温带阔叶树种不同生活史阶段的资源获取能力和防御能力关系的变化

doi:10.11707/j.1001-7488.LYKX20240427

摘要/Abstract

摘要：

目的: 基于东北不同阔叶树种，探讨植物资源获取型与防御型性状的相关性及其驱动因素，揭示木本植物资源获取能力与防御能力的关联，解析木本植物对环境的动态适应机制，为森林生态系统管理提供科学依据。方法: 测定温带森林中5种不同耐荫性阔叶树种白桦、水曲柳、裂叶榆、紫椴、五角槭的叶片、新枝、老枝在3个不同生活史阶段（幼苗、幼树、成年树）的20个植物功能性状，将其划分为资源获取型和防御型性状，探讨木本植物2类性状的相互关联及影响因子。结果: 叶片资源获取能力与植株地上部分防御能力的相关性较弱，不同生活阶段资源获取能力与防御能力的相关性存在差异。幼苗期间，2种能力存在协作关系（夹角< 90°）；幼树期间，2种能力存在解耦关系（夹角≈ 90°）；成年树阶段，2种能力存在权衡关系（夹角> 90°）。综合考虑环境因子（土壤养分、土壤pH、土壤含水率）和植物特性（生活史阶段、耐荫性）对植物资源获取型和防御型性状的影响时，环境因子的影响（PC1 56%、PC2 73%）大于植物特性的影响（PC1 44%、PC2 27%）。结论: 植物在不同生活史阶段对资源获取与防御的协调策略具有阶段性变化，环境因子是主要驱动因素，但植物特性亦不容忽视。

关键词: 资源获取能力, 防御能力, 驱动因素, 环境因子, 生活史

Abstract:

Objective: In view of different broadleaved tree species in Northeast China, this study explored the correlation between plant resource acquisition traits and defense traits and their driving factors, to reveal the correlation between the resource acquisition ability and defense ability of woody plants, to analyze the dynamic adaptation mechanism of woody plants to the environment, and to provide a scientific basis for forest ecosystem management. Method: In this study, we measured 20 phytofunctional traits in leaves, new twigs, and old branchs of five broadleaf species with different shade tolerance in temperate forests, including Betula platyphylla, Fraxinus mandshurica, Ulmus laciniata, Tilia amurensis, and Acer pictum subsp. mono, at three different life-history stages (seedling, sapling, and mature tree), and these traits were divided into resource-acquisition traits and defense traits, to explore the interrelationships and influencing factors of the two types of traits in woody plants. Result: There was weak correlation between leaf resource acquisition capacity and aboveground defense capacity, and the correlation between resource acquisition capacity and defense capacity varied at different life stages. At the seedling stage, the two capacities had a collaborative relationship (angle<90°); at the sapling stage, the two capacities were decoupled (angle≈90°); and at the mature tree stage, the two capacities had a trade-off relationship (angle>90°). Combining the effects of environmental factors (soil nutrients, soil pH, soil water content) and plant characteristics (life history stages, shade tolerance) on plant resource acquisition traits and defense traits, it was found that environmental factors (PC1 56%, PC2 73%) had a stronger effect on the two types of traits than plant characteristics (PC1 44%, PC2 27%). Conclusion: The coordination strategies between resource acquisition and defense vary across different growth stages. While environmental factors serve as the primary driving forces, plant characteristics also play a crucial role.

Key words: resource acquisition capacity, defense capacity, drivers, environmental factors, life history

中图分类号:

S718.5

王凯波,金光泽,刘志理. 温带阔叶树种不同生活史阶段的资源获取能力和防御能力关系的变化[J]. 林业科学, 2025, 61(8): 219-230.

Kaibo Wang,Guangze Jin,Zhili Liu. Changes of Resource Acquisition and Defense Capabilities of Temperate Broadleaf Tree Species at Different Life History Stages[J]. Scientia Silvae Sinicae, 2025, 61(8): 219-230.

