环境异质性（environmental heterogeneity）是自然环境中的一种普遍现象，其在磷分布和光照条件上表现得尤为明显。磷异质性对植物生长影响显著，当植物生在在离养分富集斑块较远时，会导致植株生长受限、营养失衡、根系发育异常及生殖能力降低等一系列问题，极大影响植物的健康状况。对于一些阳生植物，光照不均分布可能抑制光合作用、干扰植物的光周期调控，甚至引起光合作用的下降，对植物的生长发育和生理节律造成负面影响。植物为了适应这些异质性，会对应采取多种生存策略。有研究表明，通过引入丛枝菌根真菌（Arbuscular mycorrhizal fungi，AMF）或适当增施肥料，可以有效减轻这些异质性对植物的不良影响。尽管如此，但目前研究往往只关注单一类型的异质性对植物的影响，对多种异质性因素共同作用下的综合影响研究鲜有报道。因此，本研究旨在探讨磷异质性条件下植物性状和根系觅食行为的变化，并对比当不同光照强度加入时，植物性状和根系策略变化情况。主要结果如下：

试验一：磷异质对菌根植物红三叶农艺性状及根策略的影响。

本试验旨在研究在磷异质条件下添加氮和AMF对红三叶性状及根系策略的影响。在根盒中使用红三叶进行了盆栽试验。研究不同磷分布（磷均匀分布，Pe；磷近分布，Pc；磷远分布，Pf）、施氮量（N0,0 mg kg^-1；N1，20 mg kg^-1）和AMF（添加，A+；不添加，A–）对植物生长的影响。结果表明，磷分布不均会导致植株地上生物量减少，而同时添加氮和AMF可以显著增加地上生物量。在结构方程模型中显示，AMF与根瘤数和比叶面积呈显著正相关。在本试验缺氮土壤中，氮肥与根瘤数和地上生物量呈显著正相关。根瘤数和比叶面积与地上部生物量呈显著正相关。随后，在随机森林模型中发现，在Pf处理下，细根长度是影响地上生物量的主要因素，其次是最粗根长度。相反，在Pe处理中，最粗的根长是最重要的。

综上所述，在磷分布不均匀的情况下，植物会同时采取了不同的根系觅食策略。AMF在调节根策略、提高根瘤数和比叶面积中起关键作用。氮添加对缓解磷异质对植物生长具有正面影响。

试验二：在不同的光照条件下，磷异质对红三叶农艺性状以及根策略的影响。

基于试验一研究结果，为继续研究多种环境异质相互作用调节菌根植物的根系觅食策略，在磷异质的基础上，加入异质性光环境。本研究通过根盒试验，研究磷异质（磷均质分布，PE；磷远分布，PF）、光照强度（正常光，L1；弱光+正常光，L2；弱光，L3）和AMF及其相互作用下对红三叶表型及根策略的影响。结果表明，光照强度对生物量影响显著，根冠比在L3条件下最大，比叶面积只与光照相关，且光越弱植物的比叶面积越大。随机森林模型中显示，L1处理下粗根和最粗根会显著影响地上部生物量；L3处理下最粗根和细根对地上部生物量贡献显著。在结构方程模型中显示，光照强度和AMF与平均根直径呈显著正相关，根直径与地上部生物量显著正相关。AMF与比根长呈显著负相关。光照强度与比叶面积和比根长呈显著负相关。磷异质性分布与根瘤数和地上部生物量呈显著负相关。

综上所述，红三叶在多重压力下会同时采用不同的根系策略来缓解生存压力，以维持自身的生长。AMF与红三叶形成共生后，可以部分代替植物细根的觅食任务，为植物获取更多养分，缓解植物在异质性环境中的压力。

Environmental heterogeneity is a common phenomenon in natural environments, particularly evident in phosphorus distribution and light conditions. Phosphorus heterogeneity significantly affects plant growth, leading to a series of issues such as restricted growth, nutrient imbalance, abnormal root development, and reduced reproductive capacity when plants grow farther away from nutrient-rich patches. This greatly impacts the health of plants. Uneven distribution of light can inhibit photosynthesis, interfere with plant photoperiod regulation, and even cause a decrease in photosynthetic activity, resulting in negative effects on plant growth, development, and physiological rhythms, especially for some heliophilous plants. To adapt to these heterogeneities, plants adopt a variety of survival strategies.

