More is not necessarily better: the impact of limiting and excessive nutrients on herbivore population growth rates

Abstract 1. The body tissues of insect herbivores contain higher concentrations of nitrogen and phosphorus than do their host plants, leading to an elemental mismatch that can limit herbivore growth, fecundity and ultimately influence population dynamics. While low nutrient availability can limit herbivore growth and reproduction, nutrient levels that exceed an organism’s nutritional requirements, i.e. an organisms’ threshold elemental ratio, can also decrease performance. 2. We conducted a laboratory experiment to examine the impacts of nitrogen and phosphorus additions on population growth rates of a phloem‐feeding insect herbivore. 3. Herbivore per capita population growth rates were highest at intermediate foliar nitrogen concentrations, indicating a performance cost on the highest nitrogen foliage. While there was no direct effect of foliar phosphorus concentration on insect performance, there was a strong and unexpected indirect effect. High soil phosphorus availability increased both foliar nitrogen concentrations and aphid tissue nitrogen, resulting in low population growth rates when both soil nitrogen and phosphorus availabilities were high. 4. In this study, experimental increases in foliar nitrogen levels led to a decrease in herbivore performance suggesting that excessive nutrient levels can limit herbivore population growth rates.     

Detrital nutrient content determines growth rate and elemental composition of bromeliad‐dwelling insects

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Allometric and trait‐based patterns in parasite stoichiometry

We measured the elemental content (%C, N and P) and ratios (C:N, C:P, N:P) of a diverse assemblage of parasitic helminths to ask whether taxonomy or traits were related to stoichiometric variation among species. We sampled 27 macroparasite taxa, spanning four phyla, infecting vertebrate and invertebrate hosts from freshwater ecosystems in New Jersey. Macroparasites varied widely in elemental content, exhibiting 4.7‐fold variation in %N, 4.6‐fold variation in %P, and 11.5‐fold variation in N:P. Across all species, parasite %P scaled negatively and C:P scaled positively with body size. Similar relationships between parasite P content and body size occurred at the phylum level and within individual species. The allometric scaling of P across species supports the growth rate hypothesis, which predicts that smaller taxa require more P to support relatively higher growth rates. Life cycle stage was related to %N and C:N, with non‐reproductive parasite stages lower in %N and higher in C:N than actively reproducing parasites. Parasite phylum, functional feeding group, and trophic level did not explain elemental variation among species. Organismal stoichiometry is linked to ecological function, and wide variation in macroparasite stoichiometry likely generates diverse patterns in host–parasite nutrient dynamics and variable relationships between parasitism and nutrient cycling.     

Effects of soil nutrients on the sequestration of plant defence chemicals by the specialist insect herbivore,Danaus plexippus

1. Although anthropogenic nitrogen (N) enrichment has significantly changed the growth, survival and reproduction of herbivorous insects, its effects on the defensive sequestration of secondary chemicals by insect herbivores are less well understood. Previous studies have shown that soil nutrient availability can affect sequestration directly through changing concentrations of plant defence chemicals, or indirectly through altering growth rates of herbivores. There has been less exploration of how nutrient deposition affects the consumption of secondary chemicals and subsequent sequestration efficiency. In the current study, the overall effect of soil N availability on cardenolide sequestration by the monarch caterpillar Danaus plexippus was examined. Specifically, the effects of soil nutrient availability on growth, consumption, excretion and sequestration efficiency of cardenolides by D. plexippus larvae fed on the tropical milkweed Asclepias curassavica were measured. 2. The results showed that soil N and phosphorus (P) fertilisation significantly reduced caterpillar growth rate and the sequestration efficiency of cardenolides by monarch caterpillars feeding on A. curassavica. The lowered sequestration efficiency was accompanied by higher concentrations of cardenolides in frass. Although the total cardenolide contents of caterpillars were lower under high N or P fertilisation levels, caterpillar cardenolide concentrations were constant across fertilisation treatments because of lower growth rates (and therefore lower body mass) under high fertilisation. It is concluded that anthropogenic N deposition may have multiple effects on insect herbivores, including their ability to defend themselves from predators with sequestered plant defences.     

