植物养分捕获的菌根塑性——外生菌根的塑性

植物根资源捕获塑性是地下生态学研究的重点之一, 在过去二三十年间有长足的进步。菌根塑性是根资源捕获塑性的重要方面, 但由于研究手段的限制, 目前仅有概念上的探讨。缺乏菌根塑性的根塑性研究至少是不全面的。菌根生物学的迅速发展, 尤其是分子生物学手段的介入, 使对菌根塑性进行深入研究成为可能。该文对外生菌根塑性进行讨论, 在简要介绍了外生菌根的生物学基本知识后, 着重讨论了外生菌根形态塑性和生理塑性的定义与内涵。通过文献综述, 分析讨论了外生菌根塑性的研究现状: 很少有研究聚焦在菌根塑性本身, 现有的材料多为其他研究的隐示或研究结果的引申, 并多在形态塑性方面。外生菌根的生理塑性未见有直接的实验数据。该文还对外生菌根研究中发展的、可用于菌根塑性研究的方法进行了综述。由于外生菌根塑性的复杂性, 对菌根塑性的研究会较植物根本身塑性的研究复杂得多, 问题也会相对复杂, 比如植物和外生菌根菌之间的营养需求关系、植物外生菌根塑性的生态意义、实验方法的缺陷等等。对今后外生菌根塑性研究的方向进行了探讨。     

翠菊根系养分捕获形态塑性及其生理机制

为验证以下3个假设: 1) NO₃ ^-和NH₄ ⁺及其不同供给方式显著影响根系生长; 2) NO₃ ^-和NH₄ ⁺以及不同供给方式对根内激素含量影响显著; 3)根构型(1级根长、单位2级根上1级侧根密度(分枝强度)和1级根在2级根上的根间距)与根内激素(生长素(IAA)、脱落酸(ABA)和细胞分裂素(玉米素核苷+玉米素) (CK (ZR + Z))含量显著相关, 采用营养液培养方法, 使实验植物翠菊(Callistephus chinensis)在两种氮肥(NO₃ ^-和NH₄ ⁺)、不同施氮浓度(NO₃ ^-: 0.2、1.0和18.0 mmol·L ^-1; NH₄ ⁺: 0.2、4.0和20.0 mmol·L ^-1), 以及脉冲和稳定两种施用方式处理下生长。在处理35天后收获植物, 测定根系生物量、根系构型指标(根系1级根长、单位2级根上1级侧根数和1级根在2级根上的根间距)和根系中激素含量(IAA、ABA和CK (ZR + Z))。结果显示: 1)实验处理对根生物量和根系中IAA、ABA和CK (ZR + Z)含量均有不同程度的显著影响: 施用NH₄ ⁺使根生物量和根内IAA含量显著低于施用NO₃ ^-; 高浓度NO₃ ^-和NH₄ ⁺处理亦使根生物量和IAA降低; 相对于稳定处理, 脉冲施氮显著降低根生物量和根内IAA含量; NO₃ ^-使根内CK (ZR + Z)含量显著高于施用NH₄ ⁺, 且与施氮浓度及施氮方式无关; NO₃ ^-处理下, 高浓度使根内ABA含量提高, 且脉冲处理使ABA含量升高。NH₄ ⁺处理下, 高浓度使根内ABA含量降低, 而施氮方式对其没有显著影响。2)根构型因素与根内激素关系各异: 各激素与1级根间距无显著关系; IAA和CK (ZR + Z)与1级根长和侧根密度有显著回归关系。3)根构型因素与根生物量的关系是根生物量与1级根长和侧根密度有显著正回归关系, 与1级根间距无显著回归关系。实验结果表明翠菊根生长的 “反常”可能是由于其对脉冲高浓度NH₄ ⁺耐受阈值低所致。该研究通过实验建立了氮养分种类/供应方式通过改变激素、影响根构型而影响根生长的联系, 进一步探究了植物根养分捕获塑性机制。     

水曲柳幼苗根系对土壤养分和水分空间异质性的反应

通过沙培试验方法,研究了温室条件下水曲柳幼苗在施肥和浇水区,非施肥和非浇水区中根系生长,生物量分布,地下部分与地上部分关系,细根直径等特征。结果表明,土壤养分和水分的空间异质性对水曲柳幼苗根系生长和分布有明显影响。在施肥区和浇水区根系生长快,密度大,生物量高,而在非施肥和非浇水区根系生长受到抑制,根系密度小,生物量低,与非施肥区相比,施肥区细根直径下降,有利于根系对养分和水分的运输,但是在非浇水区     

