Methods of estimating the half-life of biological activity of toxic chemicals in water
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Lake eutrophication is an economic, recreational, and aesthetic problem that affects every lake of the world. Eutrophication is the natural process of lake aging, and progresses irrespective of man's activities. Pollution, however, can hasten the natural rate of aging and shorten the life expectancy of a body of water. The eutrophication of a lake consists of the gradual progression from one life stage to another based on the degree of nourishment or productivity. The extinction of a lake is attributed to enrichment by nutritive materials, biological productivity, decay, and sedimentation. Presently used methods for retarding eutrophication are the abatement of cultural enrichment, treatment of eutrophic symptoms, and control of fundamental causes.
Detecting individuals of amphibian and reptile species can be a daunting task. Detection can be hindered by various factors such as cryptic behavior, color patterns, or observer experience. These factors complicate the estimation of state variables of interest (e.g., abundance, occupancy, species richness) as well as the vital rates that induce changes in these state variables (e.g., survival probabilities for abundance; extinction probabilities for occupancy). Although ad hoc methods (e.g., counts uncorrected for detection, return rates) typically perform poorly in the face of no detection, they continue to be used extensively in various fields, including herpetology. However, formal approaches that estimate and account for the probability of detection, such as capture-mark-recapture (CMR) methods and distance sampling, are available. In this paper, we present classical approaches and recent advances in methods accounting for detectability that are particularly pertinent for herpetological data sets. Through examples, we illustrate the use of several methods, discuss their performance compared to that of ad hoc methods, and we suggest available software to perform these analyses. The methods we discuss control for imperfect detection and reduce bias in estimates of demographic parameters such as population size, survival, or, at other levels of biological organization, species occurrence. Among these methods, recently developed approaches that no longer require marked or resighted individuals should be particularly of interest to field herpetologists. We hope that our effort will encourage practitioners to implement some of the estimation methods presented herein instead of relying on ad hoc methods that make more limiting assumptions.
The U.S. Geological Survey (USGS) is focused on the development of early detection and effective response tools that promote an adaptive management approach to chronic wasting disease (CWD). USGS scientists across the United States are working to understand the biology of CWD, assess and predict the spread and persistence in wildlife and the environment, and develop tools for early detection and control.
The purpose of this study is 1. to document the geographic distribution of Holocene (deposited in the last 11,000 years) sedimentary lithologies in and along the Spokane River, 2. to document the downstream variation of metal contents (especially lead [Pb] and zinc [Zn] ) by sedimentary lithology, and 3. to compare the metal contents of different sedimentary lithologies. This data is used to gain some understanding of the physical and chemical processes that control those metal contents. It is hoped this study can be used to guide potential future remedial actions aimed at reducing the biologic impact of metal-enriched sediments in this area. This work was undertaken in cooperation with the Washington Department of Ecology and the Environmental Protection Agency.
We compared the influence of biological traits (morphology, physiology, reproduction, and life history), ecological traits (geographic distribution, habitat associations, food habits), and introduction attributes (propagule pressure, human use of a species, residence time) on invasion success of native and introduced stream fishes during the colonisation and spread stages in an Appalachian (U.S.A.) river basin. Colonisation success was positively related to residence time, benthic feeding, an equilibrium life-history strategy, and nest spawning. Successful spread was associated with tolerance to increased temperature and an equilibrium life-history strategy. The spread of introduced fishes was negatively related to gamefish status. No effect of propagule pressure was detected. Traits linked to invasion success were consistent with the hypothesis that human land-use practices increase the invasibility of highland catchments by creating novel conditions better suited to lowland and equilibrium invaders. We found biological traits to be more useful than ecological traits in predicting invasion success and suggesting invasion mechanisms. Parental care and nest association can facilitate invasions over large spatial extents for both native and introduced fishes. Analyses of suites of traits can reveal mechanisms of invasions and tactics for controlling them; such mechanisms and tactics may be system-specific and scale-dependent.
