Effects of elevated CO<sub>2</sub> concentration and nitrogen addition on soil physicochemical properties and enzyme activities in the root zone of <i>Lycium barbarum</i>

ZHANG Lei; LI Jingzu; QI Chuanxin; MA Yaping; CAO Bing

doi:10.11833/j.issn.2095-0756.20250127

Volume 42 Issue 6

Nov. 2025

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ZHANG Lei, LI Jingzu, QI Chuanxin, et al. Effects of elevated CO2 concentration and nitrogen addition on soil physicochemical properties and enzyme activities in the root zone of Lycium barbarum[J]. Journal of Zhejiang A&F University, 2025, 42(6): 1232−1242 doi: 10.11833/j.issn.2095-0756.20250127

Citation:

ZHANG Lei, LI Jingzu, QI Chuanxin, et al. Effects of elevated CO₂ concentration and nitrogen addition on soil physicochemical properties and enzyme activities in the root zone of Lycium barbarum[J]. Journal of Zhejiang A&F University, 2025, 42(6): 1232−1242 doi: 10.11833/j.issn.2095-0756.20250127

Effects of elevated CO₂ concentration and nitrogen addition on soil physicochemical properties and enzyme activities in the root zone of Lycium barbarum

DOI: 10.11833/j.issn.2095-0756.20250127

ZHANG Lei^1
,,
LI Jingzu¹,
QI Chuanxin¹,
MA Yaping²,
CAO Bing^{2, 3
,
,}

1.
College of Enology and Horticulture, Ningxia University, Yinchuan 750021, Ningxia, China
2.
College of Forestry and Prataculture, Ningxia University, Yinchuan 750021, Niangxia, China
3.
Ningxia Technical College of Wine and Desertification Prevention, Yinchuan 750001, Ningxia, China

Received Date: 2025-01-19
Accepted Date: 2025-07-04
Rev Recd Date: 2025-06-30

Available Online: 2025-09-15

Publish Date: 2025-12-20

Abstract

Objective This aim was to investigate the changes of the key physicochemical properties and enzyme activities in the rhizosphere soil of Lycium barbarum under the interaction effects of elevated atmospheric CO₂ concentration and nitrogen addition. Method The four-year old potted plants of L. barbarum ‘Ningqi 1’ were used as experimental materials, with the open-top chamber (OTC) control system to simulate the treatment of elevated CO₂ concentration [CO₂, (840±20) μmol·mol⁻¹], and at the same time, the CO₂ concentration in the natural environment was used as the control [ck, (420±20) μmol·mol⁻¹]; The treatments included three levels of nitrogen addition were set: N0 (0 g·kg⁻¹), N1 (1 g·kg⁻¹), and N2 (2 g·kg⁻¹), and the total nitrogen, total phosphorus, organic matter content, and indicators of catalase and polyphenol oxidase were measured and analyzed in the soil of the rhizosphere zone of L. barbarum. Result (1) The interaction between elevated CO₂ concentration and nitrogen addition has an extremely significant impact on soil total potassium, ammonium nitrogen, available phosphorus, and available potassium throughout the entire treatment period(P＜0.01); (2) After 120 days of elevated CO₂ concentration, soil polyphenol oxidase, acid invertase, and urease activities increased by 40.58%, 30.67%, and 36.41% respectively compared to ambient CO₂ concentration, while catalase activity decreased by 61.50%; (3) Significant correlations persisted between total potassium content and soil enzyme activities during the entire treatment duration. Conclusion Under the effects of elevated CO₂ concentration and nitrogen addition, the contents of soil total phosphorus, organic matter, polyphenol oxidase, acid invertase, and urease in the root zone of L.barbarum exhibited an increasing trend, while the contents of soil total nitrogen and catalase showed a decreasing trend. The results of this study provide a reference and guidance basis for the adaptive cultivation and regulation of L.barbarum in Ningxia under the background of global climate change. [Ch, 5 fig. 1 tab. 49 ref.]
- Lycium barbarum,
- CO₂ concentration,
- nitrogen addition,
- organic matter,
- catalase

