A millennial‐scale chronicle of evolutionary responses to cultural eutrophication in Daphnia

For an accurate assessment of the anthropogenic impacts on evolutionary change in natural populations, we need long‐term environmental, genetic and phenotypic data that predate human disturbances. Analysis of c. 1600 years of history chronicled in the sediments of South Center Lake, Minnesota, USA, revealed major environmental changes beginning c. 120 years ago coinciding with the initiation of industrialised agriculture in the catchment area. Population genetic structure, analysed using DNA from dormant eggs of the keystone aquatic herbivore, Daphnia pulicaria, suggested no change for c. 1500 years prior to striking shifts associated with anthropogenic environmental alterations. Furthermore, phenotypic assays on the oldest resurrected metazoan genotypes (potentially as old as c. 700 years) indicate significant shifts in phosphorus utilisation rates compared to younger genotypes. Younger genotypes show steeper reaction norms with high growth under high phosphorus (P), and low growth under low P, while ‘ancient’ genotypes show flat reaction norms, yet higher growth efficiency under low P. Using this resurrection ecology approach, environmental, genetic and phenotypic data spanning pre‐ and post‐industrialised agricultural eras clearly reveal the evolutionary consequences of anthropogenic environmental change.     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Growth responses of littoral mayflies to the phosphorus content of their food

We examined how mayfly growth rates and body stoichiometry respond to changing phosphorus (P) content in food. In two experiments, mayfly nymphs were given high or low quantities of food at different carbon:phosphorus (C:P) ratios and their growth was measured. Low food quantity resulted in negative growth rates in both experiments, regardless of food P content. However, under high food availability, mayfly growth was affected by the type of food eaten, with low C:P ratio food producing more rapid growth. In addition, mayfly growth increased somewhat when P-poor food was artificially enriched with inorganic P although this effect was not statistically significant. Mayfly body P content was inversely related to body size but increased in animals fed artificially P-enriched food. A model was constructed to simulate mass balance constraints on mayfly growth imposed by the relative supply of two elements (C and P) in food. The model shows that mayfly growth should be limited by food P content at moderately low C:P ratios ( c . 120, by mass). Given high C:P ratios (mean c.  270, by mass) in periphyton from oligotrophic boreal lakes, our experimental and theoretical results indicate that stoichiometric constraints are important factors affecting benthic food webs in lakes from the Canadian Shield and perhaps in other systems with similarly high C:P ratios in periphyton.     

Photosynthetic and growth responses of three freshwater algae to phosphorus limitation and daylength

1. Three common species of freshwater phytoplankton, the diatom Nitzschia sp., green alga Sphaerocystis schroeteri and cyanobacterium Phormidium luridum, were grown under contrasting daylengths [18 : 6 h light : dark cycles (LD) versus 6 : 18 h LD] and phosphorus (P) regimes (P‐sufficient versus 1 μm P). The rates of growth and photosynthesis, as well as growth efficiencies and pigment concentrations, were compared among treatments. 2. The growth and photosynthetic parameters of the three species depended on both P status and daylength in a species‐specific way. The responses to P limitation depended on daylength and, conversely, the responses to daylength depended on P status. 3. Growth rates and the maximum rates of photosynthesis (P_max) of all species decreased under P limitation under both light regimes. However, the decrease of P_max because of P limitation was greater under long daylength. The P_max of the green alga S. schroeteri decreased the most (ca. sixfold) under P limitation compared with the other two species. The photosynthesis saturation parameter I_k also decreased under P limitation; the decline was significant in Nitzschia and Sphaerocystis. P‐limitation significantly increased photoinhibition (β) in Nitzschia and Sphaerocystis, but not in Phormidium. The excess photochemical capacity (the ratio of the maximum photosynthesis rate to the photosynthesis rate at the growth irradiance), characterising the ability to utilise fluctuating light, was significantly lower under P limitation. 4. The growth efficiency (growth rate normalised to daylength) declined with increasing daylength in all species. Under short daylength the cyanobacterium Phormidium had the lowest growth efficiency of the three species. 5. The cellular chlorophyll a concentration in both Nitzschia and Sphaerocystis was significantly higher under short daylength, but only under P‐sufficient conditions. In Nitzschia, under short daylength, P‐limitation significantly decreased cellular chlorophyll concentration. In contrast, P‐limitation increased cellular chlorophyll concentration in Sphaerocystis, but under long daylength only. The ratio of chlorophyll a to b in the green alga also declined under short daylength and under P‐limited conditions.     