中国主要湿地植被氮和磷生态化学计量学特征

研究湿地植物氮(N)和磷(P)的生态化学计量学特征对揭示植物与生境的耦合关系具有重要意义。通过收集中国52个采样区湿地植物不同器官和全株样本的N和P含量, 对其进行分类和统计分析, 探讨植物器官、生长期、植物类型、湿地类型和气候带对湿地植物N和P生态化学计量学特征的影响。结果表明: 1)湿地植物各器官N、P和N:P的几何平均值均表现为叶片(N, 16.07 mg·g^-1; P, 1.85 mg·g^-1; N:P, 8.67) >地上部分(N, 13.54 mg·g^-1; P, 1.72 mg·g^-1; N:P, 7.96) >茎(N, 7.86 mg·g^-1; P, 1.71 mg·g^-1; N:P, 4.58); 2)叶片N含量随时间变化呈现“三峰”型变化, 峰值分别出现在5月、7月和9月; 茎的N含量随时间变化表现为“双峰”型, 峰值出现在5月和9月; 成熟期之前, 植物叶片的N:P与N趋同波动, N:P主要受N含量控制; 衰老期N:P受P含量控制。3)湿地类型是影响植物叶片N和P生态化学计量特征的关键因素, N和P含量最高值出现在河流, 最低值出现在沼泽湿地, N:P的变化趋势大致与之相反。4)植物叶片N、P和N:P的几何平均值都表现为热带>温带>亚热带, 但总体差异不显著(p > 0.05)。5)中国大部分湿地植物叶片N:P < 14, 表现为N限制。     

Coupling of growth rate and body stoichiometry in Daphnia: a role for maintenance processes?

1. The growth rate hypothesis predicts positive relationships among growth rate (μ), body RNA (%RNA of dry mass) and body P (%P of dry mass) contents. 2. We tested this within‐ and across‐species by growing five species/clones of Daphnia (Daphnia magna, Daphnia pulex, Daphnia galeata and two isolates of Daphnia pulicaria) with different combinations of food quantity and stoichiometric food quality. 3. Within each species, positive correlations among μ, %RNA and %P were seen and across species there was a strong association between%RNA and %P, consistent with the growth rate hypothesis. However, coupling of growth to %RNA and to %P differed for different species. In particular, the %RNA–μ and %P–μ relationships had similar slopes but considerably different y‐intercepts (i.e.%P or %RNA at zero growth), with D. pulicaria and D. galeata having higher intercepts than D. magna and especially D. pulex. As a result of these displacements, the relative rankings of the species on the basis of %P and %RNA did not correspond to their rankings based on μ. 4. These findings suggest that within a narrow clade (e.g. the daphnids), interspecific differences in body P content may reflect not growth rate‐related RNA allocation but instead the amount of RNA required for support of maintenance processes.     

Stoichiometric traits of stickleback: Effects of genetic background,rearing environment,and ontogeny

Phenotypes can both evolve in response to, and affect, ecosystem change, but few examples of diverging ecosystem‐effect traits have been investigated. Bony armor traits of fish are good candidates for this because they evolve rapidly in some freshwater fish populations, and bone is phosphorus rich and likely to affect nutrient recycling in aquatic ecosystems. Here, we explore how ontogeny, rearing environment, and bone allocation among body parts affect the stoichiometric phenotype (i.e., stoichiometric composition of bodies and excretion) of threespine stickleback. We use two populations from distinct freshwater lineages with contrasting lateral plating phenotypes (full vs. low plating) and their hybrids, which are mostly fully plated. We found that ontogeny, rearing environment, and body condition were the most important predictors of organismal stoichiometry. Although elemental composition was similar between both populations and their hybrids, we found significant divergence in phosphorus allocation among body parts and in phosphorus excretion rates. Overall, body armor differences did not explain variation in whole body phosphorus, phosphorus allocation, or phosphorus excretion. Evolutionary divergence between these lineages in both allocation and excretion is likely to have important direct consequences for ecosystems, but may be mediated by evolution of multiple morphological or physiological traits beyond plating phenotype.     