Studies have shown that introducing arbuscular mycorrhizal fungi (AMF) or appropriately increasing fertilization can effectively mitigate the adverse effects of these heterogeneities on plants. Despite this, current research often only focuses on the impact of a single type of heterogeneity on plants, and there are few reports on the combined effects of multiple heterogeneity factors. Therefore, this study aims to investigate the changes in plant traits and root foraging behavior under P heterogeneity conditions, and to compare the changes in plant traits and root strategies when different light intensity are introduced. The main results are as follows:

Experiment 1:The impact of phosphorus heterogeneity on the agronomic traits and root strategies of the mycorrhizal plant red clover.

This experiment aims to investigate the effects of nitrogen and AMF addition on the traits and root strategies of red clover under P heterogeneity conditions. A rhizobox experiment was conducted using red clover in rhizoboxes to study the impact of different phosphorus distributions (even distribution of P, Pe; close distribution of P, Pc; far distribution of P, Pf), nitrogen (N) application rates (N0, 0 mg kg^-1; N1, 20 mg kg^-1), and AMF (added, A+; not added, A– ) on plant growth. The results indicated that uneven P distribution led to a reduction in aboveground biomass, while the simultaneous addition of N and AMF significantly increased aboveground biomass. The structural equation model showed that AMF was significantly positively correlated with the number of root nodules and specific leaf area (SLA). N fertilizer was significantly positively correlated with the number of root nodules and aboveground biomass. The number of root nodules and SLA were significantly positively correlated with aboveground biomass. Subsequently, the random forest model revealed that, under the Pf treatment, fine root length was the primary factor affecting aboveground biomass, followed by the length of the thickest roots. In contrast, in the Pe treatment, the length of the thickest roots was the most important factor.

  In summary, under conditions of P heterogeneity, plants adopt different root foraging strategies simultaneously. AMF plays a key role in regulating root strategies and increasing the number of root nodules and SLA. N addition has a positive effect on reducing phosphorus heterogeneity and plant growth.

Experiment 2: Under varying light conditions, the influence of phosphorus heterogeneity on the agronomic traits and root strategies of red clover.

Building on the results of experiment 1, this study aims to investigate how multiple environmental heterogeneities interact to regulate the root foraging strategies of mycorrhizal plants. On the basis of P heterogeneity, a heterogeneous light environment is introduced. In natural environments, light conditions are influenced by factors such as topography and vegetation cover. As the energy source for photosynthesis, light directly affects plant growth, development, and morphological construction. Through rhizoboxes experiments, this study examines the effects of P heterogeneity (P even distribution, PE; P far distribution, PF), light intensity (normal light, L1; low light + normal light, L2; low light, L3), and AMF, as well as their interactions, on the phenotype and root strategies of red clover. The effect of light intensity on biomass was significant, with the root-to-shoot ratio being the highest under L3 conditions. The SLA is only related to light conditions, and the weaker the light, the larger the SLA. The random forest model shows that under the L1 treatment, coarse roots and the coarsest roots significantly affect aboveground biomass; under the L3 treatment, the coarsest roots and fine roots contribute significantly to aboveground biomass. The structural equation model shows that the light intensity and AMF are positively correlated with the mean root diameter (ARD), and that ARD was significantly positively correlated with aboveground biomass. AMF showed a significant negative correlation with specific root length (SRL). Light intensity was significantly negatively correlated with SLA and SRL. The distribution of P heterogeneity was significantly negatively correlated with the number of root nodules and aboveground biomass.

In all, under multiple stress conditions, red clover adopts different root strategies simultaneously to alleviate survival pressure and maintain its growth. After forming a symbiotic relationship with AMF, the plant can partially replace the foraging function of fine roots, obtaining more nutrients for the plant and alleviating the stress of plants in heterogeneous environments.

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