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    雌雄异株植物东北鼠李生态化学计量特征的性别二态性

    Sexual dimorphism in the ecological stoichiometric characteristics of dioecious Rhamnus schneideri var. manshurica

    • 摘要:
      目的 植物的生态化学计量特征与生态内稳性是其适应环境变化的关键体现。鉴于雌雄异株植物繁殖成本存在性别分化,其资源分配策略在不同物候期及年际间可能呈现动态的性别二态性。揭示这一性别二态性对理解雌雄异株植物在环境波动中的适应性具有重要意义。本研究以雌雄异株植物东北鼠李为研究对象,探究其在不同生长阶段叶片碳(C)、氮(N)、磷(P)生态化学计量特征及生态内稳性的性别二态性,基于繁殖成本理论解析性别差异,并结合环境因子的年际变化分析生境变化对于东北鼠李性别二态性的影响。
      方法 依托吉林蛟河针阔混交林长期固定样地,于2021年和2025年的花期与果期,采集东北鼠李雌雄植株叶片及对应根际表层土壤样品。利用激光高光谱元素分析仪测定叶片C、N、P含量,结合土壤全量养分数据,采用Wilcoxon秩和检验及重抽样置换检验分析生态化学计量特征的性别差异,并通过线性回归模型计算内稳性指数,评估其生态内稳性特征。
      结果 (1)2021年花期,雌株叶片C含量和C∶N比显著高于雄株(p < 0.05),C∶P、N∶P比值极显著高于雄株(p < 0.001),P含量极显著低于雄株(p < 0.001);果期雌株叶片C含量以及C∶N、C∶P、N∶P比值极显著低于雄株(p < 0.001),而N、P含量极显著高于雄株(p < 0.001)。2025年两性差异减弱,仅花期雌株C、N含量显著高于雄株(p < 0.05),其余计量特征均无显著性别差异。(2)生态内稳性分析表明,2021年雄株花期C元素以及花期、果期C∶N比值呈现弱稳态,表现出一定的环境敏感性;果期C元素呈现稳态,P元素呈现弱稳态,其余指标均维持在绝对稳态;雌株仅果期C∶N比值呈现稳态,其余指标均维持在绝对稳态。2025年随着水热条件改善,两性植株的所有生态内稳性指标均维持在绝对稳态。
      结论 东北鼠李雌雄植株的化学计量特征及生态内稳性存在性别二态性,其根本原因在于两性繁殖成本的分化。雌株因承担果实与种子发育的高能耗任务,在果期通过主动富集N、P元素提升叶片光合效率,以补偿繁殖消耗并维持较高的生态内稳性;雄株繁殖投入主要集中在花期,其内稳性相对易受环境影响。这种由繁殖成本驱动的资源分配策略差异,是雌雄异株植物性别二态性的重要形成机制。

       

      Abstract:
      Objective The ecophysiological characteristics and ecological homeostasis of plants are key indicators of their abaptability to environmental changes. Given sex-specific differences in reproductive costs in dioecious plants, resource allocation strategies may exhibit dynamic sexual dimorphism across phenological stages and between years. Uncovering this sexual dimorphism is of crucial for understanding the adaptability of dioecious plants to environmental fluctuations. This study investigated sexual dimorphism in the leaf carbon (C), nitrogen (N), and phosphorus (P) stoichiometry and ecological homeostasis of the dioecious plant Rhamnus schneideri var. manshurica across different growth stages. Based on reproductive cost theory, we analyzed these sexual differences and examined the impact of habitat changes on the sexual dimorphism of Rhamnus schneideri var. manshurica by assessing interannual variations in environmental factors.
      Method Leaf samples from male and female Rhamnus schneideri var. manshurica plants, along with corresponding samples, were collected during the flowering and fruiting periods in 2021 and 2025 from a long-term permanent plot in a needle-broadleaved mixed forest in Jiaohe, Jilin Province. Leaf C, N and P contents were determined using a laser hyperspectral elemental analyzer. Combined with soil total nutrient data, the Wilcoxon rank-sum test and permutation test were employed to analyze sexual differences in ecological stoichiometric characteristics, and a linear regression model was used to calculate the homeostatic index to evaluate ecological homeostasis.
      Result (1) During 2021 flowering period, female plants exhibited significantly higher leaf C content and C∶N ratio (p < 0.05), extremely significantly higher C∶P and N∶P ratios (p < 0.001), but extremely significantly lower P content (p < 0.001) than male plants. In the fruiting period of 2021, female plants had extremely significantly lower leaf C content and C∶N, C∶P, N∶P ratios (p < 0.001), but extremely significantly higher N and P contents (p < 0.001) than male plants. In 2025, sexual differences were substantially weakened: only C and N contents of females were significantly higher than those of males during flowering (p < 0.05), and no significant sexual differences were detected for other stoichiometric traits (p ≥ 0.05). (2) Ecological homeostasis analysis showed that in 2021, male plants exhibited weak homeostasis for leaf C content during flowering and for the C∶N ratio during both flowering and fruiting periods, indicating some environmental sensitivity. Male plants also showed a steady state for C content and a weak steady state for P content in the fruiting period, while all other indices remained in absolute homeostasis. Female plants maintained a steady state only for the C∶N ratio in the fruiting period and absolute homeostasis for all other indices. In 2025, with improved water and heat conditions, all ecological homeostasis indices for both sexes returned to absolute homeostasis.
      Conclusion Sexual dimorphism in the stoichiometric characteristics and ecological homeostasis of male and female plants of Rhamnus schneideri var. manshurica is fundamentally attributable to differences in reproductive costs. As female plants bear the energy-intensive task of fruit and seed development, they actively accumulate nitrogen and phosphorus during the fruiting stage to enhance leaf photosynthetic efficiency, thereby compensating for reproductive expenditure and maintaining a high level of ecological homeostasis. In contrast, the reproductive investment of male plants is primarily concentrated during the flowering stage, rendering their homeostasis relatively susceptible to environmental influences. This difference in resource allocation strategies, driven by reproductive costs, constitutes a key mechanism underlying sexual dimorphism in dioecious plants.

       

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