水稻根系在根袋处理条件下对氮养分的反应

通过大田试验对 1 0个水稻品种根系与产量的关系研究表明 ,抽穗期和成熟期根冠比与产量呈极显著的负相关关系 ,相关系数分别为 - 0 .861 6和 - 0 .8889。随之在大田试验基础上选择根冠比大的品种粳籼89,设计水分和养分能自由通过 ,而根系不能穿过的根袋 ,根袋从小到大不同 ,以便产生不同大小的水稻植株根冠比。通过水培实验研究在根袋处理后对不同养分条件的反应。水培液设 3种氮素养分水平 ,即2 0 mg/kg,40 mg/kg,60 mg/kg。结果表明 ,在不同氮素养分条件下 ,经过根袋处理后在抽穗期根系干重都有下降趋势 ,根冠比显著降低 ,而根系活性吸收面积在抽穗期有不同程度的增加 ,茎鞘贮存性碳水化合物含量明显增加 ,叶绿素含量则无明显影响。在抽穗期较大的根袋处理根系总吸收面积、活跃吸收面积及所占比例与对照相比增加效果较为明显 ,而较小的根袋处理根系吸收的能力降低 ,根系吸收能力大小顺序为 :大袋 >中袋 >对照 >小袋。随养分浓度的增加 ,不同根袋处理在抽穗期的根系总吸收面积和活跃吸收面积有下降的趋势。较大的根袋处理在 2 0 mg/kg和 60 mg/kg氮素养分条件下能适当减少根系直径 ,增强根系的活性吸收比例 ,从而提高根系的活力 ;但在成熟期根袋处理对根系的活性吸收无明显影响     

根际微生物调控植物根系构型研究进展

根系构型是最重要的植物形态特征之一,具有可塑性,既由遗传因素控制,又受到许多环境因子的调控。近年的大量研究表明,根际微生物能够调控植物的根系构型,进而影响植物的一系列生理与生态过程。综述丛枝菌根真菌(AMF)、根瘤菌、植物根际促生菌(PGPR)等重要根际微生物类群对植物根系构型的调控模式以及相应的调控机理,并对进一步的研究进行了展望,旨在为今后的相关研究和实际应用提供参考。     

根压与根系的主动吸水

植物根系吸收水分有两种类型:被动吸水和主动吸水,被动吸水的动力是蒸腾拉力,主动吸水的动力是根压。我认为根压不是植物根系吸水的动力,而应当是植物根系吸水的结果。理由有三点: 第一,根压的定义没有“根压是根系主动吸水的动力”的含义。“根系的生理活功使液流上升的压力称为根压”。“根压是由于根系代谢活动的结果,使木质部导管分子内产生     

宁南山区典型植物根系分解特征及其对土壤养分的影响

根系分解是陆地生态系统碳和养分循环的重要地下生态过程,研究宁南山区典型植物根系分解特征及其对土壤养分的影响,能够丰富和完善陆地生态系统的物质和能量循环机制,为我国黄土高原植被恢复过程中植物与土壤之间的养分循环提供依据。连续2年研究了宁南山区3种典型植物(长芒草、铁杆蒿和百里香)根系的分解特征及其对土壤养分的影响。结果表明,长芒草、铁杆蒿和百里香根系年分解指数(K)分别0.00891、0.01128、0.01408,分解速率依次表现为百里香铁杆蒿长芒草。分解16个月后3种典型植物根系释放大量养分,其中碳的释放量在57.05—124.39 g/kg;氮的释放量在0.12—0.47 g/kg。3种典型植物根系对土壤养分的影响主要表现为:试验结束时,0—5 cm表层土壤有机碳含量提高了0.17—0.35 g/kg,5—20 cm土层土壤有机碳含量提高了0.26—0.35 g/kg。相关性分析可知,植物根系养分释放量与土壤养分含量之间存在一定的负相关关系,当土壤养分含量较低时,根系会增加养分释放量进行补充。由此可知,根系分解提高了土壤养分含量,有效的促进了养分在根系-土壤中的循环。     