Early growth responses of mangroves to different rates of nitrogen and phosphorus supply

Experiments were conducted in an outdoor facility to quantify growth responses of six mangrove species to rates of dissolved inorganic nitrogen and phosphorus supply mimicking the range of N and P mineralization rates in natural soils. Growth of all six species on nitrogen was nonlinear. Stem extension rates of Rhizophora apiculata and Xylocarpus granatum were enhanced to the highest rate of N supply (50 mmol m^− 2 d^− 1); Bruguiera gymnorrhiza, Avicennia marina, and Xylocarpus moluccensis stem growth leveled off by 10 mmol m^− 2 d^− 1. Stem growth of Ceriops tagal peaked at 24-26 mmol N m^− 2 d^− 1. Except for A. marina and C. tagal, rates of biomass increase declined at the highest supply rate, indicating NH₄⁺ toxicity. At different rates of P supply, stem extension rates and rates of biomass increase of R. apiculata and C. tagal best-fit Gaussian curves and B. gymnorrhiza stem growth and biomass increase best-fit sigmoidal and Gaussian curves, respectively; X. moluccensis stem and biomass growth increased linearly, but stem and biomass growth rates of A. marina did not vary in relation to P supply. Stem growth of X. granatum was Gaussian but rates of biomass increase best-fit a quadratic equation. Changes in leaf and root N and P content mirrored the growth responses. As rates of N and P mineralization in natural mangrove soils overlap with the lowest rates of N and P supplied in these experiments, the growth responses imply that mangroves are intrinsically nutrient-limited at mineralization rates often encountered in nature. Such species specificity may have significant implications for recruitment success and the establishment of species gradients within mangrove forests.     

Plasticity in metabolic allometry: the role of dietary stoichiometry

Metabolism involves multiple elements. While we know much about the allometry in metabolic response of organisms to energy (carbon, C) availability, little is known about how different-sized organisms respond to the relative availability of elements. I experimentally manipulated availability of phosphorus (P) relative to C, to test whether dietary C : P affects metabolism in four species of Daphnia , spanning an order of magnitude in body mass. Results indicated that the slope of the relationship between individual respiration and body mass was M ^0.83 under a balanced diet (C : P c. 150), and M ^0.67 under an imbalanced diet (C : P c. 800). Increased respiration under dietary imbalance was not due to increased ingestion. The change in the scaling exponent was due to the greater respiratory response of smaller species to altered diets. Diet-induced metabolic plasticity contributes to variation in metabolic allometry, at least at such small scales of body size.     

车桑子幼苗生物量分配与叶性状对氮磷浓度的响应差异

调整叶性状和生物量分配格局是植物适应环境变化的主要途径, 研究车桑子(Dodonaea viscosa)幼苗生物量分配与叶性状对氮磷浓度的响应对认识车桑子在氮磷浓度变化下的适应策略具有重要意义。该研究通过砂培法, 测定不同氮浓度(3、5、15、30 mmol·L^-1)与不同磷浓度(0.25、0.5、1、2 mmol·L^-1)下车桑子幼苗的生长、生物量分配、叶性状的响应特征及其相互关系。结果表明: 高浓度氮(30 mmol·L^-1)促进了车桑子幼苗生长、叶片氮含量和生物量积累, 其余氮添加条件(3、5、15 mmol·L^-1)下车桑子幼苗各性状无显著差异, 但相比高氮水平, 其生物量积累和叶片氮含量显著降低, 根冠比和氮利用效率显著增加。随着磷添加浓度的增加, 车桑子幼苗生物量显著增加, 低磷条件(0.25、0.5 mmol·L^-1)限制了车桑子幼苗生长和生物量积累, 其根冠比和磷利用效率均没有发生显著变化, 但比叶面积和叶/茎生物量比例显著增加, 叶干物质含量显著降低。氮处理下, 叶片氮含量与根冠比显著负相关; 磷处理下, 叶片氮含量与比叶面积显著正相关。同时, 氮处理下, 车桑子幼苗株高、基径、总生物量等生长性状均与根冠比显著负相关, 与叶片氮含量显著正相关, 表明根冠比和叶片氮含量的调整在车桑子适应氮限制中发挥重要作用; 而磷处理下, 株高、基径、总生物量与比叶面积显著负相关, 与叶干物质含量显著正相关, 表明叶片结构性状的调整在车桑子适应低磷环境中具有重要意义。该研究表明, 车桑子幼苗生物量分配和叶性状及性状间的权衡策略对氮、磷的响应具有明显差异性, 在今后的研究中, 应关注氮和磷对植物性状影响的差异性。     

