间作作物种间相互作用对马铃薯根际土壤丛枝菌根真菌的影响

为揭示间作作物种间相互作用对土壤丛枝菌根(AM)真菌的影响,以马铃薯单作(T0)为对照,基于高通量测序平台的方法,研究了连续3年马铃薯‖玉米(T1)、马铃薯‖蚕豆(T2)下马铃薯根际土壤AM真菌群落组成、多样性与土壤环境因子间的相互关系。结果表明:共获得2893个AM真菌操作分类单元(OTUs),分属1门、3纲、4目、8科、11属、104种;幼套近明囊霉(Claroideoglomus)、球囊霉属(Glomus)、类球囊霉属(Paraglomus)为AM真菌优势属。玉米、蚕豆与马铃薯间作均改变了马铃薯根际土壤AM真菌属水平上的相对丰度;与T0相比,T1处理下幼套近明囊霉相对丰度降低,球囊霉属和类球囊霉属相对丰度增加;T2处理下幼套近明囊霉相对丰度增加,但球囊霉属和类球囊霉属相对丰度却降低。Metastats检验表明:T1和T2处理下,马铃薯根际土壤类球囊霉属AM真菌相对丰度存在显著差异(P=0. 024),耳孢囊霉属(Otospora)在T0和T2处理间存在极显著差异(P=0.005);土壤AM真菌香农指数在T1与T2间有显著差异(P<0.05)。主成分分析表明:T0和T1处理下,马铃薯根际土壤AM真菌群落组成较为接近;土壤碱解氮与AM真菌香农多样性指数呈显著负相关(P=0.043),基于属水平上的AM真菌幼套近明囊霉相对丰度与碱解氮呈极显著正相关(P=0.006),原囊霉属(Archaeospora)的相对丰度与速效磷呈显著正相关(P=0.046)。马铃薯间作玉米、蚕豆为主的种植模式具有间作优势,种间相互作用能改变马铃薯根际土壤AM真菌群落组成,这对于合理间作作物选择,稳定农业生态系统具有重要意义。     

Interspecific interactions affect N and P uptake rather than N:P ratios of plant species: evidence from intercropping

种间相互作用影响植物氮和磷的吸收量而不是氮磷比 量化不同农艺措施下作物氮和磷吸收量(即从农田中移除的量)的化学计量特征对理解农田生态系统中的养分收支和优化氮、磷肥施用至关重要。目前还不清楚在不同的氮肥和覆膜措施下,单作和间作体系作物氮和磷吸收量以及氮磷化学计量特征随整个生长季的变化。本研究探讨了植物种间养分竞争如何对(1) 5种种植模式(小麦、玉米和大麦单作、小麦/玉米和大麦/玉米间作),(2)两种施氮水平(0和225 kg N ha⁻¹)和(3)两种玉米覆膜处理(覆膜和不覆膜)下的作物氮、磷吸收量(以及氮磷比)时间动态的影响。研究结果表明,小麦和大麦的氮、磷竞争能力强于玉米,导致间作体系共生期的小麦和大麦氮、磷吸收量相比于单作增加,而玉米氮、磷吸收量相比于单作减少。3种作物植株氮磷比随作物生长而降低。作物氮磷比不受间作的影响,也不随施氮呈现一致的变化,覆膜降低了玉米的氮磷比。两种间作体系群落水平的氮磷比在成熟期与相应单作不同。由于(1)间作从土壤移除的氮和磷的比例不同于单作,以及(2)作物对氮和磷的吸收在施氮和覆膜下均是不耦合的,这些发现可能对间作系统的养分收支有启示意义。     

不同氮水平下作物养分吸收与利用对玉米马铃薯间作产量优势的贡

玉米与马铃薯间作是重要的间作种植模式,具有较突出的资源利用和产量优势,但养分吸收和利用对作物产量优势的贡献及这种贡献对施氮量的响应机制尚不清楚.本研究采用玉米单作、马铃薯单作和玉米与马铃薯间作3种种植模式,分别设置N_0(0 kg·hm~(-2))、N_1(125 kg·hm~(-2))、N_2(250 kg·hm~(-2))和N_3(375 kg·hm~(-2))4个氮水平,通过2年田间小区试验,研究不同氮水平下间作产量优势的营养基础.结果表明:随着施氮量的增加,氮、磷、钾的单作加权平均吸收量逐渐增加,间作则先增加后减少.间作在N_1水平时具最高的养分吸收优势,分别较单作加权平均值增加氮吸收14.9%、磷吸收38.6%、钾吸收27.8%;间作在N0和N3时具有更高的养分利用效率,较单作可提高氮利用效率3.5%~14.3%、磷利用效率3.5%~18.5%、钾利用效率10.6%~31.6%.N_0和N_1时玉米与马铃薯间作具有显著产量优势,其营养基础在N0时主要是提高了作物养分利用效率,而N_1时则是促进养分吸收的结果.充分发挥间作促进养分吸收对玉米与马铃薯间作产量优势的贡献,需要合理控制氮肥的投入.     