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大气二氧化碳(CO₂)摩尔分数升高是引起全球气候变化的主要因素^[1]。全球平均CO₂摩尔分数在2023年已经达424 μmol·mol⁻¹，预计到21世纪末达700 μmol·mol⁻¹^[2−3]。CO₂摩尔分数升高引起温室效应，同时也直接影响着植物的生长发育和代谢过程。作为光合作用的关键底物，CO₂摩尔分数变化显著影响植物的碳同化效率^[4]。已有研究表明：高CO₂摩尔分数通过提高植物光合速率与水分利用效率，降低蒸腾作用，促进植物生长，影响生态系统养分元素的分配与利用，导致植物生理代谢和果实营养品质发生变化^[5−6]。值得注意的是，植物对高CO₂摩尔分数的生理响应受氮素供应水平的显著调控^[7−8]。在氮素充足的情况下，核酮糖-1, 5-二磷酸羧化酶/加氧酶(Rubisco酶)的活性可以得到更好的维持和增强，这对于提高植物的光合效率和促进生长具有重要意义^[9]。如在长期CO₂摩尔分数升高条件下增施铵态氮或硝态氮可提升闽楠Phoebe bournei幼苗光合关键酶活性及有机物积累量，促进闽楠幼苗的生长^[10]。在全球气候变化的背景下，氮沉降往往与大气CO₂摩尔分数的升高同时发生，有必要开展其交互作用对植物生长发育与代谢及根区环境的影响^[11−12]。

宁夏枸杞Lycium barbarum为茄科Solanaceae枸杞属Lycium落叶灌木，是典型的药食同源植物^[13]，主要分布在中国西北干旱半干旱地区^[14−15]。其果实富含生物活性化合物，如多糖、类黄酮、类胡萝卜素、甜菜碱和氨基酸等，在营养保健和功能性食品研发中具有重要作用^[16−17]。已有研究表明：CO₂摩尔分数升高可以促进宁夏枸杞果实半乳糖、葡萄糖和蔗糖积累，显著提高糖代谢相关基因表达及相关酶活性^[18]；长期高CO₂摩尔分数处理会使根区土壤微生物总数显著增加，促进宁夏枸杞根区土壤微生物功能多样性，提高其碳源利用率，并改变部分碳源利用类型^[19]。CO₂摩尔分数升高显著增加‘宁杞1号’‘Ningqi 1’根区土壤脲酶和转化酶的活性，而长时间的CO₂摩尔分数倍增可能会加剧对宁夏枸杞根区土壤微生物数量和酶活性的影响^[20]。ZHAO等^[21]发现混合氮添加和单氮添加对紫花苜蓿Medicago sativa土壤酶活性的影响有显著差异，混合施氮显著提高了3种氧化酶的活性，而单一硝态氮的施用对土壤脲酶活性没有明显影响。适当减少氮肥施用量有助于提高‘宁杞7号’‘Ningqi 7’土壤氮的利用率^[22]，并能通过改善土壤质量、微生物群落结构和功能促进可持续农业实践。

然而，目前对于CO₂摩尔分数升高与氮添加对宁夏枸杞根区土壤理化性质和酶活性的交互作用研究还相对不足。因此，本研究通过模拟大气CO₂摩尔分数升高处理和设置全年氮添加量，旨在明确CO₂摩尔分数升高与氮添加对宁夏枸杞根区土壤理化性质和酶活性的具体影响，为气候变化背景下宁夏枸杞的适应性栽培提供科学依据和指导。

1. 材料与方法

1.1. 材料

本研究于2024年在宁夏大学实验农场(106°14′22.19″N，38°13′50.34″E)进行，海拔为1 116.86 m，属于中温带干旱气候区，年均日照为3 000 h，年均气温为8.5 ℃，基地土壤为壤土。在开顶气室(open-top chamber, OTC)内选用长势一致、无病虫害的4年生‘宁杞1号’盆栽植株为试验材料。本研究在课题组前期相同处理的基础上进一步开展，初始阶段土壤中各养分分布均匀^[23]。表1为土壤的基础性质。