Sakhalin taimen ( Parahucho perryi ) range from the Russian Far East mainland along the Sea of Japan coast, and Sakhalin, Kuril, and Hokkaido Islands and are considered to primarily be an anadromous species. We used otolith strontium-to-calcium ratios (Sr/Ca) to determine the chronology of migration between freshwater and saltwater and identify migratory contingents of taimen collected from the Koppi River, Russia. In addition, we examined taimen from the Sarufutsu River, Japan and Tumnin River, Russia that were captured in marine waters. Transects of otolith Sr/Ca for the Sarufutsu River fish were consistent with patterns observed in anadromous salmonids. Two fish from the Tumnin River appeared to be recent migrants to saltwater and one fish was characterized by an otolith Sr/Ca transect consistent with marine migration. Using these transects as benchmarks, all Koppi River taimen were classified as freshwater residents. These findings suggest more work is needed to assess life history variability among locations and the role of freshwater productivity in controlling migratory behavior in taimen.
Hydraulic characteristics inside two research circular tanks (1.5-m and 1.2-m diameter) with the same volume of water were studied to understand how they might affect experimental bias by influencing the behavior and development of juvenile fish. Water velocities inside each tank were documented extensively and flow behavior studied. Surface inflow to the 1.5-m tank created a highly turbulent and aerated surface, and produced unevenly distributed velocities within the tank. A low-flow velocity, or "dead" zone, persisted just upstream of the surface inflow. A single submerged nozzle in the 1.2-m tank created uniform flow and did not cause undue turbulence or introduce air. Flow behavior in the 1.5-m tank is believed to have negatively affected the feeding behavior and physiological development of a group of juvenile fall chinook salmon, Oncorhynchus tshawytscha. A new inflow nozzle design provided comparable flow behavior regardless of tank size and water depth. Maintaining similar hydraulic conditions inside tanks used for various biological purposes, including fish research, would minimize experimental bias caused by differences in flow behavior. Other sources of experimental bias are discussed and recommendations given for reporting and control of experimental conditions in fishery research tank experiments.
Indian Creek is one of the most urban drainage basins in Johnson County, Kansas, and environmental and biological conditions of the creek are affected by contaminants from point and other urban sources. The Johnson County Douglas L. Smith Middle Basin (hereafter referred to as the “Middle Basin”) and Tomahawk Creek Wastewater Treatment Facilities (WWTFs) discharge to Indian Creek. In summer 2010, upgrades were completed to increase capacity and include biological nutrient removal at the Middle Basin facility. There have been no recent infrastructure changes at the Tomahawk Creek facility; however, during 2009, chemically enhanced primary treatment was added to the treatment process for better process settling before disinfection and discharge with the added effect of enhanced phosphorus removal. The U.S. Geological Survey, in cooperation with Johnson County Wastewater, assessed the effects of wastewater effluent on environmental and biological conditions of Indian Creek by comparing two upstream sites to four sites located downstream from the WWTFs using data collected during June 2004 through June 2013. Environmental conditions were evaluated using previously and newly collected discrete and continuous data and were compared with an assessment of biological community composition and ecosystem function along the upstream-downstream gradient. This study improves the understanding of the effects of wastewater effluent on stream-water and streambed sediment quality, biological community composition, and ecosystem function in urban areas. After the addition of biological nutrient removal to the Middle Basin WWTF in 2010, annual mean total nitrogen concentrations in effluent decreased by 46 percent, but still exceeded the National Pollutant Discharge Elimination System (NPDES) wastewater effluent permit concentration goal of 8.0 milligrams per liter (mg/L); however, the NPDES wastewater effluent permit total phosphorus concentration goal of 1.5 mg/L or less was achieved at the Middle Basin WWTF. At the Tomahawk Creek WWTF, after the addition of chemically enhanced primary treatment in 2009, effluent discharges also had total phosphorus concentrations below 1.5 mg/L. After the addition of biological nutrient removal, annual total nitrogen and phosphorus loads from the Middle Basin WWTF decreased by 42 and 54 percent, respectively, even though effluent volume increased by 11 percent. Annual total phosphorus loads from the Tomahawk Creek WWTF after the addition of chemically enhanced primary treatment decreased by 54 percent despite a 33-percent increase in effluent volume. Total nitrogen and phosphorus from the WWTFs contributed between 30 and nearly 100 percent to annual nutrient loads in Indian Creek depending on streamflow conditions. In-stream total nitrogen primarily came from wastewater effluent except during years with the highest streamflows. Most of the in-stream total phosphorus typically came from effluent during dry years and from other urban sources