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。     

间作栽培对连作马铃薯根际土壤微生物群落的影响

连作严重影响了作物的产量和品质,而土壤微生物群落结构与功能对土壤生态系统和植物健康至关重要。以连作10a土壤为基质,单作马铃薯为对照,采用磷脂脂肪酸(Phospholipid fatty acids)、BIOLOGA技术和真菌形态学鉴定方法,研究了玉米、蚕豆与马铃薯间作模式下土壤微生物群落结构、功能和丛植菌根(Arbuscular Mycorrhizal)真菌对土壤环境变化的响应。结果表明:间作调控下,马铃薯根际土壤微生物主要类群结构发生显著改变;玉米间作马铃薯,土壤微生物群落总生物量降低,但群落功能多样性提高,促进了以羧酸类、多聚化合物、芳香类化合物、氨基酸类化合物为碳源的微生物类群代谢活性增强;蚕豆间作增加了土壤微生物总生物量,仅促进了以碳水化合物为碳源的微生物类群代谢活性。间作改变了作物根际土壤AM真菌的种、属数,AM真菌多样性降低,优势种由明球囊霉、地球囊霉转变为玉米间作体系里的福摩萨球囊霉、球泡球囊霉;蚕豆间作体系里福摩萨球囊霉和疣状无梗囊霉是优势种。间作栽培下AM真菌优势种群的变化可能受植物间的共生关系、微生物结构与功能等因素的制约。间作条件下,玉米显著影响了马铃薯根际土壤微生物群落功能多样性,而蚕豆则显著改变了微生物群落结构多样性;玉米、蚕豆对马铃薯根际土壤微生物群落功能与结构变化的影响不同步;间作调控后持续的土壤微生物群落结构与功能观察才有助于解释土壤微生物结构变化引起的功能响应。     

不同氮水平下作物养分吸收与利用

玉米与马铃薯间作是重要的间作种植模式,具有较突出的资源利用和产量优势,但养分吸收和利用对作物产量优势的贡献及这种贡献对施氮量的响应机制尚不清楚.本研究采用玉米单作、马铃薯单作和玉米与马铃薯间作3种种植模式,分别设置N₀(0 kg·hm^-2)、N₁(125 kg·hm^-2)、N₂(250 kg·hm^-2)和N₃(375 kg·hm^-2)4个氮水平,通过2年田间小区试验,研究不同氮水平下间作产量优势的营养基础.结果表明: 随着施氮量的增加,氮、磷、钾的单作加权平均吸收量逐渐增加,间作则先增加后减少.间作在N₁水平时具最高的养分吸收优势,分别较单作加权平均值增加氮吸收14.9%、磷吸收38.6%、钾吸收27.8%;间作在N₀和N₃时具有更高的养分利用效率,较单作可提高氮利用效率3.5%～14.3%、磷利用效率3.5%～18.5%、钾利用效率10.6%～31.6%.N₀和N₁时玉米与马铃薯间作具有显著产量优势,其营养基础在N₀时主要是提高了作物养分利用效率,而N₁时则是促进养分吸收的结果.充分发挥间作促进养分吸收对玉米与马铃薯间作产量优势的贡献,需要合理控制氮肥的投入.     

种间互作和施氮对蚕豆/玉米间作生态系统地上部和地下部生长的影响

为河西走廊绿洲灌区豆科/禾本科间作体系的养分管理提供科学依据,于2007年在武威绿洲农业试验站应用田间原位根系行分隔技术研究了蚕豆/玉米种间互作和施氮对玉米抽雄期的根系空间分布、根系形态和作物地上部生长的影响。研究结果表明:种间互作和施氮均增加了玉米和蚕豆在纵向和横向两个尺度上的根重密度、根长密度、根表面积、根系体积。根长密度和根表面积与两种作物产量和氮素吸收均呈正相关,而与蚕豆的根瘤重呈负相关;抽雄期的土壤含水量与玉米产量和养分吸收呈显著的负相关。玉米根系可以占据蚕豆地下部空间,但蚕豆的根却较少到间作玉米的地下部空间,也就是间作后增加了玉米根系水平尺度的生态位。蚕豆和玉米根系主要分布分别在0-40 cm浅土层和0-60 cm 土层,且间作玉米根系在60-120 cm比单作和分隔的多。因此,种间互作和施氮扩大了两作物根系纵向和横向的空间生态位,改变了作物根系形态,即扩展了两者水分和养分吸收的生态位,增加了作物吸收养分的有效空间,从而提高了间作生态系统的生产力。     

