Estimation of the starvation losses of nitrogen and energy in the rainbow trout (Oncorhynchus mykiss) with special regard to protein and energy maintenance requirements

Literature data are analysed regarding losses of body substances occurring during a period of food deprivation in rainbow trout ( Oncorhynchus mykiss ). Nitrogen (protein) and energy losses show a distinct dependence on fish mass (FM [g]) and water temperature (T [°C]). Several regression models for this relationship were compared with best testing estimates as follows:
Nitrogen loss [mg N 2 fish⁻¹ 2 d⁻¹] = 0.0658 e^(1.037) 2 FM^0.739
( n = 49, 9–20°C, 5–400g fish mass, P < 0.001, B = 0.826)
Nitrogen-corrected energy loss [J 2 fish⁻¹ 2 d⁻¹] = 22.09 e^(1.034) 2 FM^0.833
( n = 63, 9–25°C, 8–400 g fish mass, P < 0.001, B = 0.887).
For nitrogen loss as well as for nitrogen-corrected energy loss, the metabolic rate shows a progressive increase with rising water temperature. The temperature coefficient increases in magnitude as temperature increases. The introduction of a general common exponent (0.8 instead of 0.739 for nitrogen loss and 0.833 for energy loss) for fish mass decreases the precision of the estimate. The equations could serve as a base for estimating net protein and net energy maintenance requirements of rainbow trout. Possible limitations, caused by uncertainities in estimating the elevated metabolic rate by food intake and ingestion, are discussed.     

Laboratory growth of larval lampreys (Lampetra (Entosphenus) tridentata Richardson) at different food concentrations and animal densities

Lampreys are important research animals. This study investigates some of the Parameters important for culturing the Suspension feeding larvae: food concentration, temperature and crowding. Large larvae ( Lampetra ( Entosphenus ) tridentata Richardson) were used, weigh-ing from l·5 to 3·0 g (wet). Two food types were employed: suspended yeast cells ( Saccharo-myces cerevisiae , 0–20 mg1 – (dry), or, in a few tests, a fine particulate fish food, Liquifry® (Interpet LTD, 0–13 mg l^-1). At both 14 and 4°C, yeast could sustain weight increases comparable to those in nature: >6% month^-1 for up to 6 months, the duration of the study. In a single lest, a vitamin Supplement failed to improve growth on yeast. Growth-was fastest at 14°C (+41% month^-1, max. weight increase), although also substantial at 4°C (+11% month^-1, max). Growth could not be sustained at 20°C, due perhaps to difficulty in removing products of food decay from the aquaria. Food level being constant, growth rate varied inversely with animal density. It is suggested that larval lampreys release a growth-inhibiting substance into the sand which they inhabit. Overall, the best growth was obtained at 14° C, with <0·05g of animal (wet weight) – aquarium water and average daily yeast concentrations between 4 and 13 mg –. Liquifry was associated with lowered growth rates when present continually above 4 mg (dry weight) – (14° C), although growth did occur at lower concentrations.     

Energy and ration requirements of juvenile Pacific halibut (Hippoglossus stenolepis) based on energy consumption and growth rates

Growth of captive juvenile Pacific halibut was linearly related to energy consumption (J g⁻¹ day⁻¹) at 4°C by the following equation: growth (% body weight (b.w.) day⁻¹)=0–007 (consumption J g⁻¹ day⁻¹)– 0.192; r² =0.81. Weight gain was independent of size for fish between 9 and 7000 g when growth was expressed as a function of consumption in J g⁻¹ day⁻¹. Maintenance ration determined in feeding–growth experiments averaged 27.4 J g⁻¹ day⁻¹ at 4–0°C. Small halibut ate significantly more food than large fish. Single meals following 2 day fasts averaged 4.1% b.w. for halibut under 100 g, 1.72% b.w. for 1.2 kg fish and 1.1% B.W. for 6.8 kg fish. Both large and small size categories of halibut tended to evacuate their meal in about 3 days even though small fish ate relatively larger meals. Minimum estimates for daily ration to achieve growth rates observed in the Gulf of Alaska were approximately 0.5 to 2.4% b.w. day⁻¹ depending on fish size and whether northern shrimp or yellowfin sole were their prey.     