植物根系向地性感应的分子机理与养分吸收

植物根系向地性是决定根系空间生长趋势的主要因素之一, 对于养分吸收具有重要影响。认识根系向地性感应和根系生长变化的分子机理及其与养分吸收的关系, 可为遗传改良根系性状、提高植物养分吸收效率提供理论依据。本文从重力感应、信号转导和生长素非对称分布等方面总结了植物根系向地性感应的分子机理, 探讨了根系在养分胁迫下(特别是磷胁迫下)向地性变化的生理基础及其与养分吸收(特别是磷吸收)的关系, 最后对根系向地性研究的若干问题进行了展望。     

植物根系向地性感应的分子机理与养分吸收

植物根系向地性是决定根系空间生长趋势的主要因素之一,对于养分吸收具有重要影响.认识根系向地性感应和根系生长变化的分子机理及其与养分吸收的关系,可为遗传改良根系性状、提高植物养分吸收效率提供理论依据.本文从重力感应、信号转导和生长素非对称分布等方面总结了植物根系向地性感应的分子机理,探讨了根系在养分胁迫下(特别是磷胁迫下)向地性变化的生理基础及其与养分吸收(特别是磷吸收)的关系,最后对根系向地性研究的若干问题进行了展望.     

喀斯特地区植物根系分泌物酶活性对根际土酶活性和养分的影响

为了探究喀斯特地区植物根系分泌物酶活性对根际土酶活性和养分的影响,我们测定了灌草丛、灌木林、灌乔林和乔木林4个植被恢复阶段群落水平根系分泌物和根际土的β-1,4-葡萄糖苷酶(βG)、β-N-乙酰葡萄糖苷酶(NAG)、亮氨酸氨基肽酶(LAP)和酸性磷酸酶(ACP)的活性与土壤碳氮磷的含量,并分析了它们之间的关系.结果表明:(1)根际土以及根系分泌物的4种酶活性在植被恢复后期显著高于植被恢复前期;乔木林的根系分泌物酶活性C∶P和N∶P比值显著高于其他植被恢复阶段,而根际土酶活性这2个比值则正好相反.(2)相关分析显示,根系分泌物酶活性与对应的土壤酶活性呈显著正相关;相对于根系分泌物酶活性,土壤酶活性与相关养分的相关系数值更高.另外,根际土以及根系分泌物βG、NAG和LAP酶活性与根际土有机碳(SOC)和全氮(TN)呈显著正相关,根际土以及根系分泌物ACP酶活性与根际土速效磷(AP)呈显著正相关.上述结果表明,植被恢复对根系分泌物酶和土壤酶活性的提高具有正向的作用,根系分泌物酶是土壤酶的有利补充,在碳氮磷养分循环过程中起到积极的促进作用.综上所述,调控根系分泌物分泌及其酶活性可能为喀斯特生态系统的植被恢复提供新的视角.     

外来入侵植物的根系觅养行为研究进展

土壤养分分布具有高度空间异质性,植物的根系觅养行为是其对土壤养分异质性的一种适应。不同植物为了适应养分异质性会产生不同的根系觅养行为,通过调整自身的根系觅养范围、觅养精度和觅养速度来更好地吸收利用土壤中的养分。外来植物与本地植物的竞争是决定其成功入侵的重要因素,土壤养分等环境因素会影响它们之间的竞争关系。近年来,外来入侵植物的觅养行为逐渐受到人们的关注,关于入侵植物根系觅养行为的研究成果陆续出现:(1)总体来看,外来入侵植物具有较强的根系觅养能力,但根系觅养范围与觅养精度之间的权衡关系还不确定;(2)营养异质性会影响入侵植物与本地植物之间的竞争,反过来,二者之间的竞争也会影响根系觅养行为对营养异质性的响应;(3)丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)能够提高入侵植物的根系觅养能力,外来植物入侵能够改变入侵植物对AMF的偏好性,形成AMF对入侵的正反馈作用,而本地植物与AMF的相互作用也会影响入侵植物的竞争力。未来还应加强营养异质环境下种间竞争和AMF共生对入侵植物根系觅养行为的影响机制研究,以及全球变化背景下入侵植物根系觅养行为的变化与机制方面的研究,可以更深入地认识外来植物的觅养行为在其成功入侵中的作用,并为利用营养调控来防控入侵植物提供理论依据。     

Intra-plant versus Inter-plant Root Competition in Beans: avoidance,resource matching or tragedy of the commons

