旱雨季交替对入侵种拉关木与乡土种白骨壤整株碳氮分配及权衡策略的影响

doi:10.7525/j.issn.1673-5102.2026.02.012

摘要/Abstract

摘要：

植物碳（C）、氮（N）化学计量特征反映其资源分配策略，但现有研究常局限于单一器官，忽视了自然生境下整株与土壤系统的协同反馈。这一局限在受入侵威胁的红树林中尤为突出，制约了对种间竞争机制的深入理解。本研究以海南四必湾入侵种拉关木（Laguncularia racemosa）与乡土种白骨壤（Avicennia marina）为对象，探究旱、雨季纯林与混交林中叶、枝、根的C、N含量和C∶N（质量比），以及根际表层（0~10 cm）及亚表层（10~20 cm）土壤理化因子（全C、全N含量、C∶N、含水量、pH及电导率）的季节动态及关联格局。结果表明：相比于拉关木，白骨壤各器官始终维持显著高的N含量和低的C∶N，且其纯林表层土壤N含量在旱季显著高于拉关木纯林，而C∶N显著更低。与雨季相比，旱季显著增强拉关木根与枝间的化学计量关联，且其各器官C、N特征均与各土层理化因子紧密相关；相反，白骨壤仅在雨季表现出全株化学计量关联，而旱季其根与叶及枝间的关联消失，叶和枝化学计量特征转而与亚表层土壤理化因子（含水量、电导率或N含量）密切相关。树种混交在旱季增强白骨壤器官间C、N关联的普遍性，并提高叶片C含量；在雨季则显著提高拉关木叶片N含量并降低C∶N。综上，拉关木以较低的组织构建成本，及其在旱季维持与土壤资源的紧密关联，形成竞争优势。树种混交虽可增强白骨壤的器官间生理关联，却使拉关木更有效地缓解自身养分限制，从而可能扩大其入侵。因此，如果在进行生态修复的红树林群落中存在外来入侵植物，需警惕其通过种间互惠效应误导管理策略的制定，反而加速其扩张的潜在风险。

关键词: 拉关木, 白骨壤, 入侵植物, 旱雨季, 碳氮化学计量, 植物-土壤反馈, 功能性状关联

Abstract:

Plant carbon（C） and nitrogen（N） stoichiometry reflects resource allocation strategies. However， current researches often focus on individual organs， overlooking the critical feedback between whole-plant and soil systems in natural habitats. This oversight is particularly critical in invaded mangrove ecosystems， where it limits the understanding of interspecific competition mechanisms. Here， this study investigated the invasive Laguncularia racemosa and the native Avicennia marina in Hainan， China. The C and N contents and C-to-N ratios were measured across leaves， branches， and roots， alongside key physicochemical properties of rhizosphere surface（0-10 cm） and subsurface（10-20 cm） soils in monoculture and mixed stands during both dry and wet seasons. Results showed that A. marina consistently maintained significantly higher N contents and lower C-to-N ratios across all organs compared to L. racemosa. Additionally， surface soil N contents were significantly higher in A. marina monocultures than in L. racemosa stands during the dry season， while soil C-to-N ratios remained lower. The dry season significantly strengthened root-branch stoichiometric linkages in L. racemosa， where all organs showed tight correlations with soil physicochemical factors at both soil depths. Conversely， A. marina exhibited whole-plant synergy only in the wet season； in the dry season， root-leaf and root-branch stoichiometric relationships were decoupled， and leaf and branch traits shifted to correlate closely with subsurface soil factors. During dry season， species mixing promoted widespread C and N associations across organs in A. marina and increased leaf C contents， while significantly elevating leaf N contents and reducing C-to-N ratios in L. racemosa during the wet season. This study revealed that the competitive advantage of L. racemosa was driven by its lower tissue construction costs and sustained coupling with soil resources during dry season. Notably， while mixed stand improved the inter-organ physiological coordination of the native A. marina， they allowed L. racemosa to more efficiently alleviate its nutrient limitations， thereby amplifying its invasive dominance. These findings highlighted a critical risk for mangrove restoration： interspecific facilitation may inadvertently accelerate the expansion of invasive species， potentially undermining management strategies.

Key words: Laguncularia racemosa, Avicennia marina, invasive plant, dry and wet seasons, carbon and nitrogen stoichiometry, plant-soil feedback, functional trait relationship

中图分类号:

S718.5

杨华, 卢启航, 杜彦君, 王文娜. 旱雨季交替对入侵种拉关木与乡土种白骨壤整株碳氮分配及权衡策略的影响[J]. 植物研究, 2026, 46(2): 336-347.

Hua YANG, Qihang LU, Yanjun DU, Wenna WANG. Whole-plant Carbon and Nitrogen Allocation and Trade-off Strategies of Invasive Laguncularia racemosa and Native Avicennia marina across Dry and Wet Seasons[J]. Bulletin of Botanical Research, 2026, 46(2): 336-347.

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土层 Soil layer/cm	季节 Season	群落类型 Stand type	全氮 Total nitrogen/ （g·kg^-1）	全碳 Total carbon/ （g·kg^-1）	碳氮比 Carbon-to- nitrogen ratio	电导率 Electric conductivity/ （mS·cm^-1）	酸碱度 pH	含水量 Water content/%
［0，10）	雨季 Wet season	Lr-P	0.05±0.01^A*	4.27±0.63^A	93.43±8.21^A*	3.24±1.30^A	7.53±0.07^A	12.57±1.62^A
		Am-P	0.06±0.02^A*	3.25±0.53^A	55.55±0.37^B*	2.22±0.09^A	7.30±0.06^AB	12.50±2.58^A
		Lr-Am-M	0.08±0.02^A*	4.95±1.18^A	62.88±4.95^B*	2.42±0.16^A	7.20±0.12^B	13.62±2.37^A
	旱季 Dry season	Lr-P	0.32±0.04^B*	5.77±0.50^A	17.05±2.61^A*	5.75±4.09^A	7.37±0.15^A	15.25±5.09^A
		Am-P	0.62±0.07^A*	6.25±1.31^A	10.02±1.72^B*	5.07±2.01^A	7.27±0.27^A	17.52±2.88^A
		Lr-Am-M	0.39±0.02^B*	3.32±0.17^B	8.95±0.80^B*	3.48±1.07^A	7.13±0.07^A	15.98±1.30^A
［10，20］	雨季 Wet season	Lr-P	0.04±0.00^A*	2.76±0.54^A	79.14±10.57^A*	8.23±2.17^A	7.37±0.09^A	14.23±1.99^A
		Am-P	0.05±0.02^A*	3.21±1.64^A	49.41±7.10^B*	5.10±2.54^A	7.30±0.12^A	12.81±2.17^A
		Lr-Am-M	0.06±0.01^A*	3.71±0.82^A	56.70±2.78^B*	5.08±1.22^A	7.33±0.07^A	17.44±2.46^A
	旱季 Dry season	Lr-P	0.25±0.11^A*	3.80±0.99^A	17.55±1.64^A*	6.10±3.15^A	7.33±0.26^A	15.28±5.07^A
		Am-P	0.31±0.07^A*	2.81±0.69^A	9.47±1.34^B*	2.79±0.85^A	7.27±0.07^A	9.91±2.08^A
		Lr-Am-M	0.34±0.01^A*	2.84±0.17^A	8.50±0.29^B*	3.00±0.62^A	7.23±0.09^A	15.45±0.33^A

变量 Variable	全碳 Total carbon/（g·kg^-1）			全氮 Total nitrogen/（g·kg^-1）			碳氮比 Carbon-to-nitrogen ratio
变量 Variable	自由度 d_f	F	P	自由度d_f	F	P	自由度d_f	F	P
季节 Seasons	1	19.53	<0.01	1	3.84	0.06	1	0.10	0.75
树种 Tree species	1	2.14	0.15	1	228.83	<0.01	1	54.16	<0.01
器官 Organs	2	136.83	<0.01	2	130.68	<0.01	2	17.36	<0.01
季节×树种 Seasons×Tree species	1	0.12	0.73	1	2.35	0.13	1	0.02	0.88
季节×器官 Seasons×Organs	2	1.88	0.16	2	1.39	0.26	2	4.16	0.02
树种×器官 Tree species×Organs	2	1.01	0.37	2	19.14	<0.01	2	5.73	<0.01
季节×树种×器官 Seasons×Tree species×Organs	2	1.37	0.26	2	1.36	0.27	2	3.84	0.03

土壤深度 Soil depth/cm	季节 Season	树种 Tree species	关联类型 Relationship type
土壤深度 Soil depth/cm	季节 Season	树种 Tree species	叶 Leaf	土壤 Soil	相关系数 r	枝 Branch	土壤 Soil	相关系数 r	根 Root	土壤 Soil	相关系数 r
［0，10）	雨季 Wet season	拉关木Lr	N	pH	-0.883*	C	N	0.971**	C∶N	含水量	0.829*
			C∶N	pH	0.883*	C	C	1.000**
						C	含水量	0.943**
		白骨壤Am			ns	C	N	0.928**	C	电导率	0.829*
						C	含水量	0.886*	N	电导率	-0.943**
									C∶N	电导率	0.829*
									C∶N	pH	-0.883*
	旱季 Dry season	拉关木Lr	C	电导率	0.841*	C	N	-0.886*	N	C	-0.829*
			C	含水量	0.829*				N	C∶N	-0.829*
			C∶N	C	0.829*				C∶N	C	0.829*
			C∶N	C∶N	0.829*				C∶N	C∶N	0.829*
		白骨壤Am			ns			ns			ns
［10，20］	雨季 Wet season	拉关木Lr			ns			ns	C∶N	C	0.829*
		拉关木Lr			ns			ns	C∶N	含水量	0.943**
		白骨壤Am			ns	C	N	0.943*	C	电导率	-0.829*
		白骨壤Am			ns	C	C	0.829*
	旱季 Dry season	拉关木Lr	C	C	0.829*	C	C∶N	0.829*	N	C∶N	-0.829*
		拉关木Lr	C∶N	C∶N	0.829*				C∶N	C∶N	0.829*
		白骨壤Am	C∶N	含水量	0.886*	C	N	0.829*			ns
		白骨壤Am				C∶N	电导率	-0.886*			ns