玉米/花生间作对土壤微生物和土壤养分状况的影响

通过大田试验,研究了玉米/花生间作对玉米和花生根区土壤微生物和土壤养分状况的影响.结果表明：与单作相比,间作能显著提高玉米和花生根区的土壤细菌数量;间作花生根区土壤真菌和放线菌数量与单作无显著差异;间作玉米根区土壤真菌和放线菌数量比单作明显提高;间作作物根区微生物群落功能多样性和代谢活性比单作有所改善.玉米/花生间作不同程度提高了整个间作系统根区的土壤碱解氮、速效磷、有机质含量及EC值,其中,间作玉米根区土壤养分的增加更为明显,说明玉米/花生间作可以较明显地改善两种作物根区的微生物和养分状况,土壤微生态环境的改善又会促进作物地上部的生长.     

小麦/蚕豆，玉米/蚕豆和小麦/玉米间作对根际细菌群落结构的影响

利用PCR-DGGE技术研究了小麦/蚕豆、玉米/蚕豆和小麦/玉米间作对作物根际细菌群落结构的影响.结果表明：间作能够提高作物根际细菌群落多样性、改变根际细菌群落结构组成.其中,小麦/蚕豆间作对根际细菌群落结构的影响最为突出,作物花期时小麦/蚕豆间作显著提高和改变两种作物根际细菌多样性和群落结构组成.玉米/蚕豆间作主要表现出对苗期玉米根际细菌多样性的显著提高和群落结构组成的改变.小麦/玉米间作对作物根际细菌群落结构的影响程度较弱.同时,3种间作体系都具有不同程度的产量优势.结果证明了间作体系中地上部植物多样性与地下部微生物多样性存在紧密联系.     

入侵植物南美蟛蜞菊和本土蟛蜞菊生长对土壤养分的响应

植物入侵对全球生态系统造成了严重威胁,土壤养分条件(养分水平和养分异质性)对外来植物的成功入侵有重要影响。为了探讨土壤养分条件对入侵植物与同属本地植物生长的影响,本研究以入侵植物南美蟛蜞菊(Wedelia trilobata)和同属本地种蟛蜞菊(W.chinensis)为试验材料开展了温室控制实验。实验设置了低、中、高及异质性养分条件下的单种和混种两种种植方式,测定了每种植物总生物量,根、茎、叶生物量及分节数、根冠比等指标,并分析了土壤养分对二者生长的影响。结果表明:土壤养分水平增加显著促进了南美蟛蜞菊和本地蟛蜞菊的生长;土壤养分异质性显著增加了南美蟛蜞菊根生物量和根冠比,显著增加了本地蟛蜞菊的总生物量、根生物量和茎生物量;土壤养分水平和养分异质性对南美蟛蜞菊和本地蟛蜞菊的竞争响应均无显著影响;在土壤养分较高或较低时南美蟛蜞菊更容易入侵成功;在异质养分环境中,当南美蟛蜞菊生长处于高养斑块时更容易入侵成功。     

土壤氮素异质性分布和马先蒿寄生对长芒棒头草生长发育及根系分布的影响

无论在农田还是自然生态系统中,土壤养分异质性普遍存在。植物具有感知土壤养分异质性的能力,并通过调节根系生物量分配及空间分布以获取更多资源。了解寄生胁迫在不同养分条件下对寄主生长发育及根系空间分布的影响,对解析寄主应对寄生胁迫和养分胁迫的适应策略,进而指导寄生性杂草防控具有重要的指导意义。该文采用分根试验,通过对寄主分根,并控制根室两侧氮供应水平及寄生胁迫程度,考察了氮胁迫及两种寄主依赖程度不同的马先蒿的寄生对寄主长芒棒头草生长发育及根系空间分布的影响。结果表明:(1)土壤氮水平与马先蒿寄生均可显著影响长芒棒头草生物量及根冠比,并且两者之间存在显著交互作用,其中土壤氮水平为主要影响因子。(2)两种马先蒿对长芒棒头草的危害程度不同。在NPK和2NPK 处理时,三色马先蒿的寄生显著降低长芒棒头草生物量(茎叶:37.1%、51.5%; 根系:35.6%、63.6%); 在NPK处理时,大王马先蒿的寄生显著增加长芒棒头草生物量(茎叶:29.9%,根系:61.2%)。(3)长芒棒头草的根系生长和空间分布受氮营养的异质分布和寄生的影响,具有明显的感知养分空间分布及调节根系生长能力。     