Temperature and nutrient availability interact to mediate growth and body stoichiometry in a detritivorous stream insect

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Quantity–quality trade‐offs revealed using a multiscale test of herbivore resource selection on elemental landscapes

1. Herbivores consider the variation of forage qualities (nutritional content and digestibility) as well as quantities (biomass) when foraging. Such selection patterns may change based on the scale of foraging, particularly in the case of ungulates that forage at many scales.
2. To test selection for quality and quantity in free‐ranging herbivores across scales, however, we must first develop landscape‐wide quantitative estimates of both forage quantity and quality. Stoichiometric distribution models (StDMs) bring opportunity to address this because they predict the elemental measures and stoichiometry of resources at landscape extents.
3. Here, we use StDMs to predict elemental measures of understory white birch quality (% nitrogen) and quantity (g carbon/m²) across two boreal landscapes. We analyzed global positioning system (GPS) collared moose (n = 14) selection for forage quantity and quality at the landscape, home range, and patch extents using both individual and pooled resource selection analyses. We predicted that as the scale of resource selection decreased from the landscape to the patch, selection for white birch quantity would decrease and selection for quality would increase.
4. Counter to our prediction, pooled‐models showed selection for our estimates of quantity and quality to be neutral with low explanatory power and no scalar trends. At the individual‐level, however, we found evidence for quality and quantity trade‐offs, most notably at the home‐range scale where resource selection models explain the largest amount of variation in selection. Furthermore, individuals did not follow the same trade‐off tactic, with some preferring forage quantity over quality and vice versa.
5. Such individual trade‐offs show that moose may be flexible in attaining a limiting nutrient. Our findings suggest that herbivores may respond to forage elemental compositions and quantities, giving tools like StDMs merit toward animal ecology applications. The integration of StDMs and animal movement data represents a promising avenue for progress in the field of zoogeochemistry.

     

不同立地铁杆蒿化学计量特征及其与土壤养分的关系

以陕西吴起杨青川流域铁杆蒿群落为研究对象,结合坡向、坡位及土壤养分变化探究不同立地铁杆蒿群落化学计量特征及其与土壤养分的关系.结果表明:从峁顶、阳坡、半阴坡到阴坡,铁杆蒿地上部分和根有机碳、全氮、全磷含量、碳氮比均逐渐增大;地上部分碳磷比和根氮磷比呈减小趋势;地上部分氮磷比、根碳磷比先减小后增大.从峁顶、上坡位、中坡位到下坡位,地上部分有机碳、全氮、全磷含量以及根有机碳含量先增加后减小;地上部分碳氮比、氮磷比先减小后增大.铁杆蒿群落植物化学计量特征一般与土壤化学计量特征呈正相关,但碳氮比、碳磷比、氮磷比及根全磷与土壤相应指标呈负相关,且植物地上部分与土壤的相关性大于根.综上,不同立地铁杆蒿群落植物在阴坡中坡位生长状态最佳,其化学计量特征与土壤养分状况具有明显的相关关系.坡向和坡位在铁杆蒿群落植物和土壤的化学计量特征中具有重要影响,适宜的铁杆蒿群落将有利于土壤养分的恢复.     

Responses of growth of four desert species to different N addition levels

Aims The increase in atmospheric N deposition has accelerated N cycling of ecosystems, thus altering the structure and function of ecosystems, especially in those limited by N availability. Studies on the response of plant growth to artificial N addition could provide basic data for a better understanding of how the structure of grasslands in northern China responds to increasing N deposition. Methods We investigated the seasonal dynamics of plant growth of four species after 2-year multi-level N addition in a field experiment conducted in a desert steppe of Ningxia in 2011. Plant biomass and the relative growth rate (RGR) of the studied species were measured and their relationships with C:N:P ratios of plants (community and leaf levels) and soils were analyzed. Important findings Results in 2012 showed that 2-year N addition promoted the growth of the four species and the effects were different among growth forms and were species-specific. In general, the plant biomass of the studied species was significantly correlated with leaf N concentration, leaf N:P ratio, community N pool, soil total N content and soil N:P ratio, while only weak relationships were observed between plant biomass and C:N and C:P ratios of plants and soils. In contrast, there was a significant linear relationship between RGR and N:P ratios both of plants and soils.Our results suggest that short-term N addition promoted the accumulation of plant biomass, and the species-specific responses to stimulated N addition can directly affect the structure of the desert steppe ecosystem. Plant N:P ratio and soil N:P ratio could indicate nutrient limitation of plant growth to a certain extent: N addition increased soil N content and N:P ratio, and thus relieved N limitation gradually. Once more N is available to plants, the growth of plants and the accumulation of community N was stimulated in turn.     