Isotopic evidence for the provenance and turnover of organic carbon by soil microorganisms in the Antarctic dry valleys

The extremely cold and arid Antarctic dry valleys are one of the most environmentally harsh terrestrial ecosystems supporting organisms in which the biogeochemical transformations of carbon are exclusively driven by microorganisms. The natural abundance of ¹³C and ¹⁵N in source organic materials and soils have been examined to obtain evidence for the provenance of the soil organic matter and the C loss as CO₂ during extended incubation (approximately 1200 days at 10°C under moist conditions) has been used to determine the potential decay of soil organic C. The organic matter in soils remote from sources of liquid water or where lacustrine productivity was low had isotope signatures characteristic of endolithic (lichen) sources, whereas at more sheltered and productive sites, the organic matter in the soils that was a mixture mainly lacustrine detritus and moss-derived organic matter. Soil organic C declined by up to 42% during extended incubation under laboratory conditions (equivalent to 50–73 years in the field on a thermal time basis), indicating relatively fast turnover, consistent with previous studies indicating mean residence times for soil organic C in dry valley soils in the range 52–123 years and also with recent inputs of relatively labile source materials.     

Effects of different incubation temperatures on the yolk‐feeding stage of Eupallasella percnurus(Pallas)

Embryos and yolk‐feeding larvae of lake minnow Eupallasella percnurus were reared at 13, 16, 19, 22 and 25° C with no access to external food. Time from egg activation to first embryonic movements, hatching, filling of swimbladder and final yolk resorption increased with decreasing temperature. At 13° C, c . 40% of larvae were unable to fill their swimbladder. The predicted lower temperature at which development and growth ceased (biological zero, t ₀) was the same for both processes, c . 7·5–10·5° C. There was no ontogenetic shift in the t ₀ value. Temperature coefficients for development ( Q _10dev.) ranged from 2 to 3 at 19–25° C, but were higher in hatched larvae at lower temperatures. Eggs of E. percnurus had a combination of small size, high hydration and low caloric value of fresh matter. Dry mass of larval tissue on yolk, percentage of dry matter in wet matter, and specific growth rate were maximized at 22 and 25° C. At 19–25° C, energy and matter contained in the initial eggs were converted to body tissue most efficiently. Temperatures from 22 to 25° C are considered optimal for E. percnurus embryos and yolk‐feeding larvae and are recommended for their indoor rearing.     

Effect of organic loading on nitrification and denitrification in a marine sediment microcosm

Abstract The effects of organic additions on nitrification and dentrification were examined in sediment microcosms. The organic material, heat killed yeast, had a C/N ratio of 7.5 and was added to sieved, homogenized sediments. Four treatments were compared: no addition (control), 30 g dry weight (dw) m⁻² mixed throughout the 10 cm sediment column (30M), 100 g dw m⁻² mixed throughout sediments (100M), and 100 g dw m⁻² mixed into top 1 cm (100S). After the microcosms had been established for 7–11 days, depth of O₂ penetration, sediment-water fluxes and nitrification rates were measured. Nitrification rates were measured using three different techniques: N-serve and acetylene inhibition in intact cores, and nitrification potentials in slurris. Increased organic additions decreased O₂ penetration from 2.7 to 0.2 mm while increasing both O₂ consumption, from 30 to 70 mmol O₂ m⁻² d⁻¹, and NO₃⁻ flux into sediments. Nitrification rates in intact cores were similar for the two methods. Highest rates occurred in the 30M treatment, while the lowest rate was measured in the 100S treatment. Total denitrification rates (estimated from nitrification and nitrate fluxes) increased with increased organic addition, because of the high concentrations of NO₃⁻ (40 μM) in the overlaying water. The ratio of nitrification: denitrification was used as an indication of the importance of nitrification as the NO₃⁻ supply for denitrificaion. This ratio decreased from 1.55 to 0.05 iwth increase organic addition.     

Bioactivity of stored luteinizing hormone-releasing hormone analogue (LHRHa) in sea bass, Lates calcarifer Bloch

The spawning induction activity of dissolved and pelleted (D-Ala⁶, Pro⁹ N ethylamide) luteinizing hormone-releasing hormone analogue (LHRHa) stored for various periods was assessed in mature female sea bass. The spawning response of mature fish was reduced significantly after injection of dissolved LHRHa (20 μg kg⁻¹) stored for more than 90 days in a refrigerator (4–10°C) or for more than 30 days at room temperature (28–30°C). Similar to fish administered fresh preparations of LHRHa, fish spawned successfully after injection of a solution of LHRHa previously frozen, subjected to alternate freezing and thawing, exposed to sunlight or implanted pelleted LHRHa (50 ng kg⁻¹) stored at room temperature for 30–120 days. Loss of hormone bioactivity after prolonged storage may have been due to bacterial growth in solubilized preparations. Injection or implantation of stored LHRHa did not influence egg production among treated sea bass. These results demonstrated the relatively prolonged shelf life of stored LHRHa.     

The effect of temperature and fish size on growth and feed efficiency ratio of juvenile spotted wolffish Anarhichas minor

The growth properties of juvenile spotted wolffish Anarhichas minor reared at 4, 6, