Soil enzyme activity and their kinetics in broadleaf forests of Beijing mountainous areas

LIU Xiaoxiao; DAI Wei; DAI Aona

doi:10.11833/j.issn.2095-0756.2018.05.002

Volume 35 Issue 5

Sep. 2018

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LIU Xiaoxiao, DAI Wei, DAI Aona. Soil enzyme activity and their kinetics in broadleaf forests of Beijing mountainous areas[J]. Journal of Zhejiang A&F University, 2018, 35(5): 794-801. doi: 10.11833/j.issn.2095-0756.2018.05.002

Citation:

LIU Xiaoxiao, DAI Wei, DAI Aona. Soil enzyme activity and their kinetics in broadleaf forests of Beijing mountainous areas[J]. Journal of Zhejiang A&F University, 2018, 35(5): 794-801. doi: 10.11833/j.issn.2095-0756.2018.05.002

Soil enzyme activity and their kinetics in broadleaf forests of Beijing mountainous areas

doi: 10.11833/j.issn.2095-0756.2018.05.002

College of Forestry, Beijing Forestry University, Beijing 100083, China

Received Date: 2017-10-23
Rev Recd Date: 2017-12-18
Publish Date: 2018-10-20

Abstract

To study the relationship between characteristics of soil enzymes and soil physicochemical properties of broadleaf forests in the Beijing mountainous area and to determine the feasibility of using soil enzyme activity as well as kinetic parameters as the evaluation indexes of soil fertility, soil from Robinia pseudoacacia, mixed Betula platyphylla-Populus davidiana, Quercus dentata, and Castanea mollissima, four kinds of broadleaf forests in Beijing, were selected as the research objects. Variance, correlation analysis, and principal component analysis (PCA) were used for analysis of four soil enzymes activity (urease, peroxidase, polyphenol oxidase, and catalase) and kinetic characteristics in the 0-20 cm, 20-40 cm, and 40-60 cm soil depths. Results showed that enzyme activity, maximum rate of enzymatic reaction (V_max), and the indicator of enzymatic reaction rate (V_max/K_m) (where K_m is Michaelis constant) of urease and catalase significantly decreased (P < 0.05) with an increase of soil depth. However, K_m had not significant variation. Soil enzymatic reaction efficiency depends not only on enzyme activity but also on its kinetics. Enzyme activity, V_max, and V_max/K_m of urease as well as enzyme activity and K_m of peroxidase could be used to reflect soil fertility in the Beijing mountain forests with other important indexes to evaluate being soil organic matter, total N, available K, available P, and available Fe.
- forest soil science,
- broadleaf forests,
- soil enzyme characteristics,
- soil physicochemical properties,
- correlation analysis,
- principal component analysis (PCA)

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沈阳化工大学材料科学与工程学院沈阳 110142

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土壤酶作为土壤有机物转化的执行者和植物营养元素的活性库，积极参与土壤中所有的生物化学过程^[1]。由于酶对土壤及环境变化比较敏感，因此土壤酶被看作判别土壤质量和胁迫环境下土壤退化的早期预警指标^[2-3]。鉴于此，很多学者将关注焦点集中在土壤酶活性与土壤理化性质关系方面的研究，探讨利用酶活性评价土壤肥力的可能性^[4-9]。为了更加深入了解土壤酶的影响作用，学者们进一步开展了土壤酶促反应动力学特征方面的研究^[10-15]。但由于土壤酶促反应的复杂性，迄今为止，土壤酶活性作为土壤肥力指标虽被大多数者认可，但仍存有争论^[16-17]；研究内容多集中在表层土壤酶活性方面，而结合土壤酶促反应机理所开展的更深入的研究较少；研究对象多以不同农业管理方式下的土壤为主，针对森林土壤的研究很少，且不完善等^[18]。刺槐Robinia psedoacacia林，板栗Castanea mollissima林，槲树Quercus dentata林以及山杨Populus davidiana-白桦Betula platyphylla林是北京山地重要的阔叶林分类型，针对其林下土壤酶活性及酶促反应动力学特征的相关研究较少。为此，本研究选择刺槐、山杨-白桦、槲树和板栗林下土壤为对象，研究土壤脲酶、多酚氧化酶、过氧化物酶和过氧化氢酶活性及动力学特征，并进一步利用主成分分析，分析比较各特征因子评价土壤肥力的可行性，选取合理的肥力评价指标，以期为完善北京林地土壤质量评价体系提供理论依据。