四种荒漠草原植物的生长对不同氮添加水平的响应

大气氮(N)沉降增加加速了生态系统N循环, 从而会对生态系统的结构和功能产生巨大的影响, 尤其是一些受N限制的生态系统.研究N添加对荒漠草原植物生长的影响, 可为深入理解N沉降增加对我国北方草原群落结构的影响提供基础数据.该文基于2011年在宁夏荒漠草原设置的N沉降增加的野外模拟试验, 研究了两年N添加下4个常见物种(牛枝子(Lespedeza potaninii),老瓜头(Cynanchum komarovii),针茅(Stipa capillata)和冰草(Agropyron cristatum))不同时期种群生物量和6-8月份相对生长速率的变化特征.并通过分析物种生长与植物(群落和叶片水平)和土壤碳(C),N,磷(P)生态化学计量学特征的关系, 探讨C:N:P化学计量比对植物生长养分限制的指示作用.结果显示N添加促进了4个物种的生长, 但具有明显的种间差异性, 且这种差异也存在于相同生活型的不同物种间.总体而言, 4个物种种群生物量与叶片N浓度,叶片N:P,群落N库,土壤全N含量和土壤N:P存在明显的线性关系, 与植物和土壤C:N和C:P的相关关系相对较弱.几个物种相对生长速率与植物和土壤N:P也呈现一定程度的正相关关系, 但与其他指标相关性较弱.以上结果表明, 短期N沉降增加提高了植物的相对生长速率, 促进了植物生长, 且更有利于针茅和老瓜头的生物量积累, 从而可能会逐渐改变荒漠草原群落结构.植物N:P和土壤N:P对荒漠草原物种生长具有较强的指示作用: 随着土壤N受限性逐渐缓解, 土壤N含量和N:P相继升高, 可供植物摄取的N增多, 因而有利于植物生长和群落N库积累.     

Fire effects on insect herbivores in an oak savanna: the role of light and nutrients

Abstract.  1. Environmental heterogeneity created by prescribed burning provided the context for testing whether the distribution of an oak specialist (the lace bug, Corythuca arcuata ) could be explained by stoichiometric mismatches between herbivore and host plant composition.
2. Field observations showed that lace bug density was seven-fold higher in frequently burned than in unburned units.
3. Lace bug density did not increase with leaf nutrient concentrations, but was instead associated with higher light levels, higher concentrations of leaf carbon (C), lignin and total phenolics, and lower levels of cellulose. In addition, lace bugs reared on high-light leaves had higher levels of survivorship than those fed on low-light leaves.
4. Sampling restricted to full-sun leaves was used to test whether fire-related changes in leaf nitrogen (N) and phosphorus (P) concentrations have a secondary influence on lace bug success. This sampling provided only limited evidence for nutrient limitation, as decreases in leaf N and P were associated with an increase in lace bug mass but a decrease in density.
5. It is concluded that burning probably promotes lace bug population growth by increasing canopy openness, light penetration, and the availability of C-based metabolites, and thus simple stoichoimetric mismatches between herbivores and host plants are not of primary importance in this system.     

土壤氮磷化学计量特征对小麦光合气体交换参数和叶绿素荧光参数的影响

选取‘镇麦168’为实验对象,采用盆栽实验,设置16个不同土壤氮磷梯度,测定不同土壤氮磷化学计量比处理下小麦叶片叶绿素含量、光合气体交换参数和叶绿素荧光参数的响应特征,以揭示不同土壤氮磷化学计量关系对小麦叶片光合特性影响的生理生态机制。结果表明:(1)不同生育期内小麦叶片叶绿素含量均随土壤速效N或速效P含量的增加而增加。(2)随着土壤化学计量比N∶P下降,小麦叶片净光合速率(Pn)在土壤中低氮水平(≤258.4mg·kg~(-1))下持续增加,在高氮水平(308.4mg·kg~(-1))下则呈现先增后降的趋势。(3)在土壤速效N相同水平处理下,土壤化学计量比N∶P的降低显著提高了小麦叶片叶绿素荧光参数最小荧光(F0,7.27%~20.00%)、最大荧光(Fm,5.28%~16.15%)、光化学猝灭系数(qP,6.64%~20.92%)及实际光化学效率(ΦPSⅡ,6.95%~18.82%),显著降低了非光化学猝灭系数(NPQ,7.42%~25.63%,P0.05)。研究认为,在中、高氮磷养分水平下,土壤化学计量比N∶P为2.88时,‘镇麦168’叶片净光合速率和实际光化学效率均达到最高水平,表现出较强的光能利用能力。     

