Contrasting Predictors of Fern versus Angiosperm Decomposition in a Common Garden

We evaluated differences in the rates and correlates of decomposition among 32 fern and angiosperm litter types collected in Hawai'i. Leptosporangiate ferns were separated into groups based on phylogeny: 'polypod' ferns, a monophyletic clade of ferns that diversified in the Cretaceous, and all other ('non-polypod') ferns that diversified earlier. We measured initial litter chemistry (nutrients and carbon chemistry), and mass loss and nitrogen (N), phosphorus (P), and calcium (Ca) of litter tissue during a 1-yr incubation in a common garden. Nutrient concentrations and carbon (C) chemistry differed significantly among litter types, and litter turnover ( k -values) ranged from 0.29 to 8.31. Decomposition rates were more closely correlated with nutrient concentration than is typically observed. Lignin:N was the best predictor of decomposition across all litter types combined; however, among plant groups different predictors of decomposition were important. Nitrogen and P concentrations best predicted fern decomposition, whereas C chemistry, particularly lignin concentration, was more important for angiosperm (monocot and dicot) decomposition. Among native plants, non-polypod ferns decomposed significantly more slowly than both polypod ferns and angiosperms. Contrary to our hypothesis, fern litter did not decompose more slowly than angiosperm litter overall. Nutrient dynamics in litter were affected by initial litter concentration more than phylogeny; low-nutrient litter immobilized more nutrients than high-nutrient litter. Systematic differences in rates of decomposition, and the importance of nutrients in predicting fern decomposition, imply that changes in species composition within ferns and between ferns and angiosperms could influence the functioning of ecosystems where ferns are important forest components.     

Evaluation of angiosperm and fern contributions to soil organic matter using two methods of pyrolysis-gas chromatography-mass spectrometry

Background

Ferns are an important plant group, and older phylogenies of non-polypod ferns contain relatively high concentrations of aliphatic leaf waxes, lignins, and tannins that could contribute to soil organic matter (SOM) biochemistry and stability.

Methods

Pyrolysis gas-chromatography mass-spectrometry (py-GC/MS) analyzes biochemical fragments which can be related to lignin, polysaccharide, lipid, nitrogen (N)-bearing, non-lignin aromatics, and phenol source compounds. Thermochemolysis using tetramethylammonium hydroxide (TMAH) combined with py-GC/MS improves detection of lignin, cutin, and suberin-derived compounds. To examine the advantages and disadvantages of both methods for characterizing plant and soil biochemistry, we characterized non-polypod and polypod fern and angiosperm live tissues, roots and soils from the Kohala Mountains, Hawaii.

Results

Py-GC/MS provided a broad biochemical overview of compound groups including lignin, polysaccharide, lipid, N-bearing, non-lignin aromatics and phenol groups while TMAH-py-GC/MS detailed lignin units and fatty acids at the expense of the other categories. TMAH-py-GC/MS provided more detailed data on lignin, cutin, suberin and tannin-derived compounds. Both methods detected differences in lignin units between species, although p-coumaric and ferulic acids, predominantly found in ferns, were only observed with TMAH-py-GC/MS.

Conclusions

Both py-GC/MS and TMAH-py-GC/MS are methods to detect compound-specific plant biomarkers, but are most useful when combined for their complimentary results.     

Physiological mechanisms drive differing foliar calcium content in ferns and angiosperms

Recent evidence points to ferns containing significantly lower contents of foliar calcium and other cations than angiosperms. This is especially true of more ancient ‘non-polypod’ fern lineages, which predate the diversification of angiosperms. Calcium is an important plant nutrient, the lack of which can potentially slow plant growth and litter decomposition, and alter soil invertebrate communities. The physiological mechanisms limiting foliar calcium (Ca) content in ferns are unknown. While there is a lot we do not know about Ca uptake and transport in plants, three physiological processes are likely to be important. We measured transpiration rate, cation exchange capacity, and leaching loss to determine which process most strongly regulates foliar Ca content in a range of fern and co-occurring understory angiosperm species from a montane Hawaiian rainforest. We found higher instantaneous and lifetime (corrected for leaf lifespan) transpiration rates in angiosperms relative to ferns. Ferns preferentially incorporated Ca into leaves relative to strontium, which suggests that root or stem cation exchange capacity differs between ferns and angiosperms, potentially affecting calcium transport in plants. There were no differences in foliar Ca leaching loss between groups. Among the physiological mechanisms measured, foliar Ca was most strongly correlated with leaf-level transpiration rate and leaf lifespan. This suggests that inter-specific differences in a leaf’s lifetime transpiration may play a significant role in determining plant nutrition.     