河岸带生态系统植被与土壤对水文变化的响应研究进展

河岸带的植被与土壤是生态系统重要组成部分,对于维持河岸带的生态健康、生态系统服务与可持续性具有至关重要的作用。水文变化是河岸带生态系统的首要干扰因子,系统总结了水文变化对河岸带植被的特征以及植被形态、群落分布、繁殖、生存策略的影响,并阐述了河岸带水文和植被对土壤氮磷迁移转化的影响机制。根系作为土壤与植物地上部分之间物质、能量流动与信号传导的关键纽带,目前对根系的研究还较欠缺,需要加强水文变化对河岸带湿地植物根系形态、结构、功能特征的影响机理研究,以及湿地植物对水文变化的适应机制和耐受阈值方面的探究。在微观方面,应加强水文变化与植被等多因素耦合对土壤氮磷迁移转化过程的机理研究。河流形态和土壤的多样性决定着河岸带水文作用特征的复杂性,今后需注重河岸带个性特征与水文响应的关系研究。河岸带是横向的水陆生态过渡带和河流上下游的纵向生态廊道,亟需综合考虑和模拟流域土壤、植被与水文、人类活动之间的耦合关系,预测未来气候与社会经济情境下的河岸带生态系统演变规律,为河岸带生态系统的生态调节、生物多样性保护与生态恢复等提供理论依据与技术支撑。     

Changes of nitrogen and phosphorus loads to European seas

During the last decades human activity has altered the natural cycle of nitrogen and phosphorus on a global scale, producing significant emissions to waters. In Europe, the amount of nutrients discharged from rivers to coastal waters as well as the effects of mitigation measures in place are known only partially, with no consistent temporal and spatial cover. In this study, we quantify the loads and concentration of nitrogen and phosphorus discharged in the European seas over the period 1985–2005, and we discuss their impact on coastal ecosystems. To support our analysis, a catchment database covering the whole of Europe was developed together with data layers of nutrients diffuse and point sources, and the statistical model green was used to estimate the annual loads of nitrogen and phosphorus discharged in all European seas. The results of this study show that during the last 20 years, Europe has discharged 4.1–4.8 Tg yr^?1 of nitrogen and 0.2–0.3 Tg yr^?1 of phosphorus to its coastal waters. We show that beside the North Sea and part of the Baltic Sea, annual nutrient exports have not changed significantly, in spite of the implementation of measures to reduce nutrient sources, and that the N : P ratio has increased steadily, especially in the North, Mediterranean and Atlantic seas. The response of river basins to changes in inputs was not linear, but influenced by climatic variations and nutrients previously accumulated in soils and aquifers. An analysis of the effects of European environmental policies shows that measures to reduce phosphorus were more successful that those tackling nitrogen and that policies aimed at point sources were more effective or more effectively implemented than those controlling pollution from diffuse sources. The increase of the N : P ratio could fuel eutrophication in N‐limited coastal ecosystems, reducing biodiversity and the ecosystem's resilience to future additional anthropogenic stress, such as climate change.     

Morphological and physiological responses of rice (Oryza sativa) to limited phosphorus supply in aerated and stagnant solution culture

BACKGROUND AND AIMS: Rain-fed lowland rice commonly encounters stresses from fluctuating water regimes and nutrient deficiency. Roots have to acquire both oxygen and nutrients under adverse conditions while also acclimating to changes in soil-water regime. This study assessed responses of rice roots to low phosphorus supply in aerated and stagnant nutrient solution. METHODS: Rice (Oryza sativa 'Amaroo') was grown in aerated solution with high P (200 micro m) for 14 d, then transferred to high or low (1.6 micro m) P supply in aerated or stagnant solution for up to 8 d. KEY RESULTS: After only 1 d in stagnant conditions, root radial oxygen loss (ROL) had decreased by 90 % in subapical zones, whereas near the tip ROL was maintained. After 4 d in stagnant conditions, maximum root length was 11 % less, and after 8 d, shoot growth was 25 % less, compared with plants in aerated solution. The plants in stagnant solution had up to 19 % more adventitious roots, 24 % greater root porosity and 26 % higher root/shoot ratio. Rice in low P supply had fewer tillers in both stagnant and aerated conditions. After 1-2 d in stagnant solution, relative P uptake declined, especially at low P supply. Aerated roots at low P supply maintained relative P uptake for 4 d, after which uptake decreased to the same levels as in stagnant solution. CONCLUSIONS: Roots responded rapidly to oxygen deficiency with decreased ROL in subapical zones within 1-2 d, indicating induction of a barrier to ROL, and these changes in ROL occurred at least 2 d before any changes in root morphology, porosity or anatomy were evident. Relative P uptake also decreased under oxygen deficiency, showing that a sudden decline in root-zone oxygen adversely affects P nutrition of rice.     