图/表 6

表1

植物叶片、新枝、老枝功能性状及其类别"

器官 Organ	功能性状 Functional traits	单位 Units	生态学意义 Ecological meaning	生态策略 Ecological strategy
叶片 Leaf	叶绿素含量 Chlorophyll content	—	与植物光合速率相关，可表征植物的光合能力、生长状况以及胁迫状况 Correlates with plant photosynthetic rate and characterizes plant photosynthetic capacity, growth, and stress conditions	资源获取 Resource acquisition
	比叶质量 Leaf mass per area	g·cm^?2	叶片结构特征的综合指标，可表征植物的资源获取能力及对环境的响应策略 A composite index of leaf structural characteristics that characterizes the resource acquisition capacity of plants and their response strategies to the environment	资源获取 Resource acquisition
	碳含量 Carbon content	g·kg^?1	构成生命体的基本元素，维持植物生长发育和代谢过程的关键元素，植物元素含量的变异及分配规律可表征植物对环境的响应与适应机制 The basic elements that constitute living organisms, the key elements that maintain plant growth， development and metabolic processes, and the variation and distribution patterns of plant elemental content can characterize the plant’s response and adaptation mechanisms to the environment	资源获取 Resource acquisition
	氮含量 Nitrogen content	g·kg^?1		资源获取 Resource acquisition
	磷含量 Phosphorus content	g·kg^?1		资源获取 Resource acquisition
	总酚含量 Total phenolic content	g·kg^?1	能够抵御外界生物（草食动物啃食）和非生物（低温）胁迫，在植物生长、发育和防御中发挥重要作用 Able to withstand external biotic (herbivore nibbling) and abiotic (low temperatures) stresses and plays an important role in plant growth, development and defense	防御 Defense
	单宁含量 Tannin content	g·kg^?1		防御 Defense
	类黄酮含量 Flavonoid content	g·kg^?1		防御 Defense
新枝 New twig	碳含量 Carbon content	g·kg^?1	构成生命体的基本元素，维持植物生长发育和代谢过程的关键元素，植物元素含量的变异及分配规律可表征植物对环境的响应与适应机制 The basic elements that constitute living organisms, the key elements that maintain plant growth, development and metabolic processes, and the variation and distribution patterns of plant elemental content can characterize the plant’s response and adaptation mechanisms to the environment	资源获取 Resource acquisition
	氮含量 Nitrogen content	g·kg^?1		资源获取 Resource acquisition
	磷含量 Phosphorus content	g·kg^?1		资源获取 Resource acquisition
	总酚含量 Total phenolic content	g·kg^?1	能够抵御外界生物（草食动物啃食）和非生物（低温）胁迫，在植物生长、发育和防御中发挥重要作用 Able to withstand external biotic (herbivore nibbling) and abiotic (low temperatures) stresses and plays an important role in plant growth, development and defense	防御 Defense
	单宁含量 Tannin content	g·kg^?1		防御 Defense
	类黄酮含量 Flavonoid content	g·kg^?1		防御 Defense
老枝 Old branch	碳含量 Carbon content	g·kg^?1	构成生命体的基本元素，维持植物生长发育和代谢过程的关键元素，植物元素含量的变异及分配规律可表征植物对环境的响应与适应机制 The basic elements that constitute living organisms, the key elements that maintain plant growth, development and metabolic processes, and the variation and distribution patterns of plant elemental content can characterize the plant’s response and adaptation mechanisms to the environment	资源获取 Resource acquisition
	氮含量 Nitrogen content	g·kg^?1		资源获取 Resource acquisition
	磷含量 Phosphorus content	g·kg^?1		资源获取 Resource acquisition
	总酚含量 Total phenolic content	g·kg^?1	能够抵御外界生物（草食动物啃食）和非生物（低温）胁迫，在植物生长、发育和防御中发挥重要作用 Able to withstand external biotic (herbivore nibbling) and abiotic (low temperatures) stresses and plays an important role in plant growth, development and defense	防御 Defense
	单宁含量 Tannin content	g·kg^?1		防御 Defense
	类黄酮含量 Flavonoid content	g·kg^?1		防御 Defense

