skip to main content

Title: Electrolytes on the prairie: How urine‐like additions of Na and K shape the dynamics of a grassland food web

The electrolytes Na and K both function to maintain water balance and membrane potential. However, these elements work differently in plants—where K is the primary electrolyte—than in animals—where ATPases require a balanced supply of Na and K. Here, we use monthly factorial additions of Na and K to simulate bovine urine inputs and explore how these electrolytes ramify through a prairie food web. Against a seasonal trend of increasing grass biomass and decreasing water and elemental tissue concentrations, +K and +Na plots boosted water content and, when added together, plant biomass. Compared to control plots, +Na and +K plots increased element concentrations in above‐ground plant tissue early in summer and decreased them in September. Simultaneously, invertebrate abundance on Na and K additions were sequentially higher and lower than control plots from June to September and were most suppressed when grass was most nutrient rich. K was the more effective plant electrolyte, but Na frequently promoted similar changes in grass ionomes. The soluble/leachable ions of Na and K showed significant ability to shape plant growth, water content, and the 15‐element ionome, with consequences for higher trophic levels. Grasslands with high inputs of Na and K—via large mammal grazers or coastal aerosol deposition—likely enhance the ability of plants to adjust their above‐ground ionomes, with dramatic consequences for the distribution of invertebrate consumers.

more » « less
Award ID(s):
1702426 2025849
Author(s) / Creator(s):
Publisher / Repository:
Wiley Blackwell (John Wiley & Sons)
Date Published:
Journal Name:
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Sodium (Na) is an essential element for all animals, but not for plants. Soil Na supplies vary geographically. Animals that primarily consume plants thus have the potential to be Na limited and plants that uptake Na may be subject to higher rates of herbivory, but their high Na content also may attract beneficial partners such as pollinators and seed dispersers.

    To test for the effects of Na biogeochemistry on herbivory, we conducted distributed Na press experiments (monthly Na application across the growing season) in four North American grasslands.

    Na addition increased soil and plant Na concentrations at all sites. Grasses in Na addition plots had significantly higher herbivore damage by leaf miners and fungal pathogens than those in control plots. Forbs with higher foliar Na concentrations had significantly more chewing insect herbivore and fungal damage.

    While no pattern was evident across all species, several forb species had higher Na concentrations in inflorescences compared to leaves, suggesting they may allocate Na to attract beneficial partners.

    The uptake of Na by plants, and animal responses, has implications for the salinification in the Anthropocene. Increased use of road salt, irrigation with saline groundwater, rising sea levels and increasing temperatures and evapotranspiration rates with climate change can all increase inputs of Na into terrestrial ecosystems.

    Our results suggest increasing terrestrial Na availability will benefit insect herbivores and plant fungal pathogens.

    A freePlain Language Summarycan be found within the Supporting Information of this article.

    more » « less
  2. Abstract

    Plant elemental content can vary up to 1,000‐fold across grasslands, with implications for the herbivores the plants feed. We contrast the regulation, in grasses and forbs, of 12 elements essential to plants and animals (henceforth plant‐essential), 7 essential to animals but not plants (animal‐essential) and 6 with no known metabolic function (nonessential). Four hypotheses accounted for up to two thirds of the variation in grass and forb ionomes across 54 North American grasslands. Consistent with the supply‐side hypothesis, the plant‐essential ionome of both forbs and grasses tracked soil availability. Grass ionomes were more likely to harvest even nonessential elements like Cd and Sr. Consistent with the grazing hypothesis, cattle‐grazed grasslands also accumulated a handful of metals like Cu and Cr. Consistent with the NP‐catalysis hypothesis, increases in the macronutrients N and P in grasses were associated with higher densities of cofactors like Zn and Cu. The plant‐essential elements of forbs, in contrast, consistently varied as per the nutrient‐dilution hypothesis—there was a decrease in elemental parts per million with increasing local carbohydrate production. Combined, these data fit a working hypothesis that grasses maintain lower elemental densities and survive on nutrient‐poor patches by opportunistically harvesting soil nutrients. In contrast, nutrient‐rich forbs use episodes of high precipitation and temperature to build new carbohydrate biomass, raising leaves higher to compete for light, but diluting the nutrient content in every bite of tissue. Herbivores of forbs may thus be particularly prone to increases inpCO2via nutrient dilution.

    more » « less
  3. Abstract Aim

    Multiple hypotheses predict how gradients of nutrient availability, plant biomass, and temperature shape trophic pyramids. We aim to disentangle the simultaneous influence of those factors and their indirect effects on trophic structure and individual trophic levels.


    United States.

