林下植被演替过程中毛竹枝叶形态质量和抽枝展叶效率变化特征

董亚文; 谢燕燕; 陈双林; 郭子武; 张景润; 汪舍平

doi:10.11913/PSJ.2095-0837.22272

林下植被演替过程中毛竹枝叶形态质量和抽枝展叶效率变化特征

1.
中国林业科学研究院亚热带林业研究所，杭州 311400
2.
浙江省常山县林业技术推广站，浙江常山 324200

基金项目: 浙江省重点研发计划项目（2020C02008）

详细信息

作者简介:
董亚文（1996−），女，硕士研究生，研究方向为竹林结构与功能研究（E-mail：Arwend@163.com）

通讯作者:
陈双林: E-mail：cslbamboo@126.com

中图分类号: Q948
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出版历程
- 收稿日期: 2022-10-31
- 修回日期: 2022-12-22
- 网络出版日期: 2023-09-06
- 刊出日期: 2023-08-30

Characteristics of Phyllostachys edulis (Carrière) J. Houz shoot morphology, branching, and leaf expansion efficiency during understory vegetation succession

1.
Research Institute of Subtropical Forestry, Chinese Academy of Forestry, Hangzhou 311400, China
2.
Forestry Technology Promotion Station of Changshan County, Changshan, Zhejiang 324200, China

Funds: This work was supported by a grant from the Key R&D Projects in Zhejiang Province (2020C02008).

摘要

摘要:
本研究分别选取林下植被演替前毛竹（Phyllostachys edulis (Carrière) J. Houz）纯林经营、竹材6~8年采伐一次形成的林下植被演替9、21年以及纯林（对照）等3类毛竹林，调查分析了1度、2度竹的主侧枝枝叶形态质量、抽枝展叶效率及生物量异速生长关系等。结果显示：（1）随林下植被演替的进行，1度竹主枝叶片主要形态质量性状呈降低趋势，2度竹则呈“V”型变化；（2）1度、2度竹主侧枝的枝长、纤细率显著提高，枝基径、枝干物质含量、枝干重等性状呈倒“V”型或升高趋势，枝形态变化较枝质量变化明显；（3）1度竹抽枝展叶效率指标呈“V”型变化，而2度竹主侧枝抽枝展叶效率显著降低，胡波尔值明显提高；（4）一定演替年限内，立竹主侧枝枝叶生物量生长关系从等速生长向异速生长变化，尔后恢复为等速生长。可见，林下植被演替对毛竹林立竹枝叶形态质量、抽枝展叶效率和枝叶生物量异速生长关系均产生重要影响，叶片趋于变小，枝条趋于细长，且具有明显的演替年限效应和立竹年龄效应，主要对主枝叶片产生影响，1度竹较2度竹敏感；演替过程中立竹将更多的资源投资于枝的生长，以增强空间资源的竞争能力。
- 毛竹 /
- 林下植被演替 /
- 枝叶形态质量 /
- 抽枝展叶效率 /
- 异速生长关系
Abstract:
This research examined three types of moso bamboo stands: pure moso bamboo forest prior to understory vegetation succession, understory vegetation succession formed by harvesting bamboo timber once every 6–8 years over periods of 9 and 21 years, and pure forest (control). We investigated the morphological quality of the main and lateral branches and leaves, efficiency of pumping and spreading, and biomass anisotropic growth relationship of 1st- and 2nd-degree bamboo. Results showed that: (1) With the succession of understory vegetation, the primary morphological and quality traits of the main branch leaves of 1st-degree bamboo showed a downward trend, while 2nd-degree bamboo showed a significant “V”-shaped change; (2) Length and slenderness of the main and lateral branches of 1st- and 2nd-degree bamboo increased significantly, while branch diameter, branch dry matter content, and branch dry weight showed an inverted “V”-shaped or increasing trend, with the changes in branch morphology being more pronounced than those in branch quality; (3) The leaf spreading efficiency index of 1st-degree bamboo showed a “V”-shaped change, whereas that of the main and lateral branches of 2nd-degree bamboo decreased significantly, with the Huber value increasing significantly; (4) Within a certain succession period, the biomass growth relationship of the main and lateral branches of the standing bamboo transitioned from isochronous to anisotropic growth, then reverted to isochronous growth. Thus, understory vegetation succession exhibited an important effect on the morphological quality of bamboo branches and leaves, efficiency of branching and spreading, and relationship between anisotropic growth and branch and leaf biomass in moso bamboo forests. Notably, leaves tended to be smaller and branches slenderer. Furthermore, successional age and age of standing bamboo had obvious effects, mainly on the leaves of the main branches, with 1st-degree bamboo being more sensitive than 2nd-degree bamboo. Hence, during the succession process, bamboo invests more resources in branch growth to enhance competition for spatial resources.
- Phyllostachys edulis /
- Undergrowth vegetation succession /
- Shape of branches and leaves /
- Shooting and leaf developing efficiency /
- Allometric growth relationship