Nests of subterranean fungus-growing termites (Isoptera, Macrotermitinae) as nutrient patches for grasses in savannah ecosystems

Savannah are ecosystems in which mineral nitrogen is considered as a limiting factor for plant productivity. They are heterogeneous and spatially structured in patches, or islands, where mineral nitrogen content is concentrated. Among the soil macrofauna, termites of the Macrotermitinae subfamily are major determinants of soil heterogeneity through the biogenic underground nest structures (fungus‐comb chambers) they produce. To study the role of the heterogeneity created by termites on Pennisetum pedicellatum, an herbaceous grass species was grown in greenhouse. This was carried out using an homogeneous soil poor in mineral nitrogen, and an heterogeneous soil with patch, made of (i) Ancistrotermes cavithorax fungus‐comb chamber wall and (ii) soil with the same mineral nitrogen content as the termite handled soil. Plants exhibited a better growth on patch of termite‐modified soil whereas no significant differences were shown with the supply of mineral nitrogen. The presence of fungus‐comb chamber wall material resulted in an increase of fine root biomass and root/shoot ratio. We conclude that termites, through their building activities, may create nutrient patches available to grasses. Concurrently, our data illustrate that the higher mineral nitrogen content in termite‐built structures is not the only factor responsible for plant growth.     

Forest Invasibility in Communities in Southeastern New York

While biological invasions have been the subject of considerable attention both historically and recently, the factors controlling the susceptibility of communities to plant invasions remain controversial. We surveyed 44 sites in southeastern New York State to examine the relationships between plant community characteristics, soil characteristics, and nonnative plant invasion. Soil nitrogen mineralization and nitrification rates were strongly related to the degree of site invasion (F= 30.2, P < 0.0001 and F= 11.8, P < 0.005, respectively), and leaf C : N ratios were negatively correlated with invasion (R ²= 0.22, P < 0.0001). More surprisingly, there was a strong positive relationship between soil calcium levels and the degree of site invasion (partial r= 0.70, P < 0.01), and there were also positive relationships between invasion and soil magnesium and phosphorus. We found, in addition, a positive factor-ceiling relationship between native species diversity and invasive species diversity. This positive relationship between native and invasive diversity contradicts earlier hypotheses concerning the relationships between species diversity and invasion, but supports some recent findings. Cluster analysis distinguished two broad forest community types at our sites: pine barrens and mixed hardwood communities. Invaders were significantly more abundant in mixed hardwood than in pine barrens communities (Mann–Whitney U = 682.5, P < 0.0001). Even when evaluating the mixed hardwood communities alone, invasion remained significantly positively correlated with soil fertility (calcium, magnesium, and net nitrogen mineralization rates). Soil texture and pH were not useful predictors of the degree to which forests were invaded. Nitrogen and calcium are critical components of plant development, and species better able to take advantage of increased nutrient availability may out-perform others at sites with higher nutrient levels. These results have implications for areas such as the eastern United States, where anthropogenic changes in the availability of nitrogen and calcium are affecting many plant communities.     

Spatial heterogeneity in soil nutrient supply modulates nutrient and biomass responses to multiple global change drivers in model grassland communities

Changes in the atmospheric concentration of carbon dioxide ([CO₂]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO₂], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above‐ and belowground biomass (average increases of 32% and 29% for above‐ and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO₂] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO₂], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Effects of light and nutrient availability on the growth,allocation, carbon/nitrogen balance,phenolic chemistry,and resistance to herbivory of two freshwater macrophytes

Phenotypic responses of Potamogeton amplifolius and Nuphar advena to different light (7% and 35% of surface irradiance) and nutrient environments were assessed with field manipulation experiments. Higher light and nutrient availability enhanced the growth of P. amplifolius by 154% and 255%, respectively. Additionally, biomass was allocated differently depending on the resource: high light availability resulted in a higher root/shoot ratio, whereas high nutrient availability resulted in a lower root/shoot ratio. Low light availability and high nutrient availability increased the nitrogen content of leaf tissue by 53% and 40% respectively, resulting in a 37% and 31% decrease in the C/N ratio. Root nitrogen content was also increased by low light and high nutrient availability, by 50% (P=0.0807) and 77% respectively, resulting in a 20% and 40% decrease in root C/N ratio. Leaf phenolics were significantly increased 72% by high light and 31% by high nutrient availability, but root phenolic concentrations were not altered significantly. None of these changes in tissue constituents resulted in altered palatability to crayfish. N. advena was killed by the same high nutrient treatment that stimulated growth in P. amplifolius, preventing assessment of phenotypic responses to nutrient availability. However, high light availability increased overall growth by 24%, but this was mainly due to increased growth of the rhizome (increased 100%), resulting in a higher root/shoot ratio. High light tended to increase the production of floating leaves (P=0.09) and significantly decreased the production of submersed leaves. High light availability decreased the nitrogen content by 15% and 25% and increased the phenolic concentration by 88% and 255% in floating and submersed leaves, respectively. These differences in leaf traits did not result in detectable differences in damage by herbivores.     

