抚育间伐对云南松非结构性碳和化学计量特征的影响

王丽娜; 吴俊文; 董琼; 石卓功; 胡昊程; 吴单子; 李禄平

doi:10.12171/j.1000-1522.20210115

抚育间伐对云南松非结构性碳和化学计量特征的影响

doi: 10.12171/j.1000-1522.20210115

1.
西南林业大学林学院，云南昆明 650224
2.
永仁县国有林场，云南永仁 651400

基金项目: “十三五”国家重点研发专项（2017YFD0601202），国家自然科学基金项目（31960306）

详细信息

作者简介:
王丽娜。主要研究方向：森林培育。Email：2367937610@qq.com　地址：650224云南省昆明市盘龙区西南林业大学林学院

责任作者:
吴俊文，博士，讲师。主要研究方向：森林培育、植物生理生态。Email：2569755639@qq.com　地址：同上

中图分类号: S791.257
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出版历程
- 收稿日期: 2021-03-25
- 修回日期: 2021-05-26
- 网络出版日期: 2021-07-10
- 刊出日期: 2021-08-31

Effects of tending and thinning on non-structural carbon and stoichiometric characteristics of Pinus yunnanensis

1.
College of Forestry, Southwest Forestry University, Kunming 650224, Yunnan, China
2.
Yongren County State-Owned Forest Farm, Yongren 651400, Yunnan, China

摘要

摘要: 目的探究抚育间伐对云南松各个器官非结构性碳（NSC），以及碳（C）、氮（N）、磷（P）化学计量学特征的影响机制。方法以云南省永仁县国有林场白马河林区云南松中龄林为研究对象，设置间伐（间伐强度为35.4%）与对照样地，分析抚育间伐对云南松非结构性碳和化学计量特征的影响。结果（1）抚育间伐显著影响云南松针叶和枝的NSC含量（P < 0.01），对根NSC含量则无显著影响。抚育后枝NSC含量增长9.30%，而针叶NSC含量减少3.57%。（2）抚育间伐对云南松枝C含量、针叶和干P含量有显著影响（P < 0.05），对针叶N含量有极显著影响（P < 0.01），但对根的C、N、P含量没有显著影响。抚育后云南松枝中C含量增加12.93%；针叶中N含量增长18.04%；叶和干中P含量分别降低了27.78%、55.56%。（3）云南松NSC、CNP化学计量学特征在各个器官之间表现出显著差异（P < 0.05）。相关性分析表明：针叶和根P含量与可溶性糖含量呈显著正相关（P < 0.05），而叶中C含量则和可溶性糖含量显著负相关（P < 0.05）。结论抚育间伐能促进云南松干、枝和根NSC含量以及各器官C含量的积累，降低叶NSC和各器官P含量，改变C、N、P在各器官中的分布格局；该地区云南松人工林生长在一定程度上受N限制，适当配施N肥将促进林分质量的提高。
- 云南松 /
- 抚育间伐 /
- 非结构性碳 /
- 化学计量特征 /
- 养分分布格局
Abstract: Objective Our objective was to explore the mechanism of the effects of thinning on non-structural carbon (NSC), carbon (C), nitrogen (N) and phosphorus (P) stoichiometry of Pinus yunnanensis. Method The middle-aged P. yunnanensis forests were chosen in the Baima River forest area of Yongren County, Yunnan Province of southwestern China, and two experimental sample plots of thinning (thinning intensity of 35.4%) and control were set up to analyze the effects of thinning on NSC and CNP stoichiometric characteristics of P. yunnanensis forests. Result (1) Tending thinning significantly affected the NSC content of needles and branches of P. yunnanensis (P < 0.01), whereas no significant difference on roots was observed. In addition, the NSC content of branch increased by 9.30%, while the NSC content in needles decreased by 3.57%. (2) Thinning had a significant effect on C content of P. yunnanensis branch, P content in needles and stems (P < 0.05), and a highly significant effect on N content in needles (P < 0.01), however, no significant effect was observed on C, N and P content in roots. After thinning, the C content of P. yunnanensis branch increased by 12.93%; the N content in needles increased by 18.04%, and the P content in needles and stems decreased by 27.78% and 55.56%, respectively. (3) The NSC, CNP stoichiometric characteristics showed significant differences among P. yunnanensis organs (P < 0.05). Correlation analysis showed that the P content in needles and roots was significantly positively correlated with soluble sugar content (P < 0.05), while C content in needles was significantly negatively correlated with soluble sugar content (P < 0.05). Conclusion Thinning promots the accumulation of NSC content in stem, branch and root and C content in each organ, whereas needle NSC and P content decreased in each organ, furthermore, distribution pattern of C, N and P in each organ was affected; the growth of P. yunnanensis plantation in this area was limited by N to some extent, and appropriate application of N fertilizer will promote the improvement of stand quality.
- Pinus yunnanensis /
- tending and thinning /
- non-structural carbohydrate /
- stoichiometric characteristics /
- nutrient distribution pattern

