 aB 13.79±0.04 aA 14.53±0.16 bA 0.71±0.02 cA N60 3.97±0.06 bA 23.35±0.08 dA 2.10±0.01 bA 0.48±0.02 bB 7.52±0.03 cA 14.52±0.17 bB 0.79±0.01 bA N90 3.94±0.03 bB 25.45±0.50 bA 2.08±0.08 bA 0.52±0.01 aA 8.87±0.09 bA 23.89±0.67 aA 0.86±0.02 aA 20~40 N0 4.65±0.03 aA 16.25±0.05 aB 1.42±0.02 aB 0.49±0.02 cA 1.10±0.05 dB 11.44±0.34 dA 0.38±0.01 dB N30 4.36±0.05 bA 10.84±0.08 dB 0.96±0.01 dB 0.69±0.01 aA 1.50±0.05 cB 15.36±0.30 cA 0.47±0.04 cB N60 4.11±0.06 cA 14.74±0.31 bB 1.25±0.03 bB 0.62±0.01 bA 1.93±0.01 bB 17.45±0.50 bA 0.55±0.01 bB N90 4.21±0.03 cA 12.33±0.10 cB 1.01±0.01 cB 0.48±0.01 cA 3.03±0.04 aB 18.22±0.25 aB 0.67±0.11 aB 说明:不同小写字母表示在同一土壤深度处理下不同氮添加处理间差异显著(P<0.05);不同大写字母表示同一氮添加处理下不同土 壤深度间差异显著(P<0.05) -
可重复双因素方差分析(表1)表明:氮添加和土壤深度及其复合作用,除CaCl2-P外,对其他几种生物有效性磷组分质量分数均具有显著影响(P<0.05)。图1显示:氮添加显著增加了基于生物有效性的土壤磷组分质量分数(P<0.05)。在表层土壤中,与对照组相比,低氮显著增加了CaCl2-P和Citrate-P质量分数(P<0.05),增幅分别为28.5%和43.5%;中氮显著增加了CaCl2-P、Enzyme-P和HCl-P质量分数(P<0.05),增幅分别为39.7%、32.4%和101.0%;高氮显著增加了CaCl2-P和HCl-P质量分数(P<0.05),增幅分别为63.3%和155.2%。在深层土壤中,与对照组相比,低氮显著增加了CaCl2-P和Enzyme-P质量分数(P<0.05),增幅分别为33.6%和33.6%;中氮显著增加了CaCl2-P质量分数(P<0.05),增幅为44.3%,而显著降低了Citrate-P质量分数(P<0.05),降幅为24.3%;高氮显著增加了CaCl2-P和Enzyme-P质量分数(P<0.05),增幅分别为58.6%和33.6%;氮添加对HCl-P质量分数无显著影响,HCl-P质量分数随土壤深度的增加显著降低。在中氮处理下,Citrate-P和Enzyme-P质量分数随土壤深度的增加显著降低。
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相关性结果(表3)表明:表层土壤有效磷与各磷组分均呈显著正相关(P<0.05),其中有效磷质量分数与Citrate-P质量分数的相关性最高(r=0.696),与Enzyme-P (r=0.522)和HCl-P (r=0.417)次之,与CaCl2-P最低(r=0.375);深层土壤有效磷与Enzyme-P呈显著正相关(P<0.01,r=0.711)。
表 3 不同土壤深度磷组分与有效磷之间的相关关系
Table 3. Correlation coefficients between P components and available P in different soil depth
项目 土壤深度/cm CaCl2-P Citrate-P Enzyme-P HCl-P AP 0~20 0.375* 0.696** 0.522** 0.417* 20~40 0.286 −0.151 0.711** −0.493 说明:*P<0.05;**P<0.01 由图2~5可见:土壤pH与CaCl2-P(P<0.01)、Enzyme-P (P<0.05)和HCl-P (P<0.01)呈显著负相关,土壤酸性磷酸酶活性与CaCl2-P、Enzyme-P和HCl-P呈显著正相关(P<0.01),土壤微生物生物量磷与CaCl2-P呈显著正相关(P<0.01),土壤有机碳与Citrate-P (P<0.01)和HCl-P (P<0.05)呈显著正相关。土壤pH和酸性磷酸酶活性对CaCl2-P (P<0.01)的影响更为显著。土壤有机碳对Citrate-P (P<0.01)的影响更为显著。
Effect of nitrogen addition on soil phosphorus fractions in the Phyllostachys edulis plantation
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摘要:
目的 了解氮输入对毛竹Phyllostachys edulis林土壤磷组分的影响及其转化机制,为实现毛竹林土壤磷的高效利用提供参考。 方法 以毛竹林为研究对象,设置4个氮添加梯度:0(对照)、30、60、90 kg·hm−2·a−1,利用生物有效磷分级方法测定表层土壤(0~20 cm)和深层土壤(20~40 cm)中的磷组分包括可溶性无机磷(CaCl2-P)、活性无机磷(Citrate-P)、酶水解有机磷(Enzyme-P)和可溶性活性无机磷质子(HCl-P)质量分数,探讨氮添加对毛竹林土壤磷组分的影响及其与土壤有效磷和理化性质的关系。 结果 与对照相比,氮添加显著(P<0.05)增加了所有土层土壤中的CaCl2-P质量分数(28.5%~63.3%)和深层土壤中Enzyme-P质量分数(16.3%~33.6%),而对深层土壤中HCl-P质量分数无显著影响(P>0.05)。低氮处理显著(P<0.05)增加了表层土壤中Citrate-P质量分数(43.5%),中高氮处理显著(P<0.05)增加了表层土壤中HCl-P质量分数(101.0%~155.2%)。在对照和氮添加处理中,表层土壤中的不同磷组分质量分数均显著(P<0.05)高于深层土壤。表层土壤中不同磷组分均与有效磷呈显著正相关(P<0.05),而深层土壤中仅Enzyme-P与有效磷呈显著正相关(P<0.01)。氮添加通过降低土壤pH,增加土壤有机碳、微生物生物量磷和酸性磷酸酶活性,促进了土壤不同磷组分向土壤有效磷的转化。 结论 氮添加提高了毛竹林土壤磷的生物有效性,为全球变化背景下毛竹林高效经营提供了科学参考。图5表3参43 Abstract:Objective This paper aims to understand the effect of nitrogen (N) input on soil phosphorus (P) fraction and its transformation mechanism in moso bamboo (Phyllostachys edulis) plantation, so as to provide a reference for achieving efficient use of soil phosphorus in moso bamboo plantation. Method Moso bamboo plantation were used as research subjects, four N addition gradients (0, 30, 60, 90 kg·hm−2·a−1) were set and the bioavailable P method was used to determine P fractions (CaCl2-P, Citrate-P, Enzyme-P and HCl-P) in the topsoil (0−20 cm) and subsoil (20−40 cm). The effect of N input on P fraction of moso bamboo plantation and its relationship with available P and soil physicochemical properties were explored. Result Compared to the control group, nitrogen addition significantly increased CaCl2-P content (28.5%−63.3%) in all soil layers and Enzyme-P content (16.3%−33.6%) in the subsoil, and had no significant effect on the HCl-P content in the subsoil. Low N treatment significantly increased the Citrate-P content (43.5%) in the topsoil and the medium to high N treatment significantly increased the HCl-P content in the topsoil (101.0%−155.2%). In both the control and N addition treatments, the different phosphorus fractions were significantly higher in the topsoil than in the subsoil. Each soil P fractions in the topsoil were significantly and positively correlated with available P (P<0.05). Enzyme-P in the subsoil was only significantly and positively correlated with available P (P<0.01). N addition accelerated the conversion of soil bioavailable P to available P by decreasing soil pH, increasing soil organic carbon, microbial biomass phosphorus, acid phosphatase activity. Conclusion The input of N increased the bioavailability of soil phosphorus in moso bamboo plantation, which could provide scientific reference for efficient management of moso bamboo plantation in the context of global change. [Ch, 5 fig. 3 tab. 43 ref.] -