Ecological stoichiometry of the epiphyte community in a subtropical forest canopy

Epiphytes in tree canopies make a considerable contribution to the species diversity, aboveground biomass, and nutrient pools in forest ecosystems. However, the nutrient status of epiphytes and their possible adaptations to nutrient deficiencies in the forest canopy remain unclear. Therefore, we analyzed the stoichiometry of five macroelements (C, N, P, K, and Ca) in four taxonomic groups (lichens, bryophytes, ferns, and spermatophytes) to investigate this issue in a subtropical montane moist evergreen broad‐leaved forest in Southwest China. We found that the interspecific variations in element concentrations and mass ratios were generally greater than the intraspecific variations. And there were significant stoichiometric differences among functional groups. Allometric relationships between N and P across the epiphyte community indicated that P might be in greater demand than N with an increase in nutrients. Although canopy nutrients were deficient, most epiphytes could still maintain high N and P concentrations and low N:P ratios. Moreover, ferns and spermatophytes allocated more limited nutrients to leaves than to stems and roots. To alleviate frequent drought stress in the forest canopy, vascular epiphytes maintained several times higher K concentrations in their leaves than in the tissues of lichens and bryophytes. Our results suggest that epiphytes may have evolved specific nutrient characteristics and adaptations, so that they can distribute in heterogeneous canopy habitats and maintain the stability of nutrient metabolism.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

南昌市不同植物类群叶片氮磷浓度及其化学计量比

对南昌大学前湖校区89种主要植物叶片的N、P浓度及其化学计量比进行了研究,结果表明:乔灌、常绿、针叶、种子、裸子和单子叶植物类群的N浓度分别低于相对应的草本、落叶、阔叶、蕨类、被子和双子叶植物类群,而C3和C4植物差异不显著;乔灌、常绿和裸子植物类群的P浓度含量分别低于相对应的草本、落叶和被子植物类群,而针叶和阔叶、蕨类和种子、单子叶和双子叶、C3和C4植物类群间差异不显著;乔木、阔叶、被子和双子叶植物类群叶片N/P分别高于相对应的灌草、针叶、裸子和单子叶植物类群,而常绿和落叶、蕨类和种子、C3和C4植物类群之间差异不显著.可见,不同类型植物对N和P的吸收利用存在差异,且对不同养分供应采取不同的适应对策.结合研究区土壤养分现状,建议优先选择常绿、针叶、裸子和单子叶植物类群作为城市园林植物.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

Temporal variability of foliar nutrients: responses to nitrogen deposition and prescribed fire in a temperate steppe

Plant nutrient concentrations and stoichiometry drive fundamental ecosystem processes, with important implications for primary production, diversity, and ecosystem sustainability. While a range of evidence exists regarding how plant nutrients vary across spatial scales, our understanding of their temporal variation remains less well understood. Nevertheless, we know nutrients regulate plant function across time, and that important temporal controls could strongly interact with environmental change. Here, we report results from a 3-year assessment of inter-annual changes of foliar nitrogen (N) and phosphorus (P) concentrations and stoichiometry in three dominant grasses in response to N deposition and prescribed fire in a temperate steppe of northern China. Foliar N and P concentrations and their ratios varied greatly among years, with this temporal variation strongly related to inter-annual variation in precipitation. Nitrogen deposition significantly increased foliar N concentrations and N:P ratios in all species, while fire significantly altered foliar N and P concentrations but had no significant impacts on N:P ratios. Generally, N addition enhanced the temporal stability of foliar N and decreased that of foliar P and of N:P ratios. Our results indicate that plant nutrient status and response to environmental change are temporally dynamic and that there are differential effects on the interactions between environmental change drivers and timing for different nutrients. These responses have important implications for consideration of global change effects on plant community structure and function, management strategies, and the modeling of biogeochemical cycles under global change scenarios.     

