Impact of Temperature and Nutrients on Carbon: Nutrient Tissue Stoichiometry of Submerged Aquatic Plants: An Experiment and Meta-Analysis.
ABSTRACT: Human activity is currently changing our environment rapidly, with predicted temperature increases of 1-5°C over the coming century and increased nitrogen and phosphorus inputs in aquatic ecosystems. In the shallow parts of these ecosystems, submerged aquatic plants enhance water clarity by resource competition with phytoplankton, provide habitat, and serve as a food source for other organisms. The carbon:nutrient stoichiometry of submerged aquatic plants can be affected by changes in both temperature and nutrient availability. We hypothesized that elevated temperature leads to higher carbon:nutrient ratios through enhanced nutrient-use efficiency, while nutrient addition leads to lower carbon:nutrient ratios by the luxurious uptake of nutrients. We addressed these hypotheses with an experimental and a meta-analytical approach. We performed a full-factorial microcosm experiment with the freshwater plant Elodea nuttallii grown at 10, 15, 20, and 25°C on sediment consisting of pond soil/sand mixtures with 100, 50, 25, and 12.5% pond soil. To address the effect of climatic warming and nutrient addition on the carbon:nutrient stoichiometry of submerged freshwater and marine plants we performed a meta-analysis on experimental studies that elevated temperature and/or added nutrients (nitrogen and phosphorus). In the microcosm experiment, C:N ratios of Elodea nuttallii decreased with increasing temperature, and this effect was most pronounced at intermediate nutrient availability. Furthermore, higher nutrient availability led to decreased aboveground C:P ratios. In the meta-analysis, nutrient addition led to a 25, 22, and 16% reduction in aboveground C:N and C:P ratios and belowground C:N ratios, accompanied with increased N content. No consistent effect of elevated temperature on plant stoichiometry could be observed, as very few studies were found on this topic and contrasting results were reported. We conclude that while nutrient addition consistently leads to decreased carbon:nutrient ratios, elevated temperature does not change submerged aquatic plant carbon:nutrient stoichiometry in a consistent manner. This effect is rather dependent on nutrient availability and may be species-specific. As changes in the carbon:nutrient stoichiometry of submerged aquatic plants can impact the transfer of energy to higher trophic levels, these results suggest that eutrophication may enhance plant consumption and decomposition, which could in turn have consequences for carbon sequestration.
Project description:The abundance and stoichiometry of aquatic plants are crucial for nutrient cycling and energy transfer in aquatic ecosystems. However, the interactive effects of multiple global environmental changes, including temperature rise and eutrophication, on aquatic plant stoichiometry and palatability remain largely unknown. Here, we hypothesized that (1) plant growth rates increase faster with rising temperature in nutrient-rich than nutrient-poor sediments; (2) plant carbon (C): nutrient ratios [nitrogen (N) and phosphorus (P)] respond differently to rising temperatures at contrasting nutrient conditions of the sediment; (3) external nutrient loading to the water column limits the growth of plants and decreases plant C:nutrient ratios; and that (4) changes in plant stoichiometry affect plant palatability. We used the common rooted submerged plant Vallisneria spiralis as a model species to test the effects of temperature and nutrient availability in both the sediment and the water column on plant growth and stoichiometry in a full-factorial experiment. The results confirmed that plants grew faster in nutrient-rich than nutrient-poor sediments with rising temperature, whereas external nutrient loading decreased the growth of plants due to competition by algae. The plant C: N and C: P ratios responded differently at different nutrient conditions to rising temperature. Rising temperature increased the metabolic rates of organisms, increased the nutrient availability in the sediment and enhanced plant growth. Plant growth was limited by a shortage of N in the nutrient-poor sediment and in the treatment with external nutrient loading to the water column, as a consequence, the limited plant growth caused an accumulation of P in the plants. Therefore, the effects of temperature on aquatic plant C:nutrient ratios did not only depend on the availability of the specific nutrients in the environment, but also on plant growth, which could result in either increased, unaltered or decreased plant C:nutrient ratios in response to temperature rise. Plant feeding trial assays with the generalist consumer Lymnaea stagnalis (Gastropoda) did not show effects of temperature or nutrient treatments on plant consumption rates. Overall, our results implicate that warming and eutrophication might interactively affect plant abundance and plant stoichiometry, and therefore influence nutrient cycling in aquatic ecosystems.