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图 1 毛竹林叶干重与枝干重异速生长关系

A：1度竹主枝；B：1度竹侧枝；C：2度竹主枝；D：2度竹侧枝。

Figure 1. Relationship between total leaf mass and stem mass of experimental Phyllostachys edulis stands

A: Main branches of 1st-degree bamboo; B: Lateral branches of 1st-degree bamboo; C: Main branches of 2nd-degree bamboo; D: Lateral branches of 2nd-degree bamboo.

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表 1 实验毛竹林基本情况

Table 1 Basic information on experimental Phyllostachys edulis stands

演替年限 Succession period / a	海拔 Altitude / m	坡向 Slope aspect	坡位 Slope position	毛竹 P. edulis		乔灌木 Arbor and shrubs			郁闭度 Canopy density
演替年限 Succession period / a	海拔 Altitude / m	坡向 Slope aspect	坡位 Slope position	立竹密度 Bamboo density / ind/hm²	立竹胸径 Bamboo diameter at breast height / cm	密度 Density / ind/hm²	树高 Tree height / m	地径 Basal diameter / cm	郁闭度 Canopy density
0	500 ± 10	36 ± 2.50	西	2 625 ± 25	9.40 ± 0.60	－	－	－	0.60
9	520 ± 10	36 ± 2.50	西	3 150 ± 30	9.20 ± 0.40	8 000 ± 55	0.76 ± 0.08	0.76 ± 0.07	0.70
21	490 ± 10	36 ± 2.50	西	2 275 ± 20	8.90 ± 0.40	27 511 ± 147	2.42 ± 0.20	1.69 ± 0.11	0.85

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表 2 毛竹林立竹叶片形态质量性状

Table 2 Leaf morphological characteristics of experimental Phyllostachys edulis stands