Key words:
- nitrogen input /
- bioavailable phosphorus /
- Phyllostachys edulis /
- soil microorganisms /
- acid phosphatase
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表 1 氮添加和土壤深度及其交互作用对土壤磷组分质量分数和理化性质的双因素方差分析
Table 1. Two-way ANOVA about the effects of N addition, soil depth and their interactions on the concentrations of P fraction content and soil physicochemical properties
项目 氮添加 土壤深度 氮添加×土壤深度 F η2 F η2 F η2 pH 79.76** 0.93 253.45** 0.94 12.09** 0.69 SOC 82.37** 0.94 603.42** 0.99 41.48** 0.89 TN 105.50** 0.95 213.81** 0.99 7.46* 0.58 TP 81.43** 0.94 68.09** 0.81 0.38 0.07 AP 87.08** 0.94 101.26** 0.95 82.86** 0.94 MBP 452.98** 0.98 60.66** 0.79 182.38** 0.97 ACP 512.74** 0.99 124.97** 0.94 22.77** 0.63 CaCl2-P 24.50** 0.82 8.26* 0.34 1.59 0.23 Citrate-P 118.65** 0.93 65.89** 0.92 39.42** 0.84 Enzyme-P 30.23** 0.65 8.13* 0.60 9.02* 0.63 HCl-P 574.53** 0.99 86.86** 0.94 81.68** 0.94 说明:pH指酸碱度;SOC指有机碳;TN指总氮;TP指总磷;AP指有效磷;MBP指土壤微生物生物量磷;ACP指酸性磷酸酶活性; η2指方差分析中的效应量。*P<0.05;**P<0.01 表 2 氮添加对表层和深层土壤理化性质的影响
Table 2. Soil physical and chemical properties of topsoil and subsoil under N addition treatments
土壤深度/cm 处理 pH SOC/(g·kg−1) TN/(g·kg−1) TP/(g·kg−1) AP/(mg·kg−1) MBP/(mg· kg−1) ACP/
(μmol·g−1·h−1)0~20 N0 4.20±0.07 aB 24.31±0.29 cA 2.29±0.09 aA 0.42±0.01 cA 4.63±0.09 dA 7.26±0.31 cB 0.48±0.02 dA N30 3.98±0.03 bB 30.93±0.16 aA 1.80±0.04 cA 0.52±0.01 aB 13.79±0.04 aA 14.53±0.16 bA 0.71±0.02 cA N60 3.97±0.06 bA 23.35±0.08 dA 2.10±0.01 bA 0.48±0.02 bB 7.52±0.03 cA 14.52±0.17 bB 0.79±0.01 bA N90 3.94±0.03 bB 25.45±0.50 bA 2.08±0.08 bA 0.52±0.01 aA 8.87±0.09 bA 23.89±0.67 aA 0.86±0.02 aA 20~40 N0 4.65±0.03 aA 16.25±0.05 aB 1.42±0.02 aB 0.49±0.02 cA 1.10±0.05 dB 11.44±0.34 dA 0.38±0.01 dB N30 4.36±0.05 bA 10.84±0.08 dB 0.96±0.01 dB 0.69±0.01 aA 1.50±0.05 cB 15.36±0.30 cA 0.47±0.04 cB N60 4.11±0.06 cA 14.74±0.31 bB 1.25±0.03 bB 0.62±0.01 bA 1.93±0.01 bB 17.45±0.50 bA 0.55±0.01 bB N90 4.21±0.03 cA 12.33±0.10 cB 1.01±0.01 cB 0.48±0.01 cA 3.03±0.04 aB 18.22±0.25 aB 0.67±0.11 aB 说明:不同小写字母表示在同一土壤深度处理下不同氮添加处理间差异显著(P<0.05);不同大写字母表示同一氮添加处理下不同土 壤深度间差异显著(P<0.05) 表 3 不同土壤深度磷组分与有效磷之间的相关关系
Table 3. Correlation coefficients between P components and available P in different soil depth
项目 土壤深度/cm CaCl2-P Citrate-P Enzyme-P HCl-P AP 0~20 0.375* 0.696** 0.522** 0.417* 20~40 0.286 −0.151 0.711** −0.493 说明:*P<0.05;**P<0.01 -
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