Adaptive shoot and root responses collectively enhance growth at optimum temperature and limited phosphorus supply of three herbaceous legume species

Background and Aims

Studies on the effects of sub- and/or supraoptimal temperatures on growth and phosphorus (P) nutrition of perennial herbaceous species at growth-limiting P availability are few, and the impacts of temperature on rhizosphere carboxylate dynamics are not known for any species.

Methods

The effect of three day/night temperature regimes (low, 20/13 °C; medium, 27/20 °C; and high, 32/25 °C) on growth and P nutrition of Cullen cinereum, Kennedia nigricans and Lotus australis was determined.

Key Results

The highest temperature was optimal for growth of C. cinereum, while the lowest temperature was optimal for K. nigricans and L. australis. At optimum temperatures, the relative growth rate (RGR), root length, root length per leaf area, total P content, P productivity and water-use efficiency were higher for all species, and rhizosphere carboxylate content was higher for K. nigricans and L. australis. Cullen cinereum, with a slower RGR, had long (higher root length per leaf area) and thin roots to enhance P uptake by exploring a greater volume of soil at its optimum temperature, while K. nigricans and L. australis, with faster RGRs, had only long roots (higher root length per leaf area) as a morphological adaptation, but had a higher content of carboxylates in their rhizospheres at the optimum temperature. Irrespective of the species, the amount of P taken up by a plant was mainly determined by root length, rather than by P uptake rate per unit root surface area. Phosphorus productivity was correlated with RGR and plant biomass.

Conclusions

All three species exhibited adaptive shoot and root traits to enhance growth at their optimum temperatures at growth-limiting P supply. The species with a slower RGR (i.e. C. cinereum) showed only morphological root adaptations, while K. nigricans and L. australis, with faster RGRs, had both morphological and physiological (i.e. root carboxylate dynamics) root adaptations.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

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.     

Resurrecting complexity: the interplay of plasticity and rapid evolution in the multiple trait response to strong changes in predation pressure in the water flea Daphnia magna

A resurrection ecology reconstruction of 14 morphological, life history and behavioural traits revealed that a natural Daphnia magna population rapidly tracked changes in fish predation by integrating phenotypic plasticity and widespread evolutionary changes both in mean trait values and in trait plasticity. Increased fish predation mainly generated rapid adaptive evolution of plasticity (especially in the presence of maladaptive ancestral plasticity) resulting in an important change in the magnitude and direction of the multivariate reaction norm. Subsequent relaxation of the fish predation pressure resulted in reversed phenotypic plasticity and mainly caused evolution of the trait means towards the ancestral pre‐fish means. Relaxation from fish predation did, however, not result in a complete reversal to the ancestral fishless multivariate phenotype. Our study emphasises that the study population rapidly tracked environmental changes through a mosaic of plasticity, evolution of trait means and evolution of plasticity to generate integrated phenotypic changes in multiple traits.     

Opportunities to advance the synthesis of ecology and evolution

Despite decades of research on the interactions between ecology and evolution, opportunities still remain to further integrate the two disciplines, especially when considering multispecies systems. Here, we discuss two such opportunities. First, the traditional emphasis on the distinction between evolutionary and ecological processes should be further relaxed as it is particularly unhelpful in the study of microbial communities, where the very notion of species is hard to define. Second, key processes of evolutionary theory such as adaptation should be exported to hierarchical levels higher than populations to make sense of biodiversity dynamics. Together, we argue that broadening our perspective of eco-evolutionary dynamics to be more inclusive of all biodiversity, both phylogenetically and hierarchically, will open up fertile new research directions and help us to address one of the major scientific challenges of our time, that is, to understand and predict changes in biodiversity in the face of rapid environmental change.