演替年限 Succession period / a	立竹年龄 Bamboo age	枝类 Branch class	叶长 Leaf length / cm	叶宽 Leaf width / cm	叶周长 Leaf circumference / cm	叶干重 Leaf dry weight / g	单叶面积 Leaf area / cm²	比叶面积 Specific leaf area / cm²/g	叶干物质含量 Leaf dry matter content / g/g
21	1	主枝	9.63 ± 0.28B	1.44 ± 0.03B	22.02 ± 0.30B	33.94 ± 1.79A	10.31 ± 0.30B	195.53 ± 2.61A	0.12 ± 0.00A
9			9.88 ± 0.45B	1.45 ± 0.03B	22.27 ± 0.91B	34.23 ± 1.08A	10.60 ± 0.67B	174.21 ± 1.73B	0.14 ± 0.01A
0			10.67 ± 0.31A	1.52 ± 0.03A	24.18 ± 0.48A	30.51 ± 0.07B	11.97 ± 0.34A	200.31 ± 5.13A	0.13 ± 0.01A
21		侧枝	9.94 ± 0.21A	1.46 ± 0.03A	22.65 ± 0.45A	16.93 ± 0.68A	10.81 ± 0.24A	200.03 ± 11.11A	0.12 ± 0.00A
9			10.03 ± 0.26A	1.47 ± 0.03A	22.54 ± 0.69A	18.55 ± 0.16A	10.78 ± 0.39A	169.83 ± 5.15B	0.13 ± 0.01A
0			10.20 ± 0.20A	1.47 ± 0.04A	23.18 ± 0.39A	19.52 ± 2.66A	11.20 ± 0.40A	206.76 ± 8.14A	0.12 ± 0.01A
21	2	主枝	10.66 ± 0.08A	1.54 ± 0.03A	24.26 ± 0.15A	24.37 ± 1.94A	12.34 ± 0.09A	190.20 ± 0.13A	0.15 ± 0.01A
9			9.96 ± 0.12B	1.43 ± 0.04B	22.31 ± 0.02B	19.09 ± 0.74B	10.66 ± 0.23B	182.36 ± 1.95A	0.13 ± 0.01B
0			10.79 ± 0.17A	1.56 ± 0.06A	24.24 ± 0.38 A	27.18 ± 2.00A	12.15 ± 0.19A	191.78 ± 16.56A	0.13 ± 0.00A
21		侧枝	10.56 ± 0.13A	1.49 ± 0.02A	23.41 ± 0.51AB	16.18 ± 1.40B	11.71 ± 0.19A	188.60 ± 0.13A	0.14 ± 0.00A
9			10.02 ± 0.27B	1.47 ± 0.00A	22.58 ± 0.56B	11.02 ± 0.51C	10.98 ± 0.41A	184.19 ± 4.80A	0.15 ± 0.03A
0			10.71 ± 0.20A	1.48 ± 0.02A	24.20 ± 0.53A	19.08 ± 0.89A	11.67 ± 0.48A	190.47 ± 4.64A	0.14 ± 0.00A
注：不同大写字母表示不同林下植被演替年限毛竹林同一年龄立竹主枝或侧枝叶片间差异显著。下同。
Note: Different capital letters indicate different vegetation succession years, with significant differences between the leaves of main and side branches of P. edulis stands of the same age. Same below.

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表 3 毛竹林立竹枝形态质量性状

Table 3 Branch morphological characteristics of experimental Phyllostachys edulis stands

演替年限 Succession period / a	立竹年龄 Bamboo age	枝类 Branch class	枝长 Branch length / cm	枝基径 Branch base diameter / mm	枝干重 Branch weight / g	枝纤细率 Branch fineness rate / cm/mm	枝干物质含量 Branch dry matter content / g/g
21	1	主枝	159.78 ± 10.10A	9.09 ± 0.05A	75.00 ± 1.96A	176.12 ± 6.24A	0.64 ± 0.02A
9			157.22 ± 2.98A	9.18 ± 0.18A	60.20 ± 3.35B	176.05 ± 11.18A	0.64 ± 0.01A
0			143.25 ± 3.47B	8.82 ± 0.27A	55.31 ± 2.25B	163.74 ± 1.19B	0.62 ± 0.02A
21		侧枝	132.67 ± 5.39A	7.12 ± 0.23A	36.82 ± 4.36A	187.16 ± 2.95A	0.64 ± 0.02B
9			128.94 ± 1.59A	6.94 ± 0.07A	36.56 ± 0.52A	189.03 ± 10.30A	0.69 ± 0.05A
0			123.48 ± 6.46B	7.16 ± 0.47A	37.70 ± 3.81A	178.30 ± 6.70B	0.63 ± 0.01B
21	2	主枝	155.42 ± 6.61A	8.23 ± 0.14A	57.75 ± 8.71A	189.53 ± 7.49A	0.61 ± 0.02A
9			137.39 ± 1.42B	7.70 ± 0.14B	43.20 ± 3.95B	180.42 ± 0.70A	0.66 ± 0.01A
0			139.22 ± 1.13B	7.75 ± 0.07B	46.49 ± 3.31B	178.86 ± 5.05B	0.64 ± 0.01A
21		侧枝	137.50 ± 6.50A	6.74 ± 0.19A	32.52 ± 3.55A	203.41 ± 1.92A	0.64 ± 0.01A
9			124.11 ± 2.96B	6.03 ± 0.07B	23.63 ± 0.55B	204.49 ± 4.77A	0.66 ± 0.01A
0			121.31 ± 10.73B	6.59 ± 0.58A	29.60 ± 3.56A	183.98 ± 4.02B	0.64 ± 0.03A