表1

图1

图2

资源获取型与防御型性状间的主成分分析 a：幼苗阶段不同耐荫性树种资源获取与防御型性状的主成分分析结果 Principal component analysis results of resource acquisition and defense traits of different negative tolerant tree species at seedling stage；b：幼树阶段不同耐荫性树种资源获取与防御型性状的主成分分析结果 Principal component analysis results of resource acquisition and defense traits of different negative tolerant tree species at sapling stage；c：成年树阶段不同耐荫性树种资源获取与防御型性状的主成分分析结果 Principal component analysis results of resource acquisition and defense traits of different negative tolerant tree species in mature tree stage。SPAD：叶绿素含量指数Chlorophyll content (soil and plant analyzerdevelopment)；LMA：比叶质量 Leaf mass per area；LC：叶片碳含量 Leaf carbon content；LN：叶片氮含量 Leaf nitrogen content；LP：叶片磷含量 Leaf phosphorus content；LTP：叶片总酚含量 Leaf total phenolic content；LTA：叶片单宁含量 Leaf tannin content；LFLA：叶片类黄酮含量 Leaf flavonoid content；NC：新枝碳含量 New twig carbon content；NN：新枝氮含量 New twig nitrogen content；NP：新枝磷含量 New twig phosphorus content；NTP：新枝总酚含量 New twig total phenolic content；NTA：新枝单宁含量 New twig tannin content；NFLA：新枝类黄酮含量 New twig flavonoid content；OC：老枝碳含量 Old branch carbon content；ON：老枝氮含量 Old branch nitrogen content；OP：老枝磷含量 Old branch phosphorus content；OTP：老枝总酚含量 Old branch total phenolic content；OTA：老枝单宁含量 Old branch tannin content；OFLA：老枝类黄酮含量 Old branch flavonoid content。其中，蓝色荫影代表不耐荫树种（白桦、水曲柳）95%的置信区间，黄色荫影代表耐荫树种（裂叶榆、紫椴、五角槭）95%的置信区间。若性状间夹角小于90°，正相关；若性状间夹角等于90°，不相关；若性状间夹角大于90°，负相关。Among them， the blue shade represents the 95% confidence interval of the shad intolerant tree species （B. platyphylla， F. mandshurica）， the yellow shade represents the 95% confidence interval of the shad tolerant tree species (U. laciniata， T. amurensis， A. pictum subsp. mono). If the angle between traits is less than 90°， positive correlation；if the angle between traits is equal to 90°， no correlation；if the angle between traits is greater than 90°， negative correlation."

图2

图3

图4

表1

样树状况以及不同生活史阶段叶、新枝、老枝性状基本信息①"