蕨类植物叶绿体rps4基因的适应性进化分析

在原核生物和植物叶绿体中,RPS4(ribosomal protein small subunit4)在核糖体30S小亚基形成起始过程中发挥重要作用;该蛋白在植物中由叶绿体rps4基因编码。为验证蕨类植物在白垩纪适应被子植物兴起而发生分化的观点,本文以23种蕨类植物为研究对象,利用分支模型、位点模型和分支位点模型对其叶绿体rps4基因进化适应性进行分析。分支模型检测到4个可能存在正选择的分支;位点模型和分支位点模型虽然没有检测出正选择位点,但是位点模型检测出了85个负选择位点。通过研究我们仅仅得出a、b两个代表水龙骨类的分支处于正选择压力下,这与水龙骨类在白垩纪发生辐射式演化的理论相一致。同时rps4基因处于强烈的负选择压力这一事实表明该基因的功能与结构已经趋于稳定。     

Plant invasion is associated with higher plant–soil nutrient concentrations in nutrient‐poor environments

Plant invasion is an emerging driver of global change worldwide. We aimed to disentangle its impacts on plant–soil nutrient concentrations. We conducted a meta‐analysis of 215 peer‐reviewed articles and 1233 observations. Invasive plant species had globally higher N and P concentrations in photosynthetic tissues but not in foliar litter, in comparison with their native competitors. Invasive plants were also associated with higher soil C and N stocks and N, P, and K availabilities. The differences in N and P concentrations in photosynthetic tissues and in soil total C and N, soil N, P, and K availabilities between invasive and native species decreased when the environment was richer in nutrient resources. The results thus suggested higher nutrient resorption efficiencies in invasive than in native species in nutrient‐poor environments. There were differences in soil total N concentrations but not in total P concentrations, indicating that the differences associated to invasive plants were related with biological processes, not with geochemical processes. The results suggest that invasiveness is not only a driver of changes in ecosystem species composition but that it is also associated with significant changes in plant–soil elemental composition and stoichiometry.     

福建三种常绿阔叶林碳氮磷生态化学计量特征

为探明福建三种常绿阔叶林生态系统的养分循环状况及其主要影响因素,采用样地调查采样法,研究了福建武夷山自然保护区甜槠（Castanopsis eyrei）林、天宝岩自然保护区刨花润楠（Machilus pauhoi）林和虎伯寮自然保护区薄叶润楠（Machilus leptophylla）林的植物、凋落物和表层土壤（0-10 cm）碳（C）、氮（N）、磷（P）含量、化学计量特征及其与环境因子的相关性。结果表明：（1）比较3个样点同组分C、N和P含量,除了乔木层的N含量外,虎伯寮植物和凋落物层C、N和P含量均显著高于天宝岩和武夷山;武夷山表层土壤的C和N含量显著高于虎伯寮和天宝岩,P含量排序为天宝岩>武夷山>虎伯寮。（2） P是福建天然林植物生长的主要限制因子,土壤P的缺乏是影响福建常绿阔叶林养分循环的重要因素。（3）植物叶片与凋落物中的养分含量及化学计量比呈极显著正相关,与表层土壤呈极显著负相关,可见福建常绿阔叶林生态系统养分循环过程中,土壤养分消耗速率超过了凋落物的养分供给速率,导致生态系统养分循环缓慢。（4）气温和土壤C ∶ N是影响常绿阔叶林生态系统养分循环的关键环境因子。     

Tissue nutrient concentrations in freshwater aquatic macrophytes: high inter-taxon differences and low phenotypic response to nutrient supply

1. The elemental composition and stoichiometry of aquatic plants has often been suggested to reflect the nutrient enrichment of aquatic habitats. However, the relationship is often weak. Moreover, uncertainties remain in the relevance of laboratory derived critical plant tissue nutrient concentrations to maximum yield or growth rates in the field.
2. Aquatic vascular plants and bryophytes, overlying water and sediment samples were collected to test whether freshwater aquatic macrophytes: (i) show tissue nutrient deficiencies when growing in oligotrophic freshwater habitats, and (ii) have strict homeostatic stoichiometry.
3. Plant nutrient concentrations were significantly related to total inorganic nitrogen (or nitrate), total dissolved phosphorus and sediment total phosphorus. However, these relationships were weak. Virtually all the variance in plant tissue nutrient concentrations, however, could be explained by species (taxon) identity.
4. Critical tissue nutrient concentrations for 95% maximum yield or 95% maximum growth rate in aquatic angiosperms, determined from laboratory bioassays, suggested that nutrients should not limit yield in wild aquatic macrophytes. However, there were a substantial number of samples where potential growth rate limitation was possible, particularly due to phosphorus.
5. Strict C : N : P stoichiometric ratios were found for both vascular plants and bryophytes, suggesting little scope for plants as indicators of nutrient enrichment, but provide robust stoichiometric data for studies on ecosystem metabolism and nutrient cycling.     