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表 4 毛竹林抽枝展叶效率

Table 4 Branch and leaf characters of experimental Phyllostachys edulis stands

演替年限 Succession period / a	立竹年龄 Bamboo age	枝类 Branch class	出叶强度 Intensity of leaf / ind/g	叶面积比 Leaf area ratio / mm²/g	叶密度 Leaf density / ind/mm	叶枝质量比 Leaf /stem mass ratio / g/g	胡波尔值 Huber value / mm²/mm
21	1	主枝	6.35 ± 0.24A	100.12 ± 5.72B	4.34 ± 0.54A	0.53 ± 0.03B	0.00008 ± 0B
9			5.65 ± 0.56B	93.27 ± 14.77B	3.59 ± 0.76B	0.55 ± 0.08AB	0.00012 ± 0A
0			6.12 ± 0.43AB	117.82 ± 29.07A	3.87 ± 0.70AB	0.61 ± 0.07A	0.00009 ± 0AB
21		侧枝	6.04 ± 0.44B	99.34 ± 10.89B	2.58 ± 0.34A	0.52 ± 0.06B	0.00011 ± 0B
9			5.17 ± 0.28C	84.42 ± 6.93C	2.24 ± 0.30A	0.51 ± 0.04B	0.00012 ± 0A
0			6.68 ± 0.84A	121.27 ± 23.79A	2.73 ± 1.07A	0.61 ± 0.12A	0.00011 ± 0A
21	2	主枝	4.66 ± 0.34C	83.18 ± 8.66B	2.25 ± 0.32B	0.44 ± 0.05B	0.00012 ± 0A
9			5.30 ± 0.44B	84.18 ± 10.19B	2.54 ± 0.53B	0.49 ± 0.06B	0.00013 ± 0A
0			6.16 ± 0.47A	123.26 ± 15.38A	3.49 ± 0.76A	0.64 ± 0.08A	0.00009 ± 0B
21		侧枝	4.97 ± 0.30B	85.52 ± 7.59B	1.68 ± 0.49B	0.47 ± 0.04B	0.00013 ± 0A
9			4.94 ± 0.58B	82.79 ± 14.92 B	1.51 ± 0.27B	0.52 ± 0.09B	0.00014 ± 0A
0			6.72 ± 0.45A	136.35 ± 15.92A	2.67 ± 0.90A	0.73 ± 0.09A	0.00009 ± 0B

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表 5 毛竹林枝-叶质量的SMA分析

Table 5 SMA analysis of branch leaf size of experimental Phyllostachys edulis stands

指标 Indicators	演替年限 Succession period / a	立竹年龄 Bamboo age	枝类 Branch class	数量 Number	斜率 Slope	95%置信区间 95% CI	截距 Intercept	R²	P
叶干重- 枝干重	21	1	主枝	30	1.02(1.14)	0.71，1.45	−0.32	0.25	< 0.010
	9			30	1.26(1.14)	0.92，1.72	−0.74	0.48	< 0.001
	0			30	1.15(1.14)	0.87，1.51	−0.51	0.42	< 0.001
	21		侧枝	30	1.14(1.02)	0.85，1.51	−0.55	0.51	< 0.001
	9			30	1.66	1.18，2.32	−1.34	0.23	< 0.050
	0			30	0.94(1.02)	0.74，1.20	−0.19	0.61	< 0.001
	21	2	主枝	30	1.28(0.99)	0.91，1.81	−0.87	0.36	< 0.050
	9			30	1.62	1.23，2.15	−1.39	0.46	< 0.001
	0			30	0.97(0.99)	0.86，1.09	−0.19	0.89	< 0.001
	21		侧枝	30	1.16(1.13)	0.86，1.58	−0.58	0.42	< 0.001
	9			30	1.83	1.43，2.35	−1.53	0.70	< 0.001
	0			30	1.09(1.13)	0.82，1.45	−0.31	0.47	< 0.001

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