Nitrogen is an essential, often limiting, element for biological growth that can act as a pollutant if present in excess. Nitrogen is primarily transported by water from uplands to streams and eventually to recipient lakes, estuaries, and wetlands, but can be modulated by biological uptake and transformation along these flowpaths. As a result, nitrogen can accumulate in aquatic ecosystems if supply is high or if biological retention is low. Dryland and urban ecosystems offer interesting contrasts in water supply, which limits transport and biological activity in drylands, and nitrogen supply that increases with human activity. In my dissertation, I ask: What is the relative balance among nitrogen retention, removal, and transport processes in dryland watersheds, and what is the fate of exported nitrogen? My dissertation research demonstrates that water is a major control on where and when nitrogen is retained and removed versus exported to downstream ecosystems. I used a mass-balance model based on synoptic surveys to study seasonal and spatial patterns in nitrate loading to a dryland stream network. I found that irrigation diversions transport nitrate from agricultural areas to the stream network year-round, even during dry seasons, and are an important driver of nitrate loading. I further explored how seasonal precipitation influences flood nutrient export in an intermittent desert stream by coupling long-term data of flood-water chemistry with stream discharge and precipitation data. I found that higher precipitation prior to a flood fills water storage sites in the catchment, leading to larger floods. In addition, higher antecedent precipitation stimulates biological nitrogen retention in the uplands, leading to lower nitrogen concentration in floods. Finally, I evaluated the consequences of nitrogen export from watersheds on how urban wetlands attenuate nitrate through denitrification that permanently removes nitrogen, and dissimilatory nitrate reduction to ammonium (DNRA) that retains nitrogen in another biologically reactive form. I found that DNRA becomes proportionally more important with low nitrate concentration, thereby retaining nitrogen as ammonium. Collectively, my dissertation research addresses how dryland and urban ecosystems can be integrated into models of watershed nitrogen cycling.
Three dilemmas plague governance of scientific research and technological
innovation: the dilemma of orientation, the dilemma of legitimacy, and the dilemma of control. The dilemma of orientation risks innovation heedless of long-term implications. The dilemma of legitimacy grapples with delegation of authority in democracies, often at the expense of broader public interest. The dilemma of control poses that the undesirable implications of new technologies are hard to grasp, yet once grasped, all too difficult to remedy. That humanity has innovated itself into the sustainability crisis is a prime manifestation of these dilemmas.
Responsible innovation (RI), with foci on anticipation, inclusion, reflection, coordination, and adaptation, aims to mitigate dilemmas of orientation, legitimacy, and control. The aspiration of RI is to bend the processes of technology development toward more just, sustainable, and societally desirable outcomes. Despite the potential for fruitful interaction across RI’s constitutive domains—sustainability science and social studies of science and technology—most sustainability scientists under-theorize the sociopolitical dimensions of technological systems and most science and technology scholars hesitate to take a normative, solutions-oriented stance. Efforts to advance RI, although notable, entail one-off projects that do not lend themselves to comparative analysis for learning.
In this dissertation, I offer an intervention research framework to aid systematic study of intentional programs of change to advance responsible innovation. Two empirical studies demonstrate the framework in application. An evaluation of Science Outside the Lab presents a program to help early-career scientists and engineers understand the complexities of science policy. An evaluation of a Community Engagement Workshop presents a program to help engineers better look beyond technology, listen to and learn from people, and empower communities. Each program is efficacious in helping scientists and engineers more thoughtfully engage with mediators of science and technology governance dilemmas: Science Outside the Lab in revealing the dilemmas of orientation and legitimacy; Community Engagement Workshop in offering reflexive and inclusive approaches to control. As part of a larger intervention research portfolio, these and other projects hold promise for aiding governance of science and technology through responsible innovation.
The overarching question of this dissertation is: how do hydrology, soil conditions, and plant patches affect patterns of denitrification in accidental urban wetlands? To answer this question, I took a three-pronged approach using a combination of field and greenhouse studies. First, I examined drivers of broad patterns of denitrification in accidental urban wetlands. Second, I used a field study to test if plant traits influence denitrification indirectly by modifying soil resources. Finally, I examined how species richness and interactions between species influence nitrate retention and patterns of denitrification using both a field study and greenhouse experiment.
