Chin J Plant Ecol ›› 2019, Vol. 43 ›› Issue (3): 258-272.doi: 10.17521/cjpe.2018.0299

• Research Articles • Previous Articles    

Characteristics of soil enzymes stoichiometry in rhizosphere of understory vegetation in subtropical forest plantations

GAO Yu-Qiu1,2,DAI Xiao-Qin1,2,3,*(),WANG Jian-Lei1,FU Xiao-Li1,2,3,KOU Liang1,2,3,WANG Hui-Min1,2,3,4   

  1. 1 Qianyanzhou Ecological Research Station, Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Science and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
    2 College of Resource and Environment, University of Chinese Academy of Sciences, Beijing 100190, China
    3 Zhongke-Ji’an Institute of Eco-Environmental Sciences, Ji’an, Jiangxi 343000, China;
    4 Jiangxi Province Key Laboratory of Regional Ecological Processes and Information, Taihe, Jiangxi 343725, China
  • Received:2018-11-30 Revised:2019-03-07 Online:2019-04-23 Published:2019-03-20
  • Contact: DAI Xiao-Qin E-mail:daixq@igsnrr.ac.cn
  • Supported by:
    Supported by the National Natural Science Foundation of China(31730014);The National Key R&D Program of China(2016YFD0600202)

Abstract:

Aims The objective was to explore the stoichiometry of rhizosphere soil enzymes under major understory vegetation and their responses to plantation types and seasons.


Methods Rhizosphere soils of understory shrubs (Loropetalum chinense, Adinandra millettii and Eurya muricata) and herbs (Woodwardia japonica and Dryopteris atrata) were sampled in the early growth stage (April) and the vigorous growth stage (July) in Cunninghamia lanceolata, Pinus massoniana and Pinus elliottii plantations at Qianyanzhou Ecological Research Station, Taihe, Jiangxi. Potential activities of β-1,4-glucosidase (BG, carbon (C) acquiring enzyme), β-1,4-N-acetylglucosaminidase (NAG, nitrogen (N) acquiring enzyme) and leucine aminopeptidase (LAP, N-acquiring enzyme), acid phosphatase (AP, phosphorus (P) acquiring enzyme) and their stoichiometric ratios were measured. Soil physical and chemical properties were also analyzed.


Important findings The results found that (1) rhizosphere soil extracellular enzyme activities associated with C and N acquisition and BG:AP (enzyme C:P) were significantly different among understory species, but P acquisition were not. Both forest stand types and sampling seasons influenced BG:(NAG+LAP) (enzyme C:N). Interactions of understory species, forest stand types and seasons observably affected enzyme C:P. Principal component analysis showed that rhizosphere soil enzyme activities and ecoenzymatic stoichiometry differed significantly among different understory species (Loropetalum chinense was obviously different from Eurya muricata, and both of them were evidently different from other understory species), forest stand types (Cunninghamia lanceolata was different from Pinus massoniana and Pinus elliottii plantations) and sampling seasons. Soil NO3 --N, NH4 +-N, DOC content and C:N were the main edaphic abiotic factors influencing the rhizosphere soil enzyme activities and ecoenzymatic stoichiometry. (2) Standardized major axis analysis showed that there were significantly linear relationship among lg(BG), lg(NAG+LAP) and lg(AP) of rhizosphere soils of understory species. lgBG:lg(NAG+LAP):lgAP(enzyme C:N:P) was approximately 1:1:1.3. Rhizosphere soil enzyme C:P and (NAG+LAP):AP (enzyme N:P) of understory species were 0.14 and 0.15, respectively. The regression slopes of lg(BG), lg(NAG+LAP) and lg(AP) deviated significantly from 1 because AP activities were much higher than BG activities and NAG+LAP activities. This study found that rhizosphere soil enzyme activities and ecoenzymatic stoichiometry were affected by understory species, forest stand types and sampling seasons in which substrate availability played an important role. Compared with C- and N-acquiring enzymes, microorganisms allocated more resources to the production of P-acquiring enzymes, which implied that the growth and activity of soil microorganisms were much more limited by P in rhizosphere soil of understory vegetation in subtropical plantations.