1. 材料与方法

1.1. 材料

选取坡度相近、坡向相同的刺槐林、板栗林、槲树林、以及山杨-白桦林为研究样地。设置100 m × 100 m标准样地(表 1)，每个样地分割为20 m × 20 m的25个样方，利用随机取样法选取5个样方保证代表性，分别采集深度为0~20，20~40和40~60 cm的土壤混合样品，风干过筛后，放入4 ℃冰箱中保存备用。

林型	海拔/m	腐殖质层厚/cm	坡度/（°）	坡向	平均胸径/cm	平均树高/m	密度/（株·hm^-2）	郁闭度
RP	576	0.7	18	东北	11.51	6.09	778	0.65
BP	1 270	5.5	19	东北	13.10	10.97	1 149	0.93
QD	710	0.2	22	东北	14.90	8.72	871	0.67
CM	525	0	15	东北	7.43	3.11	864	0.72
说明：RP代表刺槐林，BP代表山杨-白桦林，QD代表槲树，CM代表板栗林

Table 1. General situation of experimental sites

1.2. 研究方法

1.2.1. 土壤理化性质测定

土壤颗粒组成采用吸管法；土壤有机质用外加热法；土壤全氮采用硫酸-高氯酸消煮-凯氏定氮仪法；土壤速效磷采用碳酸氢钠浸提-钼锑抗比色法；土壤速效钾采用乙酸铵浸提-火焰光度计法；pH值采用酸度计法；土壤有效态铁、锰、锌、铜采用DTPA-TEA浸提-AAS法^[19-20]。

1.2.2. 土壤酶活性及动力学参数测定

脲酶活性采用靛酚蓝比色法，酶活性以37 ℃，24 h土壤释放铵态氮的量表示(mg·g^-1)；多酚氧化酶和过氧化物酶活性采用邻苯三酚比色法，酶活性以30 ℃，2 h土壤生成没食子素的量表示(mg·g^-1)；过氧化氢酶活性采用高锰酸钾滴定法，酶活性以20 min土壤消耗高锰酸钾的量表示(mL·g^-1)。

酶促反应特征参数：分别测定不同反应时间下底物浓度或产物的变化量，利用时间与变化量作图，曲线初期的斜率即为土壤酶促反应初速度(V₀)。再根据Lineweaver-Burk方程(1)，利用1/V₀对1/S作图，求出不同酶促反应的V_max，K_m和1/V_max^[21-23]。

式(1)中：V₀为酶促反应初速度；V_max为酶促反应的最大速度，可表征土壤酶参与酶促反应过程潜在能力的容量指标，即酶的总量；K_m为米氏常数，是酶的特征性常数，只与酶的性质有关，与总酶量无关，可表征酶与底物的亲和能力；S为底物浓度(mol·L^-1)。其中，脲酶底物浓度为0.008，0.020，0.080和0.600 mol·L^-1，反应时间分别为6.0，12.0，18.0和24.0 h；多酚氧化酶底物浓度为0.080，0.160，0.320和0.480 mol·L^-1，反应时间分别为0.5，1.0，2.0和3.0 h；过氧化物酶底物浓度为0.008，0.040，0.080和0.160 mol·L^-1，反应时间分别为0.5，1.0，2.0和3.0 h；过氧化氢酶底物浓度为0.088，0.176，0.352，0.528，0.704和0.880 mol·L^-1，反应时间分别为2.0，4.0，8.0和10.0 min。

1.2.3. 数据处理

采用SPSS 21对4种酶的活性和动力学参数进行方差分析，对4种酶特征值和土壤理化性质进行相关分析和主成分分析。

3. 讨论

土壤脲酶活性和除板栗林外的其他3种林型下土壤过氧化氢酶活性垂直剖面特点明显，表现出随土壤深度增加而降低的特征。板栗林土壤过氧化氢酶可能受到较为强烈的人为影响，因而其活性表现出独特的剖面垂直特征(图 1)。例如农药的施入会强烈抑制土壤过氧化氢酶活性^[22]，在板栗林管理中往往会使用低浓度农药防治栗疫病、白粉病等，这可能是导致板栗林表层土壤过氧化氢酶活性降低的重要原因之一。土壤多酚氧化酶和过氧化物酶活性与土壤腐殖质的形成和转化密切相关^[22]。尽管刺槐、山杨-白桦和槲树林下同层土壤显示出一定的变化规律，但如何更加准确理解不同林型下2种土壤酶活性的特征及变化，还需深入研究。