Hydroperiod of accidental urban wetlands mediated patterns of denitrification in response to monsoon floods and plant patches. Specifically, ephemeral wetlands had patterns of denitrification that were largely unexplained by monsoon floods or plant patches, which are common drivers of patterns of denitrification in non-urban wetlands. Several plant traits including belowground biomass, above- and belowground tissue chemistry and rooting depth influenced denitrification indirectly by changing soil organic matter or soil nitrate. However, several other plant traits also had significant direct relationships with denitrification, (i.e. not through the hypothesized indirect relationships through soil organic matter or soil nitrate). This means these plant traits were affecting another aspect of soil conditions not included in the analysis, highlighting the need to improve our understanding of how plant traits influence denitrification. Finally, increasing species richness did not increase nitrate retention or denitrification, but rather individual species had the greatest effects on nitrate retention and denitrification.
An understanding of the formation of spatial heterogeneity is important because spatial heterogeneity leads to functional consequences at the ecosystem scale; however, such an understanding is still limited. Particularly, research simultaneously considering both external variables and internal feedbacks (self-organization) is rare, partly because these two drivers are addressed under different methodological frameworks. In this dissertation, I show the prevalence of internal feedbacks and their interaction with heterogeneity in the preexisting template to form spatial pattern. I use a variety of techniques to account for both the top-down template effect and bottom-up self-organization. Spatial patterns of nutrients in stream surface water are influenced by the self-organized patch configuration originating from the internal feedbacks between nutrient concentration, biological patchiness, and the geomorphic template. Clumps of in-stream macrophyte are shaped by the spatial gradient of water permanence and local self-organization. Additionally, significant biological interactions among plant species also influence macrophyte distribution. The relative contributions of these drivers change in time, responding to the larger external environments or internal processes of ecosystem development. Hydrologic regime alters the effect of geomorphic template and self-organization on in-stream macrophyte distribution. The relative importance of niche vs. neutral processes in shaping biodiversity pattern is a function of hydrology: neutral processes are more important in either very high or very low discharge periods. For the spatial pattern of nutrients, as the ecosystem moves toward late succession and nitrogen becomes more limiting, the effect of self-organization intensifies. Changes in relative importance of different drivers directly affect ecosystem macroscopic properties, such as ecosystem resilience. Stronger internal feedbacks in average to wetter years are shown to increase ecosystem resistance to elevated external stress, and make the backward shifts (vegetation loss) much more gradual. But it causes increases in ecosystem hysteresis effect. Finally, I address the question whether functional consequences of spatial heterogeneity feed back to influence the processes from which spatial heterogeneity emerged through a conceptual review. Such feedbacks are not likely. Self-organized spatial patterning is a result of regular biological processes of organisms. Individual organisms do not benefit from such order. It is order for free, and for nothing.
Further, I investigated the effect of precipitation variation on functional diversity on the same experiment and found a positive response of diversity to increased interannual precipitation variance. Functional evenness showed a similar response resulting from large changes in plant-functional type relative abundance including decreased grass and increased shrub cover while functional richness showed non-significant response. Increased functional diversity ameliorated the direct negative effects of precipitation variation on ecosystem ANPP but did not control ecosystem stability where indirect effects through the dominant plant-functional type determined ecosystem stability.
Analyses of 80 long-term data sets, where I aggregated annual productivity and precipitation data into five-year temporal windows, showed that precipitation variance had a significant effect on aboveground net primary production that is modulated by mean precipitation. Productivity increased with precipitation variation at sites where mean annual precipitation is less than 339 mm but decreased at sites where precipitation is higher than 339 mm. Mechanisms proposed to explain patterns include: differential ANPP response to precipitation among sites, contrasting legacy effects and soil water distribution.
Finally, increased precipitation variance may impact global grasslands affecting plant-functional types in different ways that may lead to state changes, increased erosion and decreased stability that can in turn limit the services provided by these valuable ecosystems.