Key words: plantation, soil extracellular enzyme, ecological stoichiometry, red soils

Table 1

Characteristics of understory shrub and herb species in three subtropical plantations"

林分类型
Forest stand type
林下植被类型
Understory species
基径
Basal
diameter (mm)
高度
Height
(cm)
冠幅
Crown
width (cm)
盖度
Coverage
(%)
重要值
Importance
value (%)
杉木林
Cunninghamia
lanceolata
forest
檵木 Loropetalum chinense 11.9 ± 2.7 126 ± 31 81 ± 15 - 19.72
杨桐 Adinandra millettii 13.1 ± 1.0 152 ± 13 94 ± 7 - 30.44
格药柃 Eurya muricata 13.2 ± 1.5 142 ± 16 86 ± 9 - 32.71
狗脊蕨 Woodwardia japonica - 94 ± 12 - 71 ± 7 3.09
暗鳞鳞毛蕨 Dryopteris atrata - 60 ± 5 - 43 ± 8 1.89
马尾松林
Pinus massoniana forest
檵木 Loropetalum chinense 18.9 ± 2.0 413 ± 41 144 ± 16 - 33.68
杨桐 Adinandra millettii 26.7 ± 2.3 376 ± 64 160 ± 12 - 39.88
格药柃 Eurya muricata 13.2 ± 1.1 302 ± 50 81 ± 6 - 31.43
狗脊蕨 Woodwardia japonica - 99 ± 6 - 60 ± 7 3.85
暗鳞鳞毛蕨 Dryopteris atrata - 64 ± 6 - 53 ± 7 3.41
湿地松林
Pinus elliottii forest
檵木 Loropetalum chinense 25.7 ± 1.7 244 ± 34 180 ± 11 - 36.39
杨桐 Adinandra millettii 20.3 ± 3.1 206 ± 21 118 ± 16 - 33.11
格药柃 Eurya muricata 19.6 ± 3.8 112 ± 25 106 ± 18 - 19.22
狗脊蕨 Woodwardia japonica - 86 ± 10 - 73 ± 8 4.15
暗鳞鳞毛蕨 Dryopteris atrata - 70 ± 4 - 38 ± 5 2.07

"

变异来源 Source of variation 酶相关参数 Parameter of soil enzyme
BG NAG+LAP AP BG:(NAG+LAP) BG:AP (NAG+LAP):AP
林分类型 Forest stand types (F) 0.061 0.881 0.156 0.013* 0.092 0.938
林下植被类型 Understory species (U) 0.009** 0.010* 0.173 0.165 0.027* 0.286
取样季节 Sampling seasons (S) 0.066 0.575 0.148 0.032* 0.082 0.113
F × U 0.556 0.920 0.423 0.845 0.580 0.821
F × S 0.070 0.054 0.089 0.294 0.057 0.122
U × S 0.112 0.792 0.957 0.331 0.167 0.573
F × U × S 0.260 0.396 0.823 0.495 0.036* 0.221

Fig. 1

Rhizosphere soil enzyme activities of understory vegetation under different forest stand types in subtropical plantations (mean + SE, n = 5). CL, Cunninghamia lanceolata forest; PM, Pinus massoniana forest; PE, Pinus elliottii forest. LC, Loropetalum chinense; AM, Adinandra millettii; EM, Eurya muricata; WJ, Woodwardia japonica; DA, Dryopteris atrata. Different lowercase letters were significantly different among different understory vegetation species of the same forest stand types at the same sampling season (p < 0.05), Different uppercase letters were significantly different among different stand types at the same sampling season (p < 0.05). BG, β-1,4-glucosidase; NAG+LAP, the sum of β-1,4-N-acetylglucosaminidase and leucine aminopeptidase; AP, acid phosphatase."