处理	全氮/ (g·kg⁻¹)	碱解氮/ (mg·kg⁻¹)	全磷/ (g·kg⁻¹)	有效磷/ (mg·kg⁻¹)	速效钾/ (mg·kg⁻¹)	有机质/ (g·kg⁻¹)	pH
ck+N0	1.56±0.02 d	48.92±0.22 c	0.87±0.02 c	66.13±0.52 a	207.07±1.86 d	13.41±0.44 d	8.35±0.07 a
CO₂+N0	1.68±0.05 c	49.11±0.35 c	0.91±0.01 b	62.66±0.24 c	215.27±2.52 b	14.51±0.35 c	8.33±0.10 a
ck+N1	2.14±0.10 a	57.37±0.39 a	0.71±0.05 d	61.89±0.16 d	205.87±3.12 d	12.26±0.81 d	8.04±0.07 b
CO₂+N1	2.01±0.04 a	46.34±0.21 d	0.94±0.03 b	63.72±0.02 b	237.93±4.52 a	19.07±0.54 a	7.84±0.04 c
ck+N2	1.94±0.02 b	49.17±0.23 c	0.63±0.03 d	62.55±0.12 c	201.23±5.36 d	15.38±0.42 b	7.75±0.04 d
CO₂+N2	1.95±0.02 b	56.51±0.31 b	1.12±0.02 a	62.54±0.16 c	210.87±1.55 c	16.04±0.51 b	7.74±0.02 d
说明：同列不同小写字母表示不同处理间差异显著(P＜0.05)。

Table 1. Physico-chemical properties of potting soil before experimental treatments

1.2. 试验设计

试验采用双因素设计，以OTC模拟CO₂摩尔分数升高处理[CO₂，(840±20) μmol·mol⁻¹]，以自然环境CO₂摩尔分数为对照[ck，(420±20) μmol·mol⁻¹]；设置全年氮添加量为0 (N0)、1 (N1)、2 g·kg⁻¹ (N2)3个水平^[24]。选用3个气室进行CO₂摩尔分数升高处理，3个气室作自然CO₂摩尔分数处理；气室内种植9株，盆栽，每盆1株，每盆土壤22 kg，每个氮素添加量分别处理3株(3次重复)，共计6个气室、54株试验植株；气室内与大田常规水肥管理相同^[25]。

CO₂摩尔分数升高和氮素处理开始于2024年5月下旬，每天8:00—20:00向高CO₂摩尔分数气室通入CO₂气体，控制其摩尔分数在试验要求范围内；以汉和生物普滋苗功能性氮肥为氮源，氮质量分数≥45%，还含有0.2%整合态微量元素，以及0.01%的钼元素。称取0 (N0)、88 (N1)、176 g (N2)肥料分别溶于18 L水中，得到对应质量浓度的水溶氮肥，总计54 L，每个氮水平下每株枸杞灌溉1 L。10 d添加1次氮素，至9月上旬完成全年添加(共计添加10次)。

在枸杞盆栽区域，以树冠投影范围为参照，于盆栽周边均匀选取4个点位，同时在植株根区附近增设1个点位，利用小型土钻采集处理开始后30、60、90、120 d的0~30 cm根区土壤样品。将采集的5份土样混匀，一份置于放冰袋的保温箱中带回实验室−20 ℃保存，后续用于土壤酶活性测定；另一份风干过筛用于测定土壤理化性质。

1.3. 指标测定

取风干土壤，过0.25和1.00 mm筛，用于土壤理化性质的测定。土壤铵态氮、有效磷和速效钾采用中性、石灰性土壤铵态氮、有效磷、速效钾的测定联合浸提-比色法测定^[26]；土壤有机质、全氮、全磷、全钾的测定方法见参考文献[27]。