Humans have dramatically increased phosphorus (P) availability in terrestrial and aquatic ecosystems. As P is often a limiting nutrient of primary production, changes in its availability can have dramatic effects on ecosystem processes. I examined the effects of calcium carbonate (CaCO3) deposition, which can lower P concentrations via coprecipitation of phosphate, on P availability in two systems: streams in the Huachuca Mountains, Arizona, and a stream, Río Mesquites, in Cuatro Ciénegas, México. Calcium carbonate forms as travertine in the former and within the microbialites of the latter. Despite these differences, CaCO3 deposition led to lowered P availability in both systems. By analyzing a three-year dataset of water chemistry from the Huachuca Mountain streams, I determined that P concentrations were negatively related to CaCO3 deposition rates. I also discovered that CaCO3 was positively correlated with nitrogen concentrations, suggesting that the stoichiometric effect of CaCO3 deposition on nutrient availability is due not only to coprecipitation of phosphate, but also to P-related constraints on biotic nitrogen uptake. Building from these observations, bioassays of nutrient limitation of periphyton growth suggest that P limitation is more prevalent in streams with active CaCO3 deposition than those without. Furthermore, when I experimentally reduced rates of CaCO3 deposition within one of the streams by partial light-exclusion, areal P uptake lengths decreased, periphyton P content and growth increased, and periphyton nutrient limitation by P decreased. In Río Mesquites, CaCO3 deposition was also associated with P limitation of microbial growth. There, I investigated the consequences of reductions in CaCO3 deposition with several methods. Calcium removal led to increased concentrations of P in the microbial biomass while light reductions decreased microbial biomass and chemical inhibition had no effect. These results suggest that CaCO3 deposition in microbialites does limit biological uptake of P, that photoautotrophs play an important role in nutrient acquisition, and, combined with other experimental observations, that sulfate reduction may support CaCO3 deposition in the microbialite communities of Río Mesquites. Overall, my results suggest that the effects of CaCO3 deposition on P availability are general and this process should be considered when managing nutrient flows across aquatic ecosystems.
Non-native consumers can significantly alter processes at the population, community, and ecosystem level, and they are a major concern in many aquatic systems. Although the community-level effects of non-native anuran tadpoles are well understood, their ecosystem-level effects have been less studied. Here, I tested the hypothesis that natural densities of non-native bullfrog tadpoles (Lithobates catesbeianus) and native Woodhouse's toad tadpoles (Anaxyrus woodhousii) have dissimilar effects on aquatic ecosystem processes because of differences in grazing and nutrient recycling (excretion and egestion). I measured bullfrog and Woodhouse's carbon, nitrogen, and phosphorus nutrient recycling rates. Then, I determined the impact of tadpole grazing on periphyton biomass (chlorophyll a) during a 39-day mesocosm experiment. Using the same experiment, I also quantified the effect of tadpole grazing and nutrient excretion on periphyton net primary production (NPP). Lastly I measured how dissolved and particulate nutrient concentrations and respiration rates changed in the presence of the two tadpole species. Per unit biomass, I found that bullfrog and Woodhouse's tadpoles excreted nitrogen and phosphorus at similar rates, though Woodhouse's tadpoles egested more carbon, nitrogen, and phosphorus. However, bullfrogs recycled nutrients at higher N:C and N:P ratios. Tadpole excretion did not cause a detectable change in dissolved nutrient concentrations. However, the percent phosphorus in mesocosm detritus was significantly higher in both tadpole treatments, compared to a tadpole-free control. Neither tadpole species decreased periphyton biomass through grazing, although bullfrog nutrient excretion increased areal NPP. This result was due to higher biomass, not higher biomass-specific productivity. Woodhouse's tadpoles significantly decreased respiration in the mesocosm detritus, while bullfrog tadpoles had no effect. This research highlights functional differences between species by showing non-native bullfrog tadpoles and native Woodhouse's tadpoles may have different effects on arid, aquatic ecosystems. Specifically, it indicates bullfrog introductions may alter primary productivity and particulate nutrient dynamics.