Fig. 2

Rhizosphere soil ecoenzymatic stoichiometry of understory vegetation under different forest stand types in subtropical plantations (mean + SE, n = 5). CL, Cunninghamia lanceolata forest; PM, Pinus massoniana forest; PE, Pinus elliottii forest. LC, Loropetalum chinense; AM, Adinandra millettii; EM, Eurya muricata; WJ, Woodwardia japonica; DA, Dryopteris atrata. Different lowercase letters were significantly different among different understory vegetation species of the same forest stand types at the same sampling season (p < 0.05), Different uppercase letters were significantly different among different stand types at the same sampling season (p < 0.05). BG, β-1,4-glucosidase; NAG+LAP, the sum of β-1,4-N-acetylglucosaminidase and leucine aminopeptidase; AP, acid phosphatase."

Fig. 3

Standardized major axis regressions of the log-transformed soil C-, N-, and P-acquiring enzyme activities in subtropical plantations (n = 150). The colors represent the forest stand types: red symbols, Cunninghamia lanceolata forest; green symbols, Pinus massoniana forest; blue symbols, Pinus elliottii forest. Filled symbols and open symbols respectively represent the sampling seasons at April and July.CL, Cunninghamia lanceolata forest; PM, Pinus massoniana forest; PE, Pinus elliottii forest. LC, Loropetalum chinense; AM, Adinandra millettii; EM, Eurya muricata; WJ, Woodwardia japonica; DA, Dryopteris atrata. BG, β-1,4-glucosidase; NAG+LAP, the sum of β-1,4-N-acetylglucosaminidase and leucine aminopeptidase; AP, acid phosphatase."

Fig. 4

Principal component analysis (PCA) of soil enzyme activities and ecoenzymatic stoichiometry in subtropical plantations. The effect of sampling seasons is from the average of PCA scores of soil enzyme activities and ecoenzymatic stoichiometry of understory vegetation within the same season. The effect of forest stand types is from the average of PCA scores of soil enzyme activities and ecoenzymatic stoichiometry of understory vegetation within the same forest stand type. The effect of understory species is from the average of PCA site scores of soil enzyme activities and ecoenzymatic stoichiometry of the same understory species. The colors represent the forest stand types: red symbols, Cunninghamia lanceolata forest; green symbols, Pinus massoniana forest; blue symbols, Pinus elliottii forest. CL, Cunninghamia lanceolata forest; PM, Pinus massoniana forest; PE, Pinus elliottii forest; LC, Loropetalum chinense; AM, Adinandra millettii; EM, Eurya muricata; WJ, Woodwardia japonica; DA, Dryopteris atrata."

Fig. 5

Redundancy analysis (RDA) of soil enzyme activities, ecoenzymatic stoichiometry and physical and chemical properties in subtropical plantations. NO3--N, nitrate nitrogen; NH4+-N, ammonium nitrogen; DOC, dissolved organic carbon; C:N, the ratio of total carbon to total nitrogen. BG, β-1,4-glucosidase; NAG+LAP, the sum of β-1,4-N-acetylglucosaminidase and leucine aminopeptidase; AP, acid phosphatase."

Table 3

Results (p-value) of multi-way repeated measures ANOVAs on the effects of forest stand types, understory species, sampling seasons and their interactions on four major soil nutrient factors in subtropical plantations"

变异来源 Source of variation DOC NH4+-N NO3--N C:N
林分类型 Forest stand types (F) 0.200 0.021* 0.221 0.112
林下植被类型 Understory species (U) 0.000** 0.002** 0.002** 0.000**
取样季节 Sampling seasons (S) 0.226 0.003** 0.178 0.296
F × U 0.735 0.658 0.506 0.058
F × S 0.407 0.384 0.009** 0.623
U × S 0.311 0.095 0.084 0.267
F × U × S 0.129 0.511 0.270 0.800
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