The invention of WUS-like stem cell-promoting functions in plants predates leptosporangiate ferns

The growth of land plants depends on stem cell-containing meristems which show major differences in their architecture from basal to higher plant species. In Arabidopsis, the stem cell niches in the shoot and root meristems are promoted by WUSCHEL (WUS) and WOX5, respectively. Both genes are members of a non-ancestral clade of the WUS-related homeobox (WOX) gene family, which is absent in extant bryophytes and lycophytes. Our analyses of five fern species suggest that a single WUS orthologue was present in the last common ancestor (LCA) of leptosporangiate ferns and seed plants. In the extant fern Ceratopteris richardii, the WUS pro-orthologue marks the pluripotent cell fate of immediate descendants of the root apical initial, so-called merophytes, which undergo a series of stereotypic cell divisions and give rise to all cell types of the root except the root cap. The invention of a WUS-like function within the WOX gene family in an ancestor of leptosporangiate ferns and seed plants and its amplification and sub-functionalisation to different stem cell niches might relate to the success of seed plants, especially angiosperms.     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。     

Foliar C,N, and P stoichiometry characterize successful plant ecological strategies in the Sonoran Desert

Ecological processes are centered to water availability in drylands; however, less known nutrient stoichiometry can help explain much of their structure and ecological interactions. Here we look to the foliar stoichiometry of carbon (C), nitrogen (N), and phosphorus (P) of 38 dominant plant species from the Sonoran Desert, grouped in four different functional types to describe ecological characteristics and processes. We found that foliar N, C:N, C:P, and N:P stoichiometric ratios, but not P, were higher than those known to most other ecosystems and indicate P but not N limitations in leaves. Biological N fixers (BNF) had even higher leaf N concentrations, but bio-elemental concentrations and stoichiometry ratios were not different to other non-N-fixing legume species which underscores the need to understand the physiological mechanisms for high N, and to how costly BNF can succeed in P-limiting drylands environments. Stoichiometry ratios, and to lesser extent elemental concentrations, were able to characterize BNF and colonizing strategies in the Sonoran Desert, as well as explain leaf attribute differences, ecological processes, and biogeochemical niches in this dryland ecosystem, even when no direct reference is made to other water-limitation strategies.     

生态化学计量学特征及其应用研究进展

生态化学计量学已成为生态学研究的热点问题。作为一门新兴学科,综观国内外最新研究进展,相关研究目前尚存在着许多不足。基于此,从全球与区域尺度、功能群尺度及个体水平3个方面阐述生态化学计量学特征,从空间、时间、生境和植物类型等生物与非生物因素综述生态化学计量学特征的驱动因素。并讨论生态化学计量学特征在限制性养分判断、生态系统稳定性、生长率与C:N:P关系中的应用。     

干热河谷植物化学计量特征与生物量之间的关系

了解植物化学计量学特征对生物量变化的响应机制对预测全球变化下植物生产力以及生态系统功能具有重要意义。为了了解干热河谷地区植物化学计量学塑性变化与植物生物量变化的关系, 该研究以当地的典型燥红土为基质, 观察水分、养分以及二者的交互作用对6种植物的生长的促进作用, 并分析这种作用与植物化学计量学特征变化的关系。研究结果显示: 水分、养分、物种及其二元交互作用对植物生长具有显著的作用。养分添加处理增加了32.55%的生物量, 高频次水分处理增加了31.35%的生物量, 水分与养分复合处理下生物量增加了110.60%。植物化学计量学特征的变化与植物生物量对处理的响应具有显著相关性。其中, 植物总体K:Ca、K:Mg、K:Mn、K:Zn、Mg:Mn的变化与植物生物量的变化呈正相关关系, 表明水分和养分处理对植物生长的促进作用影响了植物养分的平衡, 主要的变化趋势是高含量元素与低含量元素的计量比随着生物量的增加而不断增加。此外, 相对于植物生物量变化, 处理类型和物种因素对多数化学计量学特征变化无显著影响, 表明水分和养分处理对化学计量学的影响具有相同的驱动机制, 即通过生物量变化最终影响化学计量学变化。植物生物量对水分和养分的响应可对植物化学计量学特征以及生态系统功能产生深远的影响。     

