Precipitation control over inorganic nitrogen import-export budgets across watersheds: a synthesis of long-term ecological research

ABSTRACT We investigated long-term and seasonal patterns of N imports and exports, as well as patterns following climate perturbations, across biomes using data from 15 watersheds from nine Long-Term Ecological Research (LTER) sites in North America. Mean dissolved inorganic nitrogen (DIN) import-export budgets (N import via precipitation-N export via stream flow) for common years across all watersheds was highly variable, ranging from a net loss of 0Ð17 š 0Ð09 kg N ha 1 mo 1 to net retention of 0Ð68 š 0Ð08 kg N ha 1 mo 1 . The net retention of DIN decreased (smaller import-export budget) with increasing precipitation, as well as with increasing variation in precipitation during the winter, spring, and fall. Averaged across all seasons, net DIN retention decreased as the coefficient of variation (CV) in precipitation increased across all sites (r 2 D 0Ð48, p D 0Ð005). This trend was made stronger when the disturbed watersheds were withheld from the analysis (r 2 D 0Ð80, p < 0Ð001, n D 11). Thus, DIN exports were either similar to or exceeded imports in the tropical, boreal, and wet coniferous watersheds, whereas imports exceeded exports in temperate deciduous watersheds. In general, forest harvesting, hurricanes, or floods corresponded with periods of increased DIN exports relative to imports. Periods when water throughput within a watershed was likely to be lower (i.e. low snow pack or El Niño years) corresponded with decreased DIN exports relative to imports. These data provide a basis for ranking diverse sites in terms of their ability to retain DIN in the context of changing precipitation regimes likely to occur in the future

Hillslope Hydrology in Global Change Research and Earth System Modeling

Earth System Models (ESMs) are essential tools for understanding and predicting global change, but they cannot explicitly resolve hillslope‐scale terrain structures that fundamentally organize water, energy, and biogeochemical stores and fluxes at subgrid scales. Here we bring together hydrologists, Critical Zone scientists, and ESM developers, to explore how hillslope structures may modulate ESM grid‐level water, energy, and biogeochemical fluxes. In contrast to the one‐dimensional (1‐D), 2‐ to 3‐mdeep, and free‐draining soil hydrology in most ESM land models, we hypothesize that 3‐D, lateral ridge‐to‐valley flow through shallow and deep paths and insolation contrasts between sunny and shady slopes are the top two globally quantifiable organizers of water and energy (and vegetation) within an ESM grid cell. We hypothesize that these two processes are likely to impact ESM predictions where (and when) water and/or energy are limiting. We further hypothesize that, if implemented in ESM land models, these processes will increase simulated continental water storage and residence time, buffering terrestrial ecosystems against seasonal and interannual droughts. We explore efficient ways to capture these mechanisms in ESMs and identify critical knowledge gaps preventing us from scaling up hillslope to global processes. One such gap is our extremely limited knowledge of the subsurface, where water is stored (supporting vegetation) and released to stream baseflow (supporting aquatic ecosystems). We conclude with a set of organizing hypotheses and a call for global syntheses activities and model experiments to assess the impact of hillslope hydrology on global change predictions

Dissimilatory nitrate uptake in Paracoccus denitrificans via a ΔµH+-dependent system and a nitrate-nitrite antiport system

Respiration-driven proton translocation has been studied with the oxidant pulse method for cells of denitrifying Paracoccus denitrificans oxidizing H2 during reduction of O2, NO- 3, NO- 2 or N2O. A simplified scheme of anaerobic electron transport and associated proton translocation is shown that is consistent with the measured H+ oxidant ratios. Furthermore, the kinetics and energetics of NO- 3 uptake in whole cells of P. denitrificans were studied. For this purpose, we measured H2 consumption or N2O production after addition of NO- 3 to a cell suspension, which indirectly gave information about uptake (and reduction) of NO- 3. It was found that a lag phase in H2 consumption or N2O production appeared whenever the membrane potential was dissipated by addition of thiocyanate, carbonyl cyanide m-chlorophenylhydrazone or triphenyl-methylphosphonium bromide. However, these lag phases were not observed when NO- 2 was present at the moment of introduction of NO- 3. On the basis of these findings we conclude that there are two uptake systems for NO- 3. One system is dependent on the proton-motive force and is probably used for initiation of NO- 3 uptake. The other is an NO- 3 NO- 2 antiport and its function is to take over NO- 3 uptake from the first system

The bioenergetics of denitrification

In anaerobically grown Paracoccus denitrificans the dissimilatory nitrate reductase is linked to the respiratory chain at the level of cytochromes b. Electron transport to nitrite and nitrous oxide involves c-type cytochromes. During electron transport from NADH to nitrate one phosphorylation site is passed, whereas two sites are passed during electron transport from NADH to oxygen, nitrite and nitrous oxide. The presentation of a respiratory chain as a linear array of electron carriers gives a misleading picture of the efficiency of energy conservation since the location of the reductases is not taken into account. For the reduction of nitrite and nitrous oxide, protons are utilized from the periplasmic space, whereas for the reduction of oxygen and nitrate, protons are utilized from the cytoplasmic side of the inner membrane. Evidence for two transport systems for nitrate was obtained. One is driven by the proton motive force; this system is used to initiate nitrate reduction. The second system is a nitrate-nitrite antiport system. A scheme for proton translocation and electron transport to nitrate, nitrite, nitrous oxide and oxygen is presented. The number of charges translocated across the membrane during flow of two electrons from NADH is the same for all nitrogenous oxides and is 67-71% of that during electron transfer to oxygen via cytochrome o. These findings are in accordance with growth yield studies. YMAX electron values determined in chemostat cultures for growth with various substrates and hydrogen acceptors are proportional to the number of charges translocated to these hydrogen acceptors during electron transport

Respiration-driven proton translocation with nitrite and nitrous oxide in Paracoccus denitrificans

(1)H+ leads to/electron acceptor ratios have been determined with the oxidant pulse method for cells of denitrifying Paracoccus denitrificans oxidizing endogenous substrates during reduction of O2, NO2- or N2O. Under optimal H+-translocation conditions, the ratios leads to H+/O, H+ leads to/N2O, H+ leads to/NO2- for reduction to N2 and H+ leads to/NO2- for reduction to N2O were 6.0-6.3, 4.02, 5.79 and 3.37, respectively. (2) With ascorbate/N,N,N,'N'-tetramethyl-p-phenylene-diamine as exogenous substrate, addition of NO2- or N2O to an anaerobic cell suspension resulted in rapid alkalinization of the outer bulk medium. H+/N2O, H+/NO2- for reduction to N2 and H+/NO2- for reduction to N2O were -0.84, -2.33 and -1.90, respectively. (3) The H+/oxidant ratios, mentioned in item 2, were not altered in the presence of valinomycin/K+ and the triphenylmethylphosphonium cation. (4) A simplified scheme of electron transport to O2, NO2- and N2O is presented which shows a periplasmic orientation of the nitrite reductase as well as the nitrous oxide reductase. Electrons destined for NO2-, N2O or O2 pass two H+-trans-locating sites. The H+ leads to/electron acceptor ratios predicted by this scheme are in good agreement with the experimental values