物种 Species	生活史 Life history	器官 Organ	SW/ （g·g^?1）	SC/ （g·kg^?1）	SN/ （g·kg^?1）	SP / （g·kg^?1）	pH	H / m	DBH/ cm	SPAD	LMA / （g·cm^?2）	C / （g·kg^?1）	N / （g·kg^?1）	P / （g·kg^?1）	TP/ （g·kg^?1）	TA/ （g·kg^?1）	FLA/ （g·kg^?1）
白桦 B. platyphylla	幼苗 Seedling n=10	L	0.91±0.45	62.57±24.72	5.47±1.38	1.06±0.21	4.31±0.12	6.30±0.79	5.13±0.25	39.33±2.48	0.004±0.001	465.38±28.78	40.15±8.38	4.02±1.17	28.57±11.05	24.02±10.84	95.2±21.65
		NT								—	—	487.42±26.18	20.57±3.16	1.31±0.10	38.82±4.15	34.08±3.84	228.61±33.91
		OB								—	—	410.21±27.13	11.12±1.83	1.16±0.14	39.76±4.37	36.06±3.98	193.55±21.90
	幼树 Sapling n=10	L	1.23±0.44	138.41±39.00	9.28±2.11	1.24±0.26	4.33±0.32	17.42±1.07	18.83±1.37	38.90±2.61	0.003±0.001	474.11±34.61	28.43±2.07	1.90±0.43	34.92±12.66	31.01±12.30	101.93±19.27
		NT								—	—	484.72±18.64	18.61±5.92	1.11±0.10	43.69±5..26	38.56±4.81	223.20±30.81
		OB								—	—	417.94±13.69	12.04±2.13	1.10±0.15	36.44±7.54	33.08±7.56	190.15±37.02
	成年树 Mature tree n=10	L	1.32±0.68	129.43±64.24	9.49±3.65	1.15±0.23	4.29±0.36	18.89±1.40	40.99±2.18	39.02±3.73	0.004±0.001	457.29±15.20	26.38±2.04	1.76±0.26	33.92±7.58	29.56±7.59	112.49±21.57
		NT								—	—	493.88±14.06	17.43±2.55	1.20±0.20	48.20±8.91	43.72±8.83	254.45±51.42
		OB								—	—	417.92±10.00	12.14±1.48	1.24±0.21	36.72±8.47	32.36±7.58	208.65±47.61
水曲柳 F. mandshurica	幼苗 Seedling n=10	L	1.13±0.63	107.74±45.24	7.68±2.37	1.43±0.34	4.51±0.60	7.04±0.88	4.83±0.66	54.18±27.19	0.011±0.003	411.74±20.60	32.43±4.51	2.24±0.44	8.79±3.76	7.34±3.01	36.42±16.86
		NT								—	—	468.04±18.10	11.53±1.44	0.75±0.24	32.44±6.39	21.51±4.76	86.41±29.79
		OB								—	—	391.15±34.55	9.92±1.58	0.85±0.23	26.48±5.68	19.02±4.09	70.45±20.58
	幼树 Sapling n=10	L	1.07±0.37	99.11±47.56	6.73±1.55	1.46±0.25	4.42±0.37	16.82±2.06	17.83±1.12	46.17±20.10	0.012±0.002	415.91±40.76	29.17±3.48	2.25±0.36	29.17±3.48	8.67±4.00	43.62±20.41
		NT								—	—	479.61±41.73	11.75±1.92	0.92±0.12	44.15±5.21	29.25±3.68	152.79±29.89
		OB								—	—	401.62±17.26	9.35±1.23	0.70±0.15	33.44±5.34	24.06±3.98	87.94±18.47
	成年树 Mature tree n=10	L	1.04±0.37	91.12±31.88	7.33±1.93	1.39±0.30	4.69±0.54	21.72±1.24	42.05±1.42	38.84±2.45	0.012±0.002	419.17±9.54	29.31±3.09	2.34±0.23	14.54±8.21	11.17±6.05	58.15±36.27
		NT								—	—	468.52±7.47	12.57±1.18	0.90±0.12	46.09±5.42	30.68±3.52	171.98±37.24
		OB								—	—	401.57±11.20	10.68±1.31	0.86±0.12	35.20±5.99	24.82±4.59	103.38±15.06
裂叶榆 U. laciniata	幼苗 Seedling n=10	L	0.97±0.67	114.58±63.72	7.05±2.85	1.23±0.48	4.61±0.40	4.73±0.88	3.80±0.62	42.57±4.75	0.010±0.002	365.23±32.23	31.87±5.75	2.58±0.63	24.82±16.96	18.28±15.65	82.71±69.72
		NT								—	—	459.02±16.08	20.90±3.58	1.38±0.17	36.90±3.35	31.75±3.39	103.29±18.31
		OB								—	—	389.63±19.15	13.91±1.20	1.52±0.15	30.02±3.18	23.80±2.81	90.00±19.61
	幼树 Sapling n=10	L	1.04±0.22	153.90±34.36	10.02±1.32	1.50±0.28	5.33±0.39	12.20±1.94	18.31±1.14	44.10±3.03	0.013±0.002	380.06±27.91	31.66±7.87	2.54±0.81	28.71±10.67	21.71±9.89	88.88±35.48