Root competition inhibits root proliferation. All else equal, a plant should invest roots in a nutrient patch devoid of roots rather than one already occupied by roots. Less clear is how a plant should respond to intra-plant versus inter-plant root competition. We consider three responses for how a plant may select habitats based on intra-versus inter-plant root competition: inter-plant avoidance, resource matching, or intra-plant avoidance. The first assumes that plants prefer to have their own space and preferentially proliferate roots away from neighboring plants. The second response, based on the ideal free distribution, assumes that plants invest so as to equalize average returns from roots, regardless of the identity of the neighboring roots. The third, based on game theory, assumes that the plant proliferates roots so as to maximize whole-plant fitness, in which case it is better to proliferate plants among a neighbor's roots than to continue proliferating amongst one's own roots. To test among these models we grew beans (Phaseolus varigaris, var. Kenya) in a greenhouse under two planting scenarios. Both scenario were tested under 0.5 and 0.1 strength of nutrient solution. Under scenario A (fence-sitters), two split-root plants each shared two patches by virtue of having roots in each. Under scenario B (owners) two plants each had their own patch. The results supported the game theory model of intra-plant avoidance (whole plant habitat selection). Fence-sitters produced 150% more root mass per individual than owners. Owners produced 90% more yield (dry mass of pods) than fence-sitters. Furthermore, owners had significantly higher shoot-root ratios than fence-sitters. These effects did not vary with high or low nutrient levels. The over-proliferation of roots under inter-plant competition (fence-sitters) was manifest by the tenth day after planting. In short, the fence-sitters engaged in a tragedy of the commons in which they competed with each other through root proliferation. At the ESS, the fitness maximizing strategy of the individual is to sacrifice collective yield in a quest to `steal' nutrients from its neighbor. The research has three implications. First, plants may be able to assess and respond to local opportunities in a manner that maximizes the good of the whole plant. Second, nutrient foraging as a game may provide a fresh perceptive for viewing root competition either intra-specifically or inter-specifically. Third, it may be possible to increase the yield of certain crop species by breeding more `docile' cultivars that do not overproduce roots in response to inter-plant competition.     

Nutrient foraging traits in 10 co-occurring plant species of contrasting life forms

1 Responses to spatial heterogeneity of soil nutrients were tested in 10 plant species that differ in life form and successional status, but which co-occur in the South Carolina coastal plain. The morphological responses of the root system were tested by assessing scale (represented by root mass and root length densities), precision (preferential proliferation of roots in nutrient-rich patches compared with less fertile patches) and discrimination (ability to detect and proliferate within the richest patches when patches vary in nutrient concentration). We also investigated sensitivity (growth benefits gained as spatial heterogeneity of nutrients increases, measured as total biomass).
2 Ten individuals of each species were grown in pots under four treatments that had differing nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge.
3 We observed high variation between species in scale, precision and sensitivity. No significant discrimination responses were observed, although greatest root mass density occurred at intermediate fertility levels for all species.
4 We rejected the hypothesis that scale and precision are negatively correlated. Indeed, in herbaceous species alone, scale and precision were positively correlated.
5 Sensitivity was not closely related to precision, indicating that proliferation of roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision, suggesting that a positive growth response in heterogeneous environments may be related to plasticity in physiology or root life span, rather than morphology.
6 Plant life form was not correlated with precision or sensitivity. However, scale of response was greater in herbs than in woody plants, possibly because the two life forms develop root systems at different rates.     

Influence of root herbivory on plant communities in heterogeneous nutrient environments

While plant species respond differently to nutrient patches, the forces that drive this variability have not been extensively examined. In particular, the role of herbivory in modifying plant-resource interactions has been largely overlooked. We conducted a glasshouse study in which nutrient heterogeneity and root herbivory were manipulated, and used differences in foraging among plant species to predict the influence of root herbivores on these species in competition. We also tracked the influence of neighborhood composition, heterogeneity, and herbivory on whole-pot plant biomass. When herbivores were added to mixed-species neighborhoods, Eupatorium compositifolium, the most precise forager, was the only plant species to display a reduction in shoot biomass. Neighborhood composition had the greatest influence on whole-pot biomass, followed by nutrient heterogeneity; root herbivory had the smallest influence. These results suggest that root herbivory is a potential cost of morphological foraging in roots. Root herbivores reduced standing biomass and influenced the relative growth of species in mixed communities, but their effect was not strong enough at the density examined to overwhelm the bottom-up effects of resource distribution.