陆地生态系统植被氮磷化学计量研究进展

因化学功能的耦合和元素的不可替代性, 植物对N、P的需求和利用存在严格的比例。植物N、P化学计量在不同功能群、生长地区、生长季、器官之间以及环境梯度下存在明显的变化规律。多数研究从N、P浓度、N:P及N、P间异速指数等角度分析了植物化学计量变化规律, 并探讨其在全球范围内的具体数值。为增进对植物响应全球变化的理解, 该文综述了N、P化学计量的影响因素及其机理的最新研究进展, 并指出未来拟重点研究的方向。     

半干旱黄土小流域不同植被类型植物与土壤生态化学计量特征

生态化学计量学是研究生态系统元素平衡与能量流动的有效方法,明确不同植被恢复类型下植物与土壤化学计量特征对揭示黄土高原植被恢复中生态系统养分循环具重要意义,可为黄土高原植被恢复类型的选择提供可行性依据。以典型半干旱黄土小流域3种植被恢复方式下（天然荒草、自然恢复、人工恢复）的5种植被类型（长芒草草地、赖草草地、苜蓿草地、柠条灌丛、山杏林）为研究对象,分析不同植被类型下叶、茎、根及土壤碳（C）、氮（N）、磷（P）含量及化学计量特征。结果表明：1）植物不同器官和植被类型对植物生态化学计量特征都具有显著影响,C、N、P含量在5种典型植被中均表现为叶>茎>根。人工恢复植被各器官C、N含量及N ∶ P均显著高于天然荒草地,与自然恢复植被无显著差异;其中,在人工恢复植被中山杏各器官C含量最高,柠条各器官N含量最高。叶、茎、根的C ∶ N则表现为自然恢复植被显著高于人工恢复植被与天然荒草地。P含量、C ∶ P则在不同植被恢复类型间无显著差异。2）不同植被恢复类型下土壤C、N、P含量及化学计量特征具一定差异。人工恢复植被土壤C、N、P含量及C ∶ P、N ∶ P均为最高,显著高于自然恢复植被土壤;人工恢复植被中柠条土壤C、N含量及C ∶ P、N ∶ P均显著高于其他植被土壤。土壤C ∶ N在各植被类型间无显著差异。3）不同植被恢复类型下C、N、P含量在植物叶片与土壤间的相关性存在差异,说明植物自身生长特性影响着养分在植物与土壤间转化与传递。以5种典型植被整体来看,植物叶、茎、根的生态化学计量特征在彼此间均呈显著正相关。在植物与土壤间,植物各器官N含量与土壤C、N、P含量呈显著正相关,植物各器官N ∶ P与土壤N ∶ P呈显著正相关,表明该黄土小流域生态系统中植物与土壤生态化学计量特征的变化是相互制约,相互影响的。     

Relationships between plant stoichiometry and biomass in an arid-hot valley,Southwest China

Aims The micro-elemental stoichiometry as well as nitrogen (N) and phosphorus (P) plays an important role in ecosystem process. However, the drivers of the variations in these stoichiometric ratios in plants are less explored in compared with N and P. Plant productivity and plant stoichiometry can response simultaneously to environmental changes, such as water and nutrient supply levels. However, the relationships between the changes in plant stoichiometry and biomass were unclear yet although both of them play important roles in ecosystem functioning. Our object was to investigate the changes in plant stoichiometry (including multiple macro- and micro-elements) and in biomass under different nutrient and water supply. Methods We collected seeds from six grass species in an arid-hot valley and performed a nutrient-water addition experiment in 2012 with a complete factorial design (nutrient × water). The concentrations of N, P, K, Ca, Mg, Zn and Mn in different organs and plant biomass were measured. The effects of species, water and nutrient on element concentration and plant biomass were analyzed by three-way ANOVA. Linear regressions were used to test the relationships between changes in plant stoichiometry and changes in biomass after nutrient and water addition. Important findings Nutrient addition increased plant biomass by 32.55% compared with control. High-level water supply increased plant biomass by 31.35% and the combination of nutrient and high-level water addition increased plant biomass by 110.60%. Nutrient, water, species identity and their two-way interactions significantly affected plant biomass. Changes in total plant K:Ca, K:Mg, K:Mn, K:Zn and Mg:Mn were significantly and positively related to changes in plant biomass. The ratio between the concentrations of macro-elements and micro-elements tended to increase with biomass. Species identity and treatment had no effects in most of these relationships, suggesting that the changes in stoichiometry were mostly driven by the variations in biomass. The relationships between changes in stoichiometry and in biomass also occurred in leaves, stems and roots. The covariation between plant stoichiometry and biomass can have profound effects on ecosystem functioning under the global environmental changes.