during wet years. During 2010 through 2013, annual mean discharge from the Middle Basin WWTF was about 75 percent of permitted design capacity. Annual nutrient loads likely will increase when the facility is operated at permitted design capacity; however, estimated maximum annual nutrient loads from the Middle Basin WWTF were 27 to 38 percent lower than before capacity upgrades and the addition of biological nutrient removal to treatment processes. Thus, the addition of biological nutrient removal to the Middle Basin wastewater treatment process should reduce overall nutrient loads from the facility even when the facility is operated at permitted design capacity. The effects of wastewater effluent on the water quality of Indian Creek were most evident during below-normal and normal streamflows (about 75 percent of the time) when wastewater effluent represented about 24 percent or more of total streamflow. Wastewater effluent had the most substantial effect on nutrient concentrations in Indian Creek. Total and inorganic nutrient concentrations at the downstream sites during below-normal and normal streamflows were 10 to 100 times higher than at the upstream sites, even after changes in treatment practices at the WWTFs. Median total phosphorus concentrations during below-normal and normal streamflows at a downstream site were 43 percent lower following improvements in wastewater treatment processes. Similar decreases in total nitrogen were not observed, likely because total nitrogen concentrations only decreased in Middle Basin effluent and wastewater contributed a higher percentage to streamflows when nutrient samples were collected during the after-upgrade period. The wastewater effluent discharges to Indian Creek caused changes in stream-water quality that may affect biological community structure and ecosystem processes, including higher concentrations of bioavailable nutrients (nitrate and orthophosphorus) and warmer water temperatures during winter months. Other urban sources of contaminants also caused changes in stream-water quality that may affect biological community structure and ecosystem processes, including higher turbidities downstream from construction areas and higher specific conductance and chloride concentrations during winter months. Chloride concentrations exceeded acute and chronic exposure criteria at all Indian Creek study sites, regardless of wastewater influence, for weeks or months during winter. Streambed sediment chemistry was affected by wastewater (elevated nutrient and organic wastewater-indicator compound concentrations) and other contaminants from urban sources (elevated polyaromatic hydrocarbon concentrations). Overall habitat conditions were suboptimal or marginal at all sites; general decline in habitat conditions along the upstream-downstream gradient likely was caused by the cumulative effects of urbanization with increasing drainage basin size. Wastewater effluent likely affected algal periphyton biomass and community composition, primary production, and community respiration in Indian Creek. Functional stream health, evaluated using a preliminary framework based on primary production and community respiration, was mildly or severely impaired at most downstream sites relative to an urban upstream Indian Creek site. The mechanistic cause of the changes in these biological variables are unclear, though elevated nutrient concentrations were positively correlated with algal biomass, primary production, and community respiration. Macroinvertebrate communities indicated impairment at all sites, and Kansas Department of Health and Environment aquatic life support scores indicated conditions nonsupporting of aquatic life, regardless of wastewater influences. Urban influences, other than wastewater effluent discharge, likely control macroinvertebrate community structure in Indian Creek. Changes in treatment processes at the Middle Basin and Tomahawk Creek WWTFs improved wastewater effluent quality and decreased nutrient loads, but wastewater effluent discharges still had negative effects on the environmental and biological conditions at downstream Indian Creek sites. Wastewater effluent discharge into Indian Creek likely contributed to changes in measures of ecosystem structure (streamflow, water and streambed-sediment chemistry, algal biomass, and algal periphyton community composition) and function (primary production and community respiration) along the upstream-downstream gradient. Wastewater effluent discharges maintained streamflows and increased nutrient concentrations, algal biomass, primary production, and community respiration at the downstream sites. Functional stream health was severely impaired downstream from the Middle Basin WWTF and mildly impaired downstream from the Tomahawk WWTF relative to the urban upstream site. As distance from the Middle Basin WWTF increased, nutrient concentrations, algal biomass, primary production, and community respiration decreased, and functional stream health was no longer impaired 9.5 kilometers downstream from the discharge relative to the urban upstream site. Therefore, although wastewater effluent caused persistent changes in environmental and biological conditions and functional stream health at sites located immediately downstream from WWTF effluent discharges, some recovery to conditions more similar to the urban upstream site occurred within a relatively short distance.