Evidence of a general 2/3-power law of scaling leaf nitrogen to phosphorus among major plant groups and biomes

Scaling relations among plant traits are both cause and consequence of processes at organ-to-ecosystem scales. The relationship between leaf nitrogen and phosphorus is of particular interest, as both elements are essential for plant metabolism; their limited availabilities often constrain plant growth, and general relations between the two have been documented. Herein, we use a comprehensive dataset of more than 9300 observations of approximately 2500 species from 70 countries to examine the scaling of leaf nitrogen to phosphorus within and across taxonomical groups and biomes. Power law exponents derived from log–log scaling relations were near 2/3 for all observations pooled, for angiosperms and gymnosperms globally, and for angiosperms grouped by biomes, major functional groups, orders or families. The uniform 2/3 scaling of leaf nitrogen to leaf phosphorus exists along a parallel continuum of rising nitrogen, phosphorus, specific leaf area, photosynthesis and growth, as predicted by stoichiometric theory which posits that plants with high growth rates require both high allocation of phosphorus-rich RNA and a high metabolic rate to support the energy demands of macromolecular synthesis. The generality of this finding supports the view that this stoichiometric scaling relationship and the mechanisms that underpin it are foundational components of the living world. Additionally, although abundant variance exists within broad constraints, these results also support the idea that surprisingly simple rules regulate leaf form and function in terrestrial ecosystems.     

Effects of light and nutrients on the net accumulation and elemental composition of epilithon in boreal lakes

1. Two experiments in the Experimental Lakes Area (ELA) in north-western Ontario, Canada examined the effects of light and two key elements on the net accumulation and elemental composition of epilithon. In Lake (L) 224, benthic algae were grown under different light intensity and phosphorus supply, while in L302S we provided three levels of two different carbon sources (bicarbonate and glucose) to algae colonizing nutrient-diffusing substrata. After 1 month of accumulation, we sampled biofilms for chlorophyll (chl), carbon (C), phosphorus (P) and algal C.
2. Increased C supply did not significantly affect algal growth (C or chl) or elemental composition (C/P ratios) in L302S. However, P enrichment increased chl and algal C, dramatically reduced the C/P ratio of epilithon, and did not affect total organic C in L224. Phosphorus enrichment also increased the proportion of algal material in the total particulate organic matter and altered the taxonomic composition of algae in L224 biofilms. Shading had no significant effect on the C/P ratio and total organic C in epilithon from the L224 experiment.
3. Our results demonstrate that P supply affects the elemental composition of organic matter that collects on rock substrata. It thus appears that low availability of P relative to C and light drives the formation and retention of high C/P organic matter on rock surfaces in oligotrophic boreal lakes.     

Stoichiometric variation of halophytes in response to changes in soil salinity

• Variation in soil salt may change the stoichiometry of a halophyte by altering plant ecophysiology, and exert different influences on various plant organs, which has potentially important consequences for the nutrition of consumers as well as nutrient cycling in a saline ecosystem.
• Using a greenhouse pot experiment, we investigated the effect of salinity variability on the growth and stoichiometry of different organs of Suaeda glauca and Salicornia europaea – two dominant species of important ecological and economic value in the saline ecosystem.
• Our results showed that appropriate salt stimulated the growth of both species during the vigorous growth period, while high salt suppressed growth. Na significantly increased with increased salt in the culture, whereas concentrations of other measured elements and K:Na ratio for both species significantly decreased at low salt treatments, and became more gradual under higher salt conditions. Furthermore, with the change of salt in culture, variations in leaf (degenerated leaf for S. europaea, considered as young stem) stoichiometry, except N:P ratio, were large and less in stems (old stems for S. europaea) than in roots, reflecting physiological and biochemical reactions in the leaf in response to salt stress, supported by sharp changes in trends.
• These results suggest that appropriate saline conditions can enhance biological C fixation of halophytes; however, increasing salt could affect consumer health and decrease cycling of other nutrients in saline ecosystems.

     

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits – growth, respiration, body size, and elemental content – that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population‐level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature–nutrient interactions influence processes at the whole‐ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature–nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.