为全面表征CO₂浓度升高与氮添加对土壤生态功能的协同影响，选择与土壤碳循环、氮循环和氧化应激有关的土壤酶，包括过氧化氢酶(S-CAT)、多酚氧化酶(S-PPO)、脲酶(S-UE)和酸性转化酶(S-AI)为研究对象。采用酶联免疫分析(enzyme-linked immunosorbent assay, ELISA)测定酶活性，标准曲线分别为：Y₁=4.306 5X₁−0.068 6，$R_1^2 $=0.998 4；Y₂=251.33X₂−15.067，$R_2^2 $=0.998 9；Y₃=45.043X₃−0.741，$R_3^2 $=0.993 6；Y₄ =707.56X₄−22.491，$R_4^2 $=0.993 6。

1.4. 数据处理与分析

采用Excel 2019进行数据整理，使用SPSS 26.0和Origin 2024b对数据进行显著性检验和方差分析(P＜0.05)，试验数据用平均值±标准误表示。

3. 讨论

3.1. CO₂摩尔分数升高和氮添加对土壤理化性质的影响

作为全球变化的两大驱动因子，大气CO₂摩尔分数升高以及氮添加对植物养分的吸收、分配和循环过程有着显著影响^[28]。CO₂摩尔分数升高和氮添加可以交互调节关键的土壤过程，如氮循环和土壤有机质分解^[29]。本研究中，在CO₂摩尔分数升高和氮添加处理下，宁夏枸杞根区土壤各养分指标均受到CO₂和氮添加交互作用的影响，这种交互作用不仅能加速土壤养分循环，还能提高土壤微生物多样性，在处理最后阶段，CO₂摩尔分数升高处理下土壤的全氮、铵态氮、全钾、速效钾和有机质含量显著高于自然CO₂处理，然而，却对全磷和有效磷的吸收产生了抑制作用。大气CO₂摩尔分数升高与外源氮对生态系统的影响并非孤立，两者共同促进植物地上部及根系生长，其互作还可能通过改变土壤微生物群落结构和植物根系分泌物等途径，进一步对土壤肥力和生态功能产生影响^[5]。为满足自身生长、代谢等生理活动需求，植物会持续从根区土壤中摄取更多的磷元素。有研究发现：在相同氮沉降下，0~10 cm土层土壤速效磷随着CO₂摩尔分数的增加呈先增后减的趋势^[30]。本研究发现：在CO₂摩尔分数升高和氮添加下，有效磷同样为先增加后降低的趋势。本研究在处理60 d时CO₂+N2有机质含量显著低于ck+N2这一反常现象，可能源于植物-土壤的反馈调节，高CO₂摩尔分数下过量氮素减少了根系分泌物中低分子量有机酸的分泌量，而有机酸可络合稳定土壤有机质^[31]。此外，这也与该处理下多酚氧化酶活性显著增加的结果相印证^[32]。也有研究表明：持续增加的氮素会促进蛋白质和叶绿素合成以及植物根系分泌物增加^[33]，加之CO₂摩尔分数升高，两者协同促进植物生长，使植物光合产物、生物量增加，最终导致土壤有机质含量上升。在CO₂摩尔分数升高的N0和N1处理下，经过90 d，土壤微生物群落结构可能发生了明显改变^[34]，一些能够产生碱性代谢产物的微生物活性增强，使土壤pH升高。当长期处于高CO₂处理且氮添加的环境时，土壤微生物群落结构和功能会发生变化^[35]，使得具有解钾作用的微生物数量或活性发生变化，且后期植株生长量增加，最终致使土壤全钾和速效钾含量降低。基于上述研究结果，在未来宁夏枸杞的种植实践中，应紧密结合作物不同生长阶段的养分需求状况，实现精准配方施肥，为区域生态保护和农业可持续发展提供坚实的科学依据。