Spawning stock biomass (SSB) is often used as an index for reproductive potential (RP) in fisheries stock assessments. This method assumes that mature female biomass is proportional to total egg production and implies that (1) the fecundity–length relationship follows a cubic function or (2) relative fecundity is constant. For many marine fishes, adequate fecundity estimates to evaluate these relationships are lacking. This study estimated fecundity and fecundity relationships for Yelloweye Rockfish Sebastes ruberrimus and evaluated an automated method of counting eggs and larvae. We collected Yelloweye Rockfish ovaries ( N = 90) from the northern Gulf of Alaska, including Prince William Sound, Alaska, during 2018–2019 and used the gravimetric method and image analysis software to count eggs from digital camera images. To evaluate the speed, accuracy, and precision of the automated counting procedure, one-third of the gravimetric samples were also manually counted. Image analysis software was approximately four times faster but equally accurate and precise for fecundity estimates relative to manual counts. Fecundity ranged from 53,249 to 3.052 × 10 6 eggs (mean ± SD = 896,762 ± 699,504 eggs), and relative fecundity increased with female FL and ranged from 68 to 435 eggs/g of body weight (mean ± SD = 226 ± 87 eggs/g). The use of SSB for Yelloweye Rockfish stock assessment could underestimate the contribution to egg production by larger (>5.6-kg) females, overestimate the contribution by smaller females, and lead to biased biological reference points. This study provides critical information to more realistically model RP and improve stock assessment inputs for the development of harvest control rules for Yelloweye Rockfish. Additionally, the use of image analysis software to count eggs in digital images proved to be an effective fecundity estimation method that could be applied to other highly fecund fish species for which the time demand of manual counting methods would be prohibitive.
This special issue of Geo-Marine Letters, "Benthic Boundary Layer Processes in the Lower Florida Keys," includes 12 papers that present preliminary results from the Key West Campaign. The Dry Tortugas and Marquesas Keys test sites were selected by a group of 115 scientists and technicians to study benthic boundary layer processes in a carbonate environment controlled by bioturbation and biogeochemical processes. Major activities included remote sediment classification; high-frequency acoustic scattering experiments; sediment sampling for radiological, geotechnical, biological, biogeochemical, physical, and geoacoustic studies; and hydrodynamic studies using an instrumented tetrapod. All these data are being used to improve our understanding of the effects of environmental processes on sediment structure and behavior.
Climate change predictions for the desert southwestern U.S. are for shifts in precipitation patterns. The impacts of climate change may be significant, because desert soil processes are strongly controlled by precipitation inputs ('pulses') via their effect on soil water availability. This study examined the response of soil respiration-an important biological process that affects soil carbon (C) storage-to variation in pulses representative of climate change scenarios for the Sonoran Desert. Because deserts are mosaics of different plant cover types and soil textures-which create patchiness in soil respiration-we examined how these landscape characteristics interact to affect the response of soil respiration to pulses. Pulses were applied to experimental plots of bare and vegetated soil on contrasting soil textures typical of Sonoran Desert grasslands. The data were analyzed within a Bayesian framework to: (1) determine pulse size and antecedent moisture (soil moisture prior to the pulse) effects on soil respiration, (2) quantify soil texture (coarse vs. fine) and cover type (bare vs. vegetated) effects on the response of soil respiration and its components (plant vs. microbial) to pulses, and (3) explore the relationship between long-term variation in pulse regimes and seasonal soil respiration. Regarding objective (1), larger pulses resulted in higher respiration rates, particularly from vegetated fine-textured soil, and dry antecedent conditions amplified respiration responses to pulses (wet antecedent conditions dampened the pulse response). Regarding (2), autotrophic (plant) activity was a significant source (???60%) of respiration and was more sensitive to pulses on coarse- versus fine-textured soils. The sensitivity of heterotrophic (microbial) respiration to pulses was highly dependent on antecedent soil water. Regarding (3), seasonal soil respiration was predicted to increase with both growing season precipitation and mean pulse size (but only for pulses between 7 and 25 mm). Thus, the heterogeneity of the desert landscape and the timing or the number of medium-sized pulses is expected to significantly impact desert soil C loss with climate change. ?? 2008 Springer Science+Business Media, LLC.