土壤酶来自于微生物、植物根系和土壤动物的分泌^[22]，其活性被认为是评价土壤肥力的重要指标。研究认为，土壤脲酶活性与土壤肥力密切相关，是评价土壤肥力的重要指标之一^{[8, 15, 21]}。本研究得到相同的结论(图 2)，验证了土壤脲酶活性可作为评价土壤肥力的普遍性指标。土壤过氧化物酶活性对土壤肥力也表现出良好的指示意义，而多酚氧化酶和过氧化氢酶活性则指示意义较低(表 2)，这一结论与前人研究或相同^[27]或不同^[28]。这种研究结论间的差异表明上述土壤酶活性受土壤条件和植被类型的深刻影响，所以表现出同一种酶活性在不同研究中呈现不同的土壤肥力指示意义，不具备土壤脲酶活性所具有的普遍性特点。因此，在进行土壤肥力评价时，选择具有说明意义的土壤酶活性指标是十分必要的。

土壤酶作为一种特殊催化剂影响着土壤中生物化学反应的过程和强度，其活性是影响酶促反应的重要因素。一些学者尝试通过土壤酶活性的变化解释酶对土壤特征的响应和影响^[29]。但本研究表明：土壤酶促反应强度并不仅取决于土壤酶活性的高低，土壤酶的活性及其与底物的结合强度等动力学特征也同样会对酶促反应产生重要影响。因此，单纯利用活性高低解释土壤酶对酶促反应的影响并不利于对酶促反应的理解。以0~20 cm土壤脲酶酶促反应催化效率为例，刺槐林土壤脲酶催化效率最高，其次分别为山杨-白桦林土壤、板栗林和槲树林土壤。这种催化效率的不同是由酶的活性、质量分数和酶-底物亲和力共同作用的结果。刺槐林土壤脲酶具有最高的活性及与底物最强的亲和力，酶促反应催化效率最高；槲树林土壤脲酶活性虽然高于板栗林土壤，但由于活性最低、酶-底物亲和力最弱，催化效率最低；山杨-白桦林土壤脲酶活性显著高于其他3种土壤，脲酶活性最高，但由于活性和酶-底物亲和力的影响，使其酶促反应催化效率低于刺槐林土壤，而高于板栗林和槲树林土壤；板栗林土壤脲酶活性虽然低于槲树林，但由于相对较高的活性及K_m值明显不同，使其酶促反应的催化效率高于槲树林土壤(图 1，表 2)。

4. 结论

土壤脲酶活性和其他3种林型下(除板栗林)的土壤过氧化氢酶活性随土壤深度的增加而减小，而土壤多酚氧化酶和过氧化物酶活性在不同林型下垂直剖面特征不同。土壤脲酶和过氧化氢酶V_max和V_max/K_m随土壤深度的增加而减小；多酚氧化酶和过氧化物酶的动力学参数均无规律性变化。土壤酶促反应效率不仅取决于土壤酶活性，其动力学特征也会对酶促反应产生重要影响，两者结合更利于对酶促反应的理解。土壤酶的种类及其活性和动力学参数对土壤肥力的解释程度不同，脲酶的活性，V_max，V_max/K_m和过氧化物酶的活性，K_m可以作为反映北京山地阔叶林土壤肥力的指标，而多酚氧化酶和过氧化氢酶的活性与动力学参数对土壤肥力的说明意义较小。土壤脲酶的活性，V_max和V_max/K_m以及过氧化物酶K_m，有机质、全氮、速效钾、速效磷、有效铁等因子可作为评价北京山地阔叶林土壤肥力状况的重要指标。