Project description:Global warming is expected to strengthen herbivore-plant interactions leading to enhanced top-down control of plants. However, latitudinal gradients in plant quality as food for herbivores suggest lower palatability at higher temperatures, but the underlying mechanisms are still unclear. If plant palatability would decline with temperature rise, then this may question the expectation that warming leads to enhanced top-down control. Therefore, experiments that directly test plant palatability and the traits underlying palatability along a temperature gradient are needed. Here we experimentally tested the impact of temperature on aquatic plant growth, plant chemical traits (including stoichiometry) and plant palatability. We cultured three aquatic plant species at three temperatures (15, 20, and 25°C), measured growth parameters, determined chemical traits and performed feeding trial assays using the generalist consumer Lymnaea stagnalis (pond snail). We found that rising temperature significantly increased the growth of all three aquatic plants. Plant nitrogen (N) and phosphorus (P) content significantly decreased, and carbon (C):N and C:P stoichiometry increased as temperature increased, for both Potamogeton lucens and Vallisneria spiralis, but not for Elodea nuttallii. By performing the palatability test, we found that rising temperatures significantly decreased plant palatability in P. lucens, which could be explained by changes in the underlying chemical plant traits. In contrast, the palatability of E. nuttallii and V. spiralis was not affected by temperature. Overall, P. lucens and V. spiralis were always more palatable than E. nuttallii. We conclude that warming generally stimulates aquatic plant growth, whereas the effects on chemical plant traits and plant palatability are species-specific. These results suggest that the outcome of the impact of temperature rise on macrophyte stoichiometry and palatability from single-species studies may not be broadly applicable. In contrast, the plant species tested consistently differed in palatability, regardless of temperature, suggesting that palatability may be more strongly linked to species identity than to intraspecific variation in plant stoichiometry.
Project description:Two submerged Elodea species have small differences in their ecophysiological responses when exposed to individual environmental factors. However, field observations showed that under eutrophic conditions with low light availability, Elodea canadensis could be displaced by Elodea nuttallii. Here we investigated the combined effect of environmental factors on the ecophysiological response of the two species in order to explain the differences in their invasion successes. We cultivated the plants in aquaria containing five different nitrogen (N) concentrations and incubated at five different light intensities. For both species increasing nitrogen concentrations resulted in increased relative growth rate, chlorophyll concentration, and actual photochemical efficiency of photosystem II (?PSII), however, they produced less roots. Lowering light intensity resulted in a lower relative growth rate, root production, and nutrient removal. In contrast, chlorophyll concentration in the leaves, and ?PSII increased. The main difference between the two Elodea species was that the light compensation point (I c) and weight loss in the dark were significantly higher and photochemical efficiency and chlorophyll concentration were significantly lower for E. canadensis than for E. nuttallii, indicating that the latter can survive under much more shady and hypertrophic conditions. The change in nitrogen concentration of the media and in tissue concentration of the plants indicated that E. nuttallii has a higher nitrogen removal capacity. The ecophysiological differences between the two species can be an explanation for invasion success of E. nuttallii over E. canadensis and thus may explain why the latter is replaced by the first.
Project description:Vegetative propagule pressure may affect the establishment and structure of aquatic plant communities that are commonly dominated by plants capable of clonal growth. We experimentally constructed aquatic communities consisting of four submerged macrophytes (Hydrilla verticillata, Ceratophyllum demersum, Elodea nuttallii and Myriophyllum spicatum) with three levels of vegetative propagule pressure (4, 8 and 16 shoot fragments for communities in each pot) and two levels of water depth (30 cm and 70 cm). Increasing vegetative propagule pressure and decreasing water level significantly increased the growth of the submerged macrophyte communities, suggesting that propagule pressure and water depth should be considered when utilizing vegetative propagules to re-establish submerged macrophyte communities in degraded aquatic ecosystems. However, increasing vegetative propagule pressure and decreasing water level significantly decreased evenness of the submerged macrophyte communities because they markedly increased the dominance of H. verticillata and E. nuttallii, but had little impact on that of C. demersum and M. spicatum. Thus, effects of vegetative propagule pressure and water depth are species-specific and increasing vegetative propagule pressure under lower water level can facilitate the establishment success of submerged macrophyte communities.
Project description:Non-indigenous species (NIS) are species living outside their historic or native range. Invasive NIS often cause severe environmental impacts, and may have large economical and social consequences. Elodea (Hydrocharitaceae) is a New World genus with at least five submerged aquatic angiosperm species living in fresh water environments. Our aim was to survey the geographical distribution of cpDNA haplotypes within the native and introduced ranges of invasive aquatic weeds Elodea canadensis and E. nuttallii and to reconstruct the spreading histories of these invasive species. In order to reveal informative chloroplast (cp) genome regions for phylogeographic analyses, we compared the plastid sequences of native and introduced individuals of E. canadensis. In total, we found 235 variable sites (186 SNPs, 47 indels and two inversions) between the two plastid sequences consisting of 112,193 bp and developed primers flanking the most variable genomic areas. These 29 primer pairs were used to compare the level and pattern of intraspecific variation within E. canadensis to interspecific variation between E. canadensis and E. nuttallii. Nine potentially informative primer pairs were used to analyze the phylogeographic structure of both Elodea species, based on 70 E. canadensis and 25 E. nuttallii individuals covering native and introduced distributions. On the whole, the level of variation between the two Elodea species was 53% higher than that within E. canadensis. In our phylogeographic analysis, only a single haplotype was found in the introduced range in both species. These haplotypes H1 (E. canadensis) and A (E. nuttallii) were also widespread in the native range, covering the majority of native populations analyzed. Therefore, we were not able to identify either the geographic origin of the introduced populations or test the hypothesis of single versus multiple introductions. The divergence between E. canadensis haplotypes was surprisingly high, and future research may clarify mechanisms that structure native E. canadensis populations.
Project description:Monocultures of functional equivalent species often negatively affect nutrient cycling and overall biodiversity of aquatic ecosystems. The importance of water and sediment nutrients for the occurrence of monocultures was analysed using field data from drainage ditches. Ranges of nutrients were identified that best explained the occurrence of monocultures of Elodea nuttallii (Planch.) St. John (Waterweed type), monocultures of duckweed (Duckweed type) and the occurrence of a diverse submerged vegetation (Mixed type). Results indicated these three vegetation types occurred at distinctive ranges of phosphorus in water and sediment. Sediment phosphorus distinguished monocultures from the Mixed type, with the two monocultures occurring at two to four times higher concentrations. The Waterweed type occurred at higher sediment phosphorus levels than the mixed type, showed a higher degree of dominance and lower number of red list species. Phosphorus concentrations in water were four to six times higher in the Duckweed type compared to the Waterweed and Mixed type. The three vegetation types had comparable total biomass which was unexpected. This comparability was likely caused by duckweed only growing at the water surface at the highest nutrient levels and the limited space in drainage ditches for increased submerged biomass development at high nutrient availability. Possible measures to limit the occurrence of monocultures, and thereby increasing the ecological quality, are discussed with focus on lowering phosphorus concentrations in both water and sediment and on removal of plant species that develop into monocultures.
Project description:The abundance of primary producers is controlled by bottom-up and top-down forces. Despite the fact that there is consensus that the abundance of freshwater macrophytes is strongly influenced by the availability of resources for plant growth, the importance of top-down control by vertebrate consumers is debated, because field studies yield contrasting results. We hypothesized that these bottom-up and top-down forces may interact, and that consumer impact on macrophyte abundance depends on the nutrient status of the water body. To test this hypothesis, experimental ponds with submerged vegetation containing a mixture of species were subjected to a fertilization treatment and we introduced consumers (mallard ducks, for 8 days) on half of the ponds in a full factorial design. Over the whole 66-day experiment fertilized ponds became dominated by Elodea nuttallii and ponds without extra nutrients by Chara globularis. Nutrient addition significantly increased plant N and P concentrations. There was a strong interactive effect of duck presence and pond nutrient status: macrophyte biomass was reduced (by 50%) after the presence of the ducks on fertilized ponds, but not in the unfertilized ponds. We conclude that nutrient availability interacts with top-down control of submerged vegetation. This may be explained by higher plant palatability at higher nutrient levels, either by a higher plant nutrient concentration or by a shift towards dominance of more palatable plant species, resulting in higher consumer pressure. Including nutrient availability may offer a framework to explain part of the contrasting field observations of consumer control of macrophyte abundance.