Differences between resource patches modify root herbivore effects on plants

While soil resource heterogeneity and root herbivory can have significant direct influences on plant growth, soil heterogeneity may also have indirect effects by influencing the foraging behavior of root herbivores. We used sand-filled greenhouse pots to assess root herbivore foraging behavior and potential interactions between patch quality, herbivore foraging, and plant biomass production (yield). Individual pots were divided into four quarters: one fertilized, and three unfertilized, two of which were planted with tree seedlings. Two treatments were used to create fertilized quarters: high-organic manure fertilizer, and slow-release mineral fertilizer. Seedlings of red maple (Acer rubrum L.) and Virginia pine (Pinus virginiana L.) were used to create two single-species and one mixed-species treatments. Root-feeding beetle larvae were added to the pots and allowed to forage freely for ∼8 weeks. At harvest, root herbivores in organic-fertilized pots were strongly attracted to fertilized quarters despite their relatively low-root biomass. Herbivore distribution was significantly different in mineral fertilized pots, where larvae were most abundant in planted quarters, which is also where most of the plant roots occurred. Whole pot plant yield was significantly reduced by larvae; this effect was stronger in the mineral fertilized pots than organic fertilized pots. While one of the plant species appeared more sensitive to herbivory, root herbivores had a greater influence on yield in mixed-species pots than in single-species pots. Overall, these results suggest that patch quality influences on herbivore foraging may indirectly alter yield and plant community composition. Responsible Editor: Angela Hodge.     

Combined effects between temporal heterogeneity of water supply,nutrient level,and population density on biomass of four broadly distributed herbaceous species

Temporal heterogeneity of water supply affects grassland community productivity and it can interact with nutrient level and intraspecific competition. To understand community responses, the responses of individual species to water heterogeneity must be evaluated while considering the interactions of this heterogeneity with nutrient levels and population density. We compared responses of four herbaceous species grown in monocultures to various combinations of water heterogeneity, nutrient level, and population density: two grasses (Cynodon dactylon and Lolium perenne), a forb (Artemisia princeps), and a legume (Trifolium repens). Treatment effects on shoot and root biomass were analyzed. In all four species, shoot biomass was larger under homogeneous than under heterogeneous water supply. Shoot responses of L. perenne tended to be greater at high nutrient levels. Although root biomass was also larger under homogeneous water supply, effects of water heterogeneity on root biomass were not significant in the grasses. Trifolium repens showed marked root responses, particularly at high population density. Although greater shoot and root growth under homogeneous water supply appears to be a general trend among herbaceous species, our results suggested differences among species could be found in the degree of response to water heterogeneity and its interactions with nutrient level and intraspecific competition.     

Soil heterogeneity and community composition jointly influence grassland biomass

Question: Does the spatial pattern of nutrient supply modify community biomass responses to changes in both species composition and richness? Location: Duke University Phytotron (Durham, North Carolina, USA). Methods: We conducted a microcosm experiment to evaluate individual plant and whole community responses to species richness, species composition and soil nutrient heterogeneity. The experiment consisted of seven levels of species composition (all possible combinations of Lolium perenne, Poa pratensis and Plantago lanceolata) crossed with three levels of soil nutrient distribution (homogeneous, heterogeneous‐up, and heterogeneous‐down, where up and down indicates the location of a nutrient patch in either the upper or the lower half of the soil column, respectively). Results: Communities containing Plantago and Lolium responded to nutrient heterogeneity by increasing above‐ and below‐ground biomass. Nutrient heterogeneity also increased size inequalities among individuals of these species. Significant species composition X nutrient heterogeneity interactions on community biomass and individual size inequality were observed when nutrient patches were located in the upper 10 cm of the soil columns. However, root proliferation in nutrient patches was equivalent regardless of the vertical placement of the patch. Conclusions: Our results suggest that nutrient heterogeneity may interact with plant species composition to determine community biomass, and that small‐scale vertical differences in the location of nutrient patches affect individual and community responses to this heterogeneity.