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图 1 不同处理间云南松非结构性碳水化合物积累及分配

不同大写字母表示抚育组不同器官间差异显著(P < 0.05)，不同小写字母表示未抚育组不同器官间差异显著(P < 0.05)。下同。Different capital letters indicate significant differences among varied organs in the tending group (P < 0.05), and different lowercase letters indicate significant differences among varied organs in the control group (P < 0.05). Same as below.

Figure 1. Accumulation and distribution of non-structural carbohydrates in Pinus yunnanensis among different treatments

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图 2 不同处理间云南松碳氮磷及其计量比变化

Figure 2. Changes of C, N, P and their stoichiometric ratios of Pinus yunnanensis in different treatments

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表 1 永仁县国有林场白马河林区样地基本情况调查表

Table 1. Basic information survey of Yongren County Baima River Forest Farm

样地编号 Sample plot No.	海拔 Altitude/ m	坡位和坡向 Slope position and slope aspect	平均胸径 Average BDH/cm	平均树高 Average tree height/m	间伐前林木数 Pre-thinning plant number	采伐株数 Felled tree number	间伐后林木数 Post-thinning plant number	林分密度/ (株·hm⁻²) Stand density/ (tree ·ha⁻¹)	单株材积 Individual volume/ m³	林分蓄积 Stand volume/ m³
WFY-1	2 275	上坡位、西北330°、半阴坡 Uphill position, northwest 330°, semi-shady slope	12.3	10.3	126	0	126	1 890	0.068 5	8.637 2
WFY-3	2 216	中坡位、西北318°、半阴坡 Middle slope, northwest 318°, semi-shady slope	15.2	12.6	95	0	95	1 425	0.120 7	11.470 4
WFY-4	2 480	上坡位、正西269°、半阳坡 Uphill position, due west 269°, half-sunny slope	14.8	11.1	118	0	118	1 770	0.104 1	12.288 4
FY-2	2 360	上坡位、北偏东92°、半阴坡 Uphill position, north by east 92°, semi-shady slope	16.5	12.9	102	36	66	990	0.144 3	9.523 2
FY-5	2 530	上坡位、西北308°、半阴坡 Uphill position, northwest 308°, semi-shady slope	16.3	10.0	85	30	55	825	0.116 2	6.392 3
FY-6	2 438	上坡位、西北300°、半阴坡 Uphill position, northwest 300°, semi-shady slope	18.8	14.0	95	33	62	930	0.198 1	12.281 7
注：抚育样地间伐强度为总株数的35%，间伐木的选择按照留优去劣、间密留匀、兼顾株间距的原则确定，于2017年年初完成。对照样地不采取任何抚育措施。Notes: the thinning intensity of tending sample plots is 35% of the total number of plants. The selection of thinning is determined in accordance with the principles of keeping the best and eliminating the bad, keeping the density evenly and giving consideration to the spacing between plants, and we have completed at the beginning of year 2017. No measure was taken on the CK land.

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表 2 样地林分调查表

Table 2. Survey of sample site stand

样地编号 Sample plot No.	林木株数 Plant number	胸径 DBH/cm		树高 Tree height/m		材积 Volume/m³		蓄积量 Accumulation volume/m³
样地编号 Sample plot No.	林木株数 Plant number	平均值 Average value	平均增长量 Average growth	平均值 Average value	平均增长量 Average growth	平均值 Average value	平均增长量 Average growth	平均值 Average value	平均增长量 Average growth
WFY-1	124	13.5	1.2	11.7	1.4	0.090 5	0.022 0	11.222 0	2.584 8
WFY-3	95	16.8	1.6	14.1	1.5	0.159 9	0.039 2	15.190 5	3.720 1
WFY-4	118	16.6	1.8	13.6	2.5	0.152 0	0.047 9	17.936 0	5.647 6
FY-2	66	18.9	2.4	14.9	2.0	0.209 8	0.065 5	13.846 8	4.323 6
FY-5	54	19.2	2.9	12.6	2.6	0.190 6	0.074 4	10.292 4	3.900 1
FY-6	62	20.0	1.2	15.8	1.8	0.245 0	0.046 9	15.190 0	2.908 3
注：平均增长量是指与表1中指标相比。 Note: the average growth index means compared with the indicators in Tab. 1.

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表 3 抚育对不同器官非结构性碳水化合物含量影响的双因素方差分析（F值）

Table 3. Two-way ANOVA on the effects of tending on the content of non-structural carbohydrates indifferent organs (F value)

因素 Factor	可溶性糖 Soluble sugar	淀粉 Starch	NSC
抚育 Tending	0.096	13.610**	3.401
器官 Organ	134.330**	73.269**	138.487**
抚育 × 器官 Tending × organ	2.220	2.520	2.222
注：表示 P < 0.01。下同。Notes: denotes P < 0.01. Same as below.

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表 4 抚育对不同器官非结构性碳水化合物含量影响的独立样本t检验分析

Table 4. Independent sample t-test analysis of the effects of tending on the content of non-structural carbohydrates in different organs

指标 Index	叶 Needle	干 Stem	枝 Branch	根 Root
可溶性糖 Soluble sugar	−6.890**	0.404	1.623	0.527
淀粉 Starch	1.197	3.069**	4.507**	0.102
NSC	−2.674**	1.967	3.230**	0.407

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表 5 抚育对不同器官C、N、P含量及其计量比的双因素方差分析（F值）

Table 5. Two-way ANOVA on the effects of tending on C, N, P contents and their stoichiometric ratios in different organs (F value)

因素 Factor	C	N	P	C∶N	C∶P	N∶P
抚育 Tending	5.828*	3.817	11.902**	1.346	13.003**	18.249**
器官 Organ	18.621**	554.421**	5.560**	14.139**	7.486**	177.257**
抚育 × 器官 Tending × organ	1.308	9.838**	4.092**	0.792	3.061*	10.077**
注：表示P < 0.05。下同。Notes: denotes P < 0.05. Same as below.

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表 6 抚育对不同器官C、N、P含量及其计量比的独立样本t检验分析

Table 6. Independent sample t-test analysis of the effects of tending on C, N, P contents and their stoichiometricratios in different organs

指标 Index	t值 t value
指标 Index	叶 Needle	干 Stem	枝 Branch	根 Root
C	1.564	0.246	2.370*	0.454
N	3.835**	−0.489	0.040	−1.482
P	−2.646*	−4.167**	−0.376	−0.553
C∶N	−1.249	0.740	0.622	0.979
C∶P	2.216*	3.873**	0.433	1.155
N∶P	4.057**	3.327**	−0.153	−0.141

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表 7 云南松叶−干−枝−根C、N、P含量相关性

Table 7. Correlations of C, N and P contents in needle-stem-branch-root of Pinus yunnanensis

指标 Index	叶C Needle C	叶N Needle N	叶P Needle P	干C Stem C	干N Stem N	干P Stem P	枝C Branch C	枝N Branch N	枝P Branch P	根C Root C	根N Root N	根P Root P
叶C Needle C	1
叶N Needle N	−0.99	1
叶P Needle P	−0.589*	−0.341	1
干C Stem C	−0.258	0.315	−0.312	1
干N Stem N	−0.148	0.062	0.177	−0.101	1
干P Stem P	−0.408	−0.493*	0.501*	0.100	−0.001	1
枝C Branch C	0.186	0.409	−0.124	0.129	0.165	−0.613**	1
枝N Branch N	0.008	−0.182	0.148	−0.148	−0.343	0.038	−0.387	1
枝P Branch P	0.086	−0.279	0.182	−0.064	−0.39	0.256	−0.289	0.537**	1
根C Root C	0.231	−0.263	0.226	−0.308	0.050	−0.052	−0.012	0.395	0.423	1
根N Root N	−0.310	−0.031	0.247	0.173	0.654**	0.302	0.059	−0.282	−0.278	−0.077	1
根P Root P	−0.200	−0.090	0.049	0.107	−0.062	0.219	−0.409	0.222	0.411	0.270	−0.061	1

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表 8 云南松叶、干、枝、根C∶N∶P相关性

Table 8. Correlations of C, N and P stoichiometry of needle-stem-branch-root of Pinus yunnanensis

指标 Index	叶C∶N Needle C∶N	叶C∶P Needle C∶P	叶N∶P Needle N∶P	干C∶N Stem C∶N	干C∶P Stem C∶P	干N∶P Stem N∶P	枝C∶N Branch C∶N	枝C∶P Branch C∶P	枝N∶P Branch N∶P	根C∶N Root C∶N	根C∶P Root C∶P	根N∶P Root N∶P
叶C∶N Needle C∶N	1
叶C∶P Needle C∶P	0.398	1
叶N∶P Needle N∶P	−0.280	0.753**	1
干C∶N Stem C∶N	−0.058	0.226	0.214	1
干C∶P Stem C∶P	−0.201	0.386	0.571*	0.168	1
干N∶P Stem N∶P	−0.210	0.298	0.516*	−0.332	0.848**	1
枝C∶N Branch C∶N	−0.200	0.023	0.201	−0.172	0.118	0.232	1
枝C∶P Branch C∶P	−0.291	−0.029	0.269	−0.303	0.174	0.337	0.635**	1
枝N∶P Branch N∶P	−0.041	−0.064	0.071	−0.310	0.110	0.223	−0.078	0.671**	1
根C∶N Root C∶N	0.502*	0.388	0.041	0.246	0.166	0.060	−0.039	−0.273	−0.329	1
根C∶P Root C∶P	0.099	0.036	−0.020	−0.150	0.425	0.405	0.252	0.223	0.082	0.293	1
根N∶P Root N∶P	−0.281	−0.287	−0.090	−0.441	−0.052	0.120	0.368	0.473*	0.301	−0.570*	0.481*	1

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表 9 云南松NSC构成与C、N、P及其计量比间相关系数

Table 9. Correlation coefficients among NSC composition, C, N, P and their stoichiometric ratios of Pinus yunnanensis

器官 Organ	指标 Index	可溶性糖 Soluble sugar	淀粉 Starch	NSC	C	N	P	C∶N	C∶P	N∶P
叶 Needle	可溶性糖 Soluble sugar	1
	淀粉 Starch	−0.120	1
	NSC	0.684**	0.642**	1
	C	−0.587*	−0.110	−0.534*	1
	N	−0.272	0.247	−0.029	−0.099	1
	P	0.561*	−0.223	0.270	−0.589*	−0.341	1
	C∶N	−0.175	−0.236	−0.308	0.664**	−0.801**	−0.120	1
	C∶P	−0.607*	0.134	−0.370	0.836**	0.149	−0.901**	0.398	1
	N∶P	−0.546*	0.357	−0.160	0.393	0.721**	−0.864**	−0.280	0.753**	1
干 Stem	可溶性糖 Soluble sugar	1
	淀粉 Starch	0.543**	1
	NSC	0.839**	0.913**	1
	C	−0.179	−0.032	−0.108	1
	N	0.110	−0.016	0.043	−0.101	1
	P	0.175	−0.194	−0.041	0.100	−0.001	1
	C∶N	−0.255	0.040	−0.098	0.370	−0.930**	−0.008	1
	C∶P	−0.225	0.100	−0.045	−0.015	−0.170	−0.858**	0.168	1
	N∶P	0.047	0.162	0.128	−0.176	0.298	0.819**	−0.332	0.848**	1
枝 Branch	可溶性糖 Soluble sugar	1
	淀粉 Starch	0.226	1
	NSC	0.903**	0.623**	1
	C	−0.002	0.494*	0.216	1
	N	0.085	0.037	0.084	−0.387	1
	P	0.464	−0.009	0.369	−0.289	0.537*	1
	C∶N	−0.102	0.129	−0.025	0.705**	−0.911**	−0.543*	1
	C∶P	−0.454	0.074	−0.332	0.377	−0.591**	−0.942**	0.635**	1
	N∶P	−0.452	−0.067	−0.392	−0.127	0.081	−0.755**	−0.078	0.671**	1
根 Root	可溶性糖 Soluble sugar	1
	淀粉Starch	0.703**	1
	NSC	0.963**	0.868**	1
	C	0.241	0.226	0.253	1
	N	0.027	0.158	0.079	−0.077	1
	P	0.472*	0.211	0.409	0.270	−0.061	1
	C∶N	−0.110	−0.238	−0.167	0.272	−0.762**	−0.150	1
	C∶P	−0.441	−0.110	−0.349	0.161	−0.038	−0.816**	0.293	1
	N∶P	−0.153	0.180	−0.039	−0.175	0.769**	−0.483*	−0.570*	0.481*	1

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