Project description:Elodea nuttallii is widely used in Chinese mitten crab (CMC) rearing practice, but it is not a native aquatic plant and cannot endure high temperature. Thus, large E. nuttallii mortality and water deterioration events could occur during high-temperature seasons. The aim of this study was to identify the use of local macrophytes in CMC rearing practice, including Ipomoea aquatic and Oryza sativa. A completely randomized field experiment was conducted to investigate the crab yield, water quality, bacterioplankton community and functions in the three different systems (E. nuttallii, I. aquatic, and O. sativa). Average crab yields in the different macrophyte systems did not differ significantly. The I. aquatic and O. sativa systems significantly decreased the total nitrogen and nitrate-N quantities in the outflow waters during the rearing period compared to the E. nuttallii system, and the I. aquatic and O. sativa plants assimilated more nitrogen than the E. nuttallii plant. Moreover, the significant changes of bacterioplankton abundances and biodiversity in the three systems implied that cleanliness of rearing waters was concomitantly attributed to the differential microbial community and functions. In addition, principle component analysis successfully differentiated the bacterioplankton communities of the three macrophytes systems. Environmental factor fitting and the co-occurrence network analyses indicated that pH was the driver of bacterioplankton community structure. Functional predictions using PICRUSt (v.1.1.3) software based on evolutionary modeling indicated a higher potential for microbial denitrification in the I. aquatic and O. sativa systems. Notably, the O. sativa plants stopped growing in the middle of the rearing period. Thus, the I. aquatic system rather than the O. sativa system could be a feasible and environmental-friendly alternative to the E. nuttallii system in CMC rearing practice.
Project description:Two contemporary effects of humans on aquatic ecosystems are increasing temperatures and increasing nutrient concentrations from fertilizers. The response of organisms to these perturbations has important implications for ecosystem processes. We examined the effects of phosphorus (P) supply and temperature on organismal carbon, nitrogen and phosphorus (C, N, and P) content, cell size and allocation into internal P pools in three strains of recently isolated bacteria (Agrobacterium sp., Flavobacterium sp., and Arthrobacter sp.). We manipulated resource C:P in chemostats and also manipulated temperatures from 10 to 30°C. Dilution rates were maintained for all the strains at ~25% of their temperature-specific maximum growth rate to simulate low growth rates in natural systems. Under these conditions, there were large effects of resource stoichiometry and temperature on biomass stoichiometry, element quotas, and cell size. Each strain was smaller when C-limited and larger when P-limited. Temperature had weak effects on morphology, little effect on C quotas, no effect on N quotas and biomass C:N, but had strong effects on P quotas, biomass N:P and C:P, and RNA. RNA content per cell increased with increasing temperature at most C:P supply ratios, but was more strongly affected by resource stoichiometry than temperature. Because we used a uniform relative growth rate across temperatures, these findings mean that there are important nutrient and temperature affects on biomass composition and stoichiometry that are independent of growth rate. Changes in biomass stoichiometry with temperature were greatest at low P availability, suggesting tighter coupling between temperature and biomass stoichiometry in oligotrophic ecosystems than in eutrophic systems. Because the C:P stoichiometry of biomass affects how bacteria assimilate and remineralize C, increased P availability could disrupt a negative feedback between biomass stoichiometry and C availability.
Project description:Submersed macrophytes have important ecological functions in many streams, but fostering growth of beneficial native species while suppressing weedy invasives may be challenging. Two approaches commonly used in management of terrestrial plant communities may be useful in this context: (1) altering resource availability and (2) establishing desirable species before weeds can invade (priority effects). However, these approaches are rarely used in aquatic systems, despite widespread need for sustainable solutions to aquatic weed problems. In artificial stream channels in California, USA, I conducted experiments with asexual propagules of non-native invasive Myriophyllum spicatum (Eurasian watermilfoil) and native Elodea nuttallii (western waterweed) to address the questions: (1) How does light availability affect relative performance of the two species?; (2) Does planting the native earlier than the invasive decrease survival or growth rate of the invasive?; and (3) Do light level and priority effects interact? The relative performance between E. nuttallii and M. spicatum had an interesting and unexpected pattern: M. spicatum had higher growth rates than E. nuttallii in the zero and medium shade levels, but had similar performance in the low and high shade levels. This pattern is most likely the result of E. nutallii's sensitivity to both very low and very high light, and M. spicatum's sensitivity to very low light only. Native priority did not significantly affect growth rate or survival of M. spicatum, possibly because of unexpectedly poor growth of the E. nuttallii planted early. This study suggests that altering light levels could be effective in reducing growth of an invasive macrophyte, and for changing the competitive balance between a native and a non-native species in the establishment phase. Further investigations into the use of priority effects and resource alteration for submersed macrophyte management are warranted, given their mixed results in other (limited) studies.