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林型	土层深度/cm	脲酶			多酚氧化酶			过氧化物酶			过氧化氢酶
林型	土层深度/cm	V_max/(μg·g^-1·h^-1)	K_m/(μg·g^-1)	V_max/K_m	V_max/(μg·g^-1·h^-1)	K_m/(μg·g^-1)	V_max/K_m	V_max/(μg·g^-1·h^-1)	K_m/(μg·g^-1)	V_max/K_m	V_max/(μg·g^-1·h^-1)	K_m/(μg·g^-1)	V_max/K_m
RP	0~20	5.76 Ba	7.13 Cb	0.81 Aa	0.26 Aa	0.11 Ab	2.44 Ba	0.75 ABb	0.04 Ba	16.81 Bc	1.19 Ba	1.05 Bb	1.13 Aa
	20~40	3.58 Ab	13.55 Ba	0.26 Ab	0.28 Aa	0.19 Aa	1.50 Bb	1.02 Aa	0.04 Aa	27.17 Bb	0.46 Bb	0.76 ABc	0.61 Bb
	40~60	2.70 Ac	11.34 Cab	0.24 Ab	0.21 ABa	0.12 Ab	1.78 Cb	0.76 Bb	0.02 Cb	43.29 Ba	0.32 Bb	1.35 Aa	0.24 Cc
BP	0~20	7.21 Aa	9.41 BCc	0.77 Aa	0.15 Ba	0.08 Ba	2.00 Ca	0.65 Bc	0.09 Aa	7.52 Cb	2.06 Aa	1.72 Aa	1.20 Aa
	20~40	2.04 Bb	12.04 Bb	0.17 ABb	0.12 Ba	0.09 Ba	1.31 Bb	0.88 Bb	0.05 Aa	17.39 Ca	0.02 Cb	0.27 Bb	0.08 Cb
	40~60	1.25 Bb	28.49 Aa	0.04 Bc	0.14 Ba	0.08 Ba	1.66 Cb	1.04 Aa	0.06 Aa	16.00 Ca	0.02 Cb	0.41 Cb	0.06 Db
QD	0~20	4.28 Ba	22.01 Ab	0.19 Ca	0.19 ABb	0.06 Ba	2.99 Ac	0.88 ABa	0.03 Ba	29.94 Ab	1.32 Ba	1.10 Ba	1.20 Aa
	20~40	1.62 Cb	28.91 Aa	0.06 Bb	0.28 Aa	0.08 Ba	3.51 ABb	0.83 Ba	0.02 Ba	35.09 Ab	1.16 Aab	1.13 Aa	1.02 Aab
	40~60	1.52 Bb	30.97 Aa	0.05 Bb	0.29 Aa	0.06 Ba	4.77 Ba	0.82 Ba	0.01 Ca	66.23 Aa	0.82 Ab	1.16 ABa	0.70 Ab
CM	0~20	3.46 Ba	12.39 Bb	0.28 Ba	0.15 Bb	0.08 Ba	2.00 Cb	1.02 Aa	0.03 Ba	30.03 Aab	0.20 Cb	0.54 Cb	0.37 Ba
	20~40	2.70 Ab	10.05 Bb	0.27 Aa	0.26 Aa	0.07 Ba	3.85 Ab	0.61 Bb	0.02 Ba	36.23 Aa	0.44 Ba	0.92 Aa	0.47 Ba
	40~60	1.33 Bc	21.52 Ba	0.06 Bb	0.20 ABb	0.03 Cb	6.05 Aa	0.69 Bb	0.03 Ba	22.94 Cb	0.38 Ba	0.81 Ba	0.46 Ba
说明：RP代表刺槐林，BP代表山杨-白桦林，QD代表槲树，CM代表板栗林。不同大写字母表示同一土层不同林型差异显著（P＜0.05）；不同小写字母表示相同林型不同土层差异显著（P＜0.05）

酶	酶特征值	活性	V_max	K_m	V_max/K_m
脲酶	活性	1.000
	V_max	0.905^**	1.000
	K_m	-0.596^*	-0.645^*	1.000
	V_max/K_m	0.918^**	0.927^**	-0.734^**	1.000
多酚氧化酶	活性	1.000
	V_max	0.614^*	1.000
	K_m	-0.446	0.260	1.000
	V_max/K_m	0.895^**	0.424	-0.666^*	1.000
过氧化物酶	活性	1.000
	V_max	0.334	1.000
	K_m	-0.550	0.022	1.000
	V_max/K_m	0.205	-0.028	-0.826^**	1.000
过氧化氢酶	活性	1.000
	V_max	0.439	1.000
	K_m	0.352	0.815^**	1.000
	V_max/K_m	0.456	0.937^**	0.698^*	1.000
说明：^表示差异显著（P＜0.05），^*表示差异极显著（P＜0.01）