		NT								—	—	448.07±21.71	22.94±2.89	1.55±0.16	39.42±3.74	33.98±2.95	125.44±39.78
		OB								—	—	376.80±10.16	15.39±2.71	1.65±0.23	32.42±5.37	25.54±4.27	98.53±26.28
	成年树 Mature tree n=10	L	1.40±0.73	193.06±54.22	10..21±3.09	1.52±0.30	5.39±0.48	16.66±1.87	41.30±0.90	45.19±2.23	0.014±0.003	385.21±18.68	30.99±3.87	2.32±0.38	29.29±7.64	21.83±6.92	94.72±34.84
		NT								—	—	446.03±9.05	21.81±1.71	1.52±0.07	33.21±8.00	27.67±7.87	121.83±26.41
		OB								—	—	386.92±8.27	14.84±1.25	1.53±0.09	31.45±4.83	25.49±4.43	90.28±30.12
紫椴 T. amurensis	幼苗 Seedling n=10	L	1.10±0.56	92.53±43.39	7.12±1.96	1.34±0.29	4.49±0.49	4.58±0.94	4.37±0.61	32.51±2.61	0.008±0.002	465.42±21.21	34.03±3.13	2.93±0.60	27.26±17.39	23.52±16.77	113.79±87.03
		NT								—	—	460.06±16.32	13.72±1.61	0.94±0.15	10.93±5.15	7.91±4.46	159.47±37.81
		OB								—	—	394.17±14.63	10.22±1.36	1.11±0.15	20.32±5.18	18.51±4.78	63.47±15.14
	幼树 Sapling n=10	L	0.83±0.17	87.77±26.62	6.88±1.43	1.25±0.22	4.34±0.51	12.60±2.41	17.70±1.48	40.77±3.42	0.011±0.003	460.19±30.84	33.83±5.95	2.51±0.31	25.93±11.59	23.02±10.88	107.63±43.09
		NT								—	—	482.02±22.37	15.15±1.30	1.11±0.15	20.82±7.08	16.00±7.20	223.24±53.28
		OB								—	—	395.44±19.37	11.35±1.43	1.22±0.13	21.99±6.10	20.21±6.01	74.27±22.69
	成年树 Mature tree n=10	L	1.09±0.26	142.34±55.45	9.30±2.09	1.24±0.29	5.56±0.38	18.92±2.60	43.04±2.80	41.70±4.71	0.011±0.002	448.10±14.76	33.41±3.42	2.62±0.23	29.58±8.08	26.11±7.11	102.48±35.54
		NT								—	—	465.33±13.17	14.09±1.75	1.06±0.17	22.07±8.56	20.01±8.83	214.13±49.36
		OB								—	—	397.20±7.93	11.07±1.44	1.24±0.08	25.00±7.94	23.11±7.65	92.00±31.89
五角槭 A. pictum subsp. mono	幼苗 Seedling n=5	L	0.70±0.13	102.59±27.80	6.24±1.09	1.13±0.17	5.18±0.43	4.62±1.15	4.04±0.72	39.75±2.50	0.007±0.001	444.38±26.59	36.01±3.04	2.22±0.52	39.62±12.83	33.41±11.96	108.61±46.61
		NT								—	—	479.64±20.50	12.25±3.90	0.81±0.26	49.30±7.13	42.30±7.41	168.66±30.97
		OB								—	—	410.31±16.40	8.93±1.79	0.81±0.27	39.23±8.46	34.16±7.66	169.88±39.02
	幼树 Sapling n=7	L	1.22±0.46	142.65±50.19	8.57±1.29	1.49±0.25	4.69±0.67	11.26±1.09	17.51±1.74	42.11±3.94	0.010±0.002	426.09±19.81	39.03±6.63	2.49±0.74	65.03±21.48	57.17±21.05	217.57±69.75
		NT								—	—	483.04±18.90	11.24±2.63	1.04±0.24	55.55±11.85	48.87±12.33	198.68±62.76
		OB								—	—	411.66±15.94	8.92±1.65	0.93±0.21	37.76±7.19	32.98±7.25	156.23±36.49
	成年树 Mature tree n=10	L	1.13±0.21	113.06±28.57	6.36±1.40	1.08±0.20	5.74±0.34	14.00±3.00	41.32±2.20	42.21±2.56	0.010±0.002	430.62±20.17	35.32±4.78	2.63±0.25	59.71±15.70	52.29±14.72	175.79±35.81
		NT								—	—	476.17±9.66	11.70±0.69	1.05±0.09	54.24±6.88	47.34±6.65	199.33±30.51
		OB								—	—	402.10±6.90	9.81±0.51	0.96±0.05	41.60±5.67	36.61±5.43	177.85±39.22

表1

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