3.2. CO₂摩尔分数升高和氮添加对土壤酶活性的影响

土壤酶活性的强弱与土壤内生物代谢和物质转化速度有关，可反映土壤环境质量和土壤微生物多样性^[36−37]。本研究中，在CO₂摩尔分数升高和氮添加处理下，宁夏枸杞根区土壤过氧化氢酶、多酚氧化酶、酸性转化酶和脲酶的活性呈现不同的趋势。具体表现为CO₂摩尔分数升高和氮添加显著促进了多酚氧化酶、酸性转化酶和脲酶的活性，而过氧化氢酶活性则呈现下降的趋势。此外，CO₂和氮添加的交互作用对4种酶的活性在不同时间点产生显著影响，这反映了土壤生态系统在应对环境变化时的复杂响应机制。在本研究中，高摩尔分数 CO₂和充足的氮素为微生物提供了丰富的碳源和氮源，而研究发现多酚氧化酶、酸性转化酶和脲酶与微生物的碳氮代谢密切相关^[38]，这解释了其活性增加的原因。相关研究显示：在高CO₂环境下，红豆树Ormosia hosiei幼苗叶片中Rubisco 酶羧化效率的提升会抑制光呼吸^[39]，叶片中 H₂O₂的主要生成途径受到抑制，使得土壤中过氧化氢酶含量降低。施氮对土壤过氧化氢酶活性具有抑制作用的研究也有报道^[40]，这些综合原因可能最终导致过氧化氢酶活性呈现下降趋势。有研究表明^[41]：在氮肥处理下，土壤脲酶的活性会随着CO₂摩尔分数升高而显著增加。CO₂摩尔分数变化对土壤中的多酚氧化酶也有着重要影响。由于多酚氧化酶的活性与土壤腐质化程度呈正相关^[42]，高摩尔分数CO₂会导致土壤腐质化程度加速，进而使多酚氧化酶的活性增加。究其原因，CO₂增加会显著促使植株及根系生物量的增长，同时根系分泌物种类与数量发生改变，而土壤酶活性的提升与植物根系分泌物的增多有着显著的关联性^[43]。

3.3. CO₂摩尔分数升高和氮添加下土壤理化性质和酶活性的相关性分析

在CO₂摩尔分数升高与氮添加的交互影响下，部分时段或整个处理周期内，各项理化指标与土壤过氧化氢酶、多酚氧化酶、酸性转化酶以及脲酶之间均呈现显著乃至极显著的相关性。就土壤全氮而言，直至处理120 d时才与土壤酶活性建立起相关性，这可能是由于处理前期土壤具备一定的缓冲能力，使得土壤全氮的变化未能直接作用于酶活性^[44−45]。在前期，土壤全磷与过氧化氢酶呈现极显著负相关，到了后期，两者转为极显著正相关；而对于其他3种酶，其与土壤全磷的相关性在处理前后与过氧化氢酶的情况恰好相反，这或许是由于过多的氮添加导致土壤中氮钾比例失衡，微生物为适应此变化而抑制相关代谢途径^[46]，导致其分泌的酶减少，最终使其变为负相关。土壤有效磷在处理60 d及后续阶段，与过氧化氢酶、多酚氧化酶、酸性转化酶和脲酶呈极显著的正相关或负相关，可能是受有效磷调节的相关基因被激活或强化^[47]，从而推动了更多酶的合成与分泌。速效钾在处理30和60 d时，与多酚氧化酶、酸性转化酶、脲酶呈显著正相关，与过氧化氢酶呈显著负相关。在CO₂摩尔分数升高与氮添加处理下，有机质与酶活性相关性持续变动，这可能源于土壤微生物代谢的转变或微生物群落向其他生态位的迁移^[48−49]，确切原因仍有待进一步深入探究。

4. 结论

在本研究中，宁夏枸杞根区土壤的理化性质和酶活性均受到CO₂摩尔分数升高与氮添加交互作用的显著影响。其中，土壤全磷含量与有机质含量随处理时间展现出上升态势。同时，土壤中多酚氧化酶、酸性转化酶以及脲酶的活性随着CO₂摩尔分数的升高呈现出显著的上升趋势。然而过氧化氢酶的活性却在CO₂摩尔分数升高与氮添加的共同作用下受到抑制，呈现出下降趋势。并且在部分或整个处理周期内，所有土壤理化指标均与土壤酶活性表现出显著甚至极显著的相关性，表明两者间存在相互促进作用，说明土壤理化性质的改善可以提升宁夏枸杞根区土壤酶活性。

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