In dryland ecosystems, natural recovery of biological soil crusts (biocrusts) following disturbance may be slow or inhibited, necessitating active restoration practices. While biocrusts can be readily propagated under environmentally controlled conditions, rehabilitation in the field is complicated by environmental stresses which may be particularly acute in degraded, destabilized soils with harsh climatic conditions at the soil surface. In this study, we first present the results of a field trial at a severely degraded rangeland site examining the stabilizing effects of various soil amendments (polysaccharide glues and polyacrylamides) in combination with biocrust inoculum. We found that a psyllium compound was the only amendment to maintain effectiveness after 19 months, and the only treatment that maintained biocrust inoculum throughout the trial. In a subsequent short‐term experiment where plots were shaded and watered, we examined how biocrust inoculation rate (0, 20, and 40% initial cover) and the psyllium‐based amendment affected biocrust growth. After 4 months, visible biocrust cover in inoculated plots was greater than in controls, but only chlorophyll a exhibited a dosage‐response to inoculum application rate, indicating preferential establishment of cyanobacteria. Psyllium did not affect biocrust development but did improve soil stability. Shade and watering buffered against temperature extremes (up to 15°C) and increased the duration of moist surface conditions necessary for biocrust growth by up to 30%, mimicking conditions more common in the fall and winter months. Our results suggest that inducing early successional biocrusts on a highly degraded site is possible with suitable microclimate conditions.
Disease emergence represent a global threat for public health, economy, and biological conservation and most of the emerging diseases have zoonotic origin from wildlife. To prevent their spread and to support the implementation of control measures, disease surveillance and reporting systems are needed, and due to globalisation, these activities should be carried at world level. To define the main gaps affecting the performances of wildlife health surveillance and reporting systems at word level, we analysed the Wildlife Disease Surveillance Survey submitted to all Members of World Organisation for Animal Health which inquire on structure and limits of surveillance and reporting systems. The response from 103 Members, covering all world areas, showed that 54.4% of them have a wildlife disease surveillance programme and 66% implemented a strategy to manage their spread. The lack of dedicated budget affected the possibility of outbreak investigations, sampling collection and diagnostic testing. Although most Members maintain records relating to wildlife mortality or morbidity events in centralised databases, data analyses and disease risk assessment are reported as main needs. Our evaluation of surveillance capacity showed an overall low level associated to great variability between Members not restricted to specific geographical area. A better recognition of the value of wildlife disease surveillance resulting in allocation of adequate resources by governments would make wildlife disease surveillance systems more functional. Moreover, the future comprehension of the influence of socio-economic, cultural and biodiversity aspects will be essential to increase better disease surveillance under One Health approach.
The accumulation of polycyclic aromatic hydrocarbons (PAH) in soil, plants, and water may impart negative effects on ecosystem and human health. We quantified the concentration and distribution of 41 PAH (n = 32), organic C, total N, and S (n = 140) and investigated PAH sources using a chronosequence of floodplain soils under a natural vegetation succession. Soil samples were collected between 0- and 260-cm depth in bare land (the control), wetland, forest, and grassland areas near a closed municipal landfill and an active asphalt plant (the contaminant sources) in the north bank of the Canadian River near Norman, OK. Principal component, cluster, and correlation analyses were used to investigate the spatial distribution of PAH, in combination with diagnostic ratios to distinguish pyrogenic vs. petrogenic PAH suites. Total PAH concentration (SigmaPAH) had a mean of 1300 ng g(-1), minimum of 16 ng g(-1), and maximum of 12,000 ng g(-1). At 0- to 20-cm depth, SigmaPAH was 3500 +/- 1600 ng g(-1) (mean +/- 1 SE) near the contaminant sources. The most common compounds were nonalkylated, high molecular weight PAH of pyrogenic origin, i.e., fluoranthene (17%), pyrene (14%), phenanthrene (9%), benzo(b)fluoranthene (7%), chrysene (6%), and benzo(a)anthracene (5%). SigmaPAH in the control (130 +/- 23 ng g(-1)) was comparable to reported concentrations for the rural Great Plains. Perylene had a unique distribution pattern suggesting biological inputs. The main PAH contamination mechanisms were likely atmospheric deposition due to asphalt production at the 0- to 20-cm depth and past landfill operations at deeper depths.
The parasitic sea lamprey (Petromyzon marinus) has been a serious pest since its introduction into the Great Lakes, where it contributed to severe imbalances in the fish communities by selectively removing large predators (Smith 1968; Christie 1974; Schneider et al.1996). Since the 1950s, restoration and maintenance of predator-prey balance has depended on the Great Lakes Fishery Commission (GLFC) sea lamprey management program. Initially, management relied primarily on stream treatments with a selective lampricide to kill larvae, on barriers to migration, and on trapping to remove potential spawners (Smith and Tibbles 1980). By the late 1970s, however, it was clear that the future of sea lamprey management lay in development of a larger array of control strategies, including more alternatives to lampricide applications (Sawyer 1980). Since then the only new alternative to chemical control to reach operational status is the release of sterilized male sea lampreys. Research on the concept began at the USGS, Hammond Bay Biological Station in Millersburg, MI (HBBS) during the 1970s (Hanson and Manion 1980). Development and evaluation continued through the 1980s, leading to the release of sterilized males in Great Lakes tributaries since 1991 (Twohey et al. 2003a). The objectives of this paper are 1) to review the implementation and evaluations of sterile-male-release technique (SMRT) as it is being applied against sea lampreys in the Great Lakes, 2) to review our current understanding of its efficacy, and 3) to identify additional research areas and topics that would increase either the efficacy of SMRT or expand its geographic potential for application.
A series of well-developed submerged coral reefs are preserved in the Huon Gulf (Papua New Guinea) and around Hawaii. Despite different tectonics settings, both regions have experienced rapid subsidence (2-6??m/ka) over the last 500??ka. Rapid subsidence, combined with eustatic sea-level changes, is responsible for repeated drowning and backstepping of coral reefs over this period. Because we can place quantitative constraints on these systems (i.e., reef drowning age, eustatic sea-level changes, subsidence rates, accretion rates, basement substrates, and paleobathymetry), these areas represent unique natural laboratories for exploring the roles of tectonics, reef accretion, and eustatic sea-level changes in controlling the evolution of individual reefs, as well as backstepping of the entire system. A review of new and existing bathymetric, radiometric, sedimentary facies and numerical modeling data indicate that these reefs have had long, complex growth histories and that they are highly sensitive, recording drowning not only during major deglaciations, but also during high-frequency, small-amplitude interstadial and deglacial meltwater pulse events. Analysis of five generalized sedimentary facies shows that reef drowning is characterized by a distinct biological and sedimentary sequence. Observational and numerical modeling data indicate that on precessional (20??ka) and sub-orbital timescales, the rate and amplitude of eustatic sea-level changes are critical in controlling initiation, growth, drowning or sub-aerial exposure, subsequent re-initiation, and final drowning. However, over longer timescales (> 100-500??ka) continued tectonic subsidence and basement substrate morphology influence broad scale reef morphology and backstepping geometries. Drilling of these reefs will yield greatly expanded stratigraphic sections compared with similar reefs on slowly subsiding, stable and uplifting margins, and thus they represent a unique archive of sea-level and climate changes, as well as a record of the response of coral reefs to these changes over the last six glacial cycles. ?? 2008 Elsevier B.V. All rights reserved.
The types and amounts of carbon (C) and nitrogen (N) inputs, as well as irrigation management are likely to influence gaseous emissions and microbial ecology of agricultural soil. Carbon dioxide (CO2) and nitrous oxide (N2O) efflux, with and without acetylene inhibition, inorganic N, and microbial biomass C were measured after irrigation or simulated rainfall in two agricultural fields under tomatoes (Lycopersicon esculentum). The two fields, located in the California Central Valley, had either a history of high organic matter (OM) inputs ("organic" management) or one of low OM and inorganic fertilizer inputs ("conventional" management). In microcosms, where short-term microbial responses to wetting and drying were studied, the highest CO2 efflux took place at about 60% water-filled pore space (WFPS). At this moisture level, phospholipid fatty acids (PLFA) indicative of microbial nutrient availability were elevated and a PLFA stress indicator was depressed, suggesting peak microbial activity. The highest N 2O efflux in the organically managed soil (0.94 mg N2O-N m-2 h-1) occurred after manure and legume cover crop incorporation, and in the conventionally managed soil (2.12 mg N2O-N m-2 h-1) after inorganic N fertilizer inputs. Elevated N2O emissions occurred at a WFPS >60% and lasted <2 days after wetting, probably because the top layer (0-150 mm) of this silt loam soil dried quickly. Therefore, in these cropping systems, irrigation management might control the duration of elevated N2O efflux, even when C and inorganic N availability are high, whereas inorganic N concentrations should be kept low during times when soil moisture cannot be controlled.