    Time period


    Major taxa studied



    To examine differences in trophic pyramid shape and abundance within trophic levels and across ecological gradients, we conducted 54 standardized surveys of invertebrate communities in North American grasslands. We tested for the direct and indirect effects of plant biomass, temperature, sodium (Na), other essential elements (e.g. N, P, and K), and toxic heavy metals, (e.g. Ar and Pb) in plant tissue on both individual trophic levels, and trophic pyramid shape, estimated as the community trophic mean (CTM).


    Plant sodium increased CTM, indicating that high plant sodium concentrations are associated with top‐heavy invertebrate trophic pyramids. Sites with higher plant biomass had higher proportions of herbivores compared to higher trophic levels. Finally, increasing temperature resulted in more top‐heavy trophic pyramids. Overall, plant biomass, temperature, and plant chemistry directly and indirectly affected the abundances within different trophic levels, highlighting the complexity of factors regulating trophic structure.

    Main conclusions

    Trophic structure of grassland invertebrate communities is strongly influenced by plant sodium, plant biomass, and to a lesser extent, temperature. Grasslands occupy 30% of the Earth’s terrestrial surface and are an imperiled ecosystem due to conversion to row crop agriculture. As biogeochemistry and temperature in the Anthropocene are increasingly modified, our results have considerable implications for the trophic structure of future grassland communities.

    more » « less
  4. Abstract

    The impacts of altered biogeochemical cycles on ecological systems are likely to vary with trophic level. Predicting how these changes will affect ecological food webs is further complicated by human activities, which are simultaneously altering the availability of macronutrients like nitrogen (N) and phosphorus (P), and micronutrients such as sodium (Na). Here we contrast three hypotheses that predict how increasing nutrient availability will shape grassland food webs. We conducted a distributed factorial fertilization experiment (N and P crossed with NaCl) across four North American grasslands, quantifying the responses of aboveground plant biomass and volume, plant tissue and soil elemental concentrations, as well as the abundance of five arthropod functional groups. Fertilization with N and P increased plant biomass and foliar N and P concentrations in grasses but not forbs. Fertilization with Na had no effect on plant biomass but increased foliar Na concentrations. Consistent with the nutrient limitation hypothesis, we found strong evidence of nutrient limitation for insect herbivores across the four sites with sucking (phloem and xylem feeding) herbivores increasing in abundance with NP fertilization and chewing herbivores increasing in response to both Na and NP fertilization, and a trend for increased response of arthropods to lower plant nutrient availability. We found no evidence for an interaction of NaCl and NP on arthropod abundance as predicted by the serial colimitation hypothesis. Finally, consistent with the ecosystem size hypothesis, predator and parasitoid abundances increased with plant volume, but not fertilization. Our results suggest these functional group‐specific responses to changes in plant nutrients and structure are key to predicting the future of grassland food webs in an era with increasing use of N and P fertilizers, and increasing terrestrial inputs of Na from road salt, saline irrigation water, and aerosols due to rising sea levels.

    more » « less
  5. Hu, Shuijin (Ed.)
    Abstract Aims Linkages formed through aquatic–terrestrial subsidies can play an important role in structuring communities and mediating ecosystem functions. Aquatic–terrestrial subsidies may be especially important in nutrient-poor ecosystems, such as the freshwater sand dunes surrounding Lake Michigan. Adult midges emerge from Lake Michigan in the spring, swarm to mate and die. Their carcasses form mounds at the base of plants, where they may increase plant productivity through their nutrient inputs. However, the effect of aquatic–terrestrial subsidies on plant productivity could depend on other biotic interactions. In particular, soil microbes might play a key role in facilitating the conversion of nutrients to plant-available forms or competing for the nutrients with plants. Methods In a greenhouse experiment, we tested how carcasses from lake emergent midges (Chironomidae) and soil microbes independently and interactively influenced the performance of a common dune grass, Calamovilfa longifolia. To determine whether midges influenced abiotic soil properties, we measured how midge additions influenced soil nutrients and soil moisture. Important Findings Midges greatly increased plant biomass, while soil microbes influenced the magnitude of this effect. In the absence of soil microbes plant biomass was seven times greater with midges than without midges. However, in the presence of soil microbes, plant biomass was only three times greater. The effect of midges might be driven by their nutrient inputs into the soil, as midges contained 100 times more N, 10 times more P and 150 times more K than dune soils did. Our results suggest that soil microbes may be competing with plants for these nutrients. In sum, we found that midges can be an important aquatic–terrestrial subsidy that produces strong, positive effects on plant productivity along the shorelines of Lake Michigan, but that the impact of aquatic–terrestrial subsidies must be considered within the context of the complex interactions that take place within ecological communities. 
    more » « less