林型	土层/cm	砂粒2.00~0.02 mm/%	粉粒0.020~0.002 mm/%	黏粒＜0.002 mm/%	pH	w_有机质/(g·kg^-1)	w_全氮/(g·kg^-1)	w_速效磷/(mg·kg^-1)	w_速效钾/(mg·kg^-1)	w_有效铁/(mg·kg^-1)	w_有效锰/(mg·kg^-1)	w_有效锌/(mg·kg^-1)	w_有效铜/(mg·kg^-1)
	0~20	54.49	38.74	6.77	6.57	31.81	1.32	2.06	148.5	23.16	17.37	1.42	0.68
RP	20~40	62.97	30.14	6.90	6.53	20.16	0.87	1.78	70.0	14.22	16.85	0.39	0.82
	40~60	49.51	42.74	7.75	6.54	8.08	0.46	1.03	56.5	8.54	12.17	0.16	0.47
	0~20	48.62	45.74	5.64	6.13	81.10	4.09	3.33	135.0	77.57	14.46	1.23	0.45
BP	20~40	35.46	53.63	10.92	6.08	29.65	1.43	1.42	62.0	43.40	6.39	0.26	0.21
	40~60	23.44	66.13	10.43	6.26	18.78	1.23	1.16	69.5	37.34	4.85	0.19	0.54
	0~20	70.51	24.01	5.49	5.71	28.43	0.97	1.71	47.5	36.02	64.82	1.03	0.55
QD	20~40	58.25	35.91	5.84	5.48	10.74	0.39	1.20	24.3	17.34	56.13	0.40	0.46
	40~60	69.72	24.79	5.49	6.40	7.50	0.15	0.85	20.5	9.49	42.71	0.16	0.17
	0~20	57.11	34.91	7.99	5.64	13.59	0.61	2.58	52.0	63.81	40.32	1.75	2.10
CM	20~40	64.48	28.31	7.22	6.87	8.74	0.36	2.26	50.0	17.82	15.71	0.69	1.32
	40~60	57.33	33.54	9.13	7.13	7.71	0.39	1.39	60.0	13.52	9.76	0.53	1.57
说明：RP代表刺槐林，BP代表山杨?鄄白桦林，QD代表槲树，CM代表板栗林

酶种类	特征值	pH值	有机质	全氮	速效磷	速效钾	黏粒	有效铁	有效锰	有效锌	有效铜
脲酶	活性	-0.043	0.634^*	0.571	0.644^*	0.870^**	-0.318	0.397	-0.090	0.706^*	0.018
	V_max	-0.258	0.793^**	0.739^**	0.784^**	0.775^**	-0.445	0.609^*	0.050	0.741^**	0.017
	K_m	0.450	-0.490	-0.405	-0.162	-0.390	0.411	-0.162	-0.234	-0.190	0.539
	V_max/K_m	-0.136	0.760^**	0.700^*	0.598^*	0.851^**	-0.425	0.403	-0.028	0.580^*	-0.217
多酚氧化酶	活性	0.268	-0.426	-0.491	-0.262	-0.388	-0.532	-0.581^*	0.420	-0.124	0.075
	V_max	0.518	-0.454	-0.475	-0.065	-0.056	-0.253	-0.640^*	-0.030	-0.018	0.412
	K_m	0.195	0.098	0.097	0.074	0.329	0.042	-0.193	-0.340	-0.104	-0.098
	V_max/K_m	0.214	-0.432	-0.460	-0.269	-0.450	-0.323	-0.390	0.334	-0.109	0.183
过氧化物酶	活性	-0.289	-0.726^**	-0.761^**	-0.552	-0.712^**	-0.087	-0.439	0.503	-0.240	0.094
	V_max	0.037	-0.171	-0.157	-0.094	-0.092	0.441	0.038	-0.127	-0.051	0.200
	K_m	-0.213	0.873^**	0.911^**	0.518	0.625^*	0.041	0.728^**	-0.318	0.191	-0.381
	V_max/K_m	0.389	-0.637^*	-0.624^*	-0.362	-0.459	0.105	-0.551	0.051	-0.211	0.487
过氧化氢酶	活性	-0.273	0.492	0.366	0.389	0.338	-0.849^**	0.137	0.564	0.467	-0.204
	V_max	0.106	0.562	0.553	0.880^**	0.589^*	-0.368	0.523	-0.109	0.729^**	0.428
	K_m	0.244	0.333	0.349	0.551	0.385	-0.438	0.175	-0.122	0.394	0.293
	V_max/K_m	0.110	0.348	0.307	0.819^**	0.497	-0.420	0.370	0.029	0.810^**	0.575
说明：^表示差异显著（P＜0.05），^*表示差异极显著（P＜0.01）

项目	特征根	方差贡献率/%	累积方差贡献率/%
第1主成分	7.64	50.93	50.93
第2主成分	2.69	17.94	68.87
第3主成分	1.91	12.74	81.61
第4主成分	1.53	10.20	91.81

Soil enzyme activity and their kinetics in broadleaf forests of Beijing mountainous areas

doi: 10.11833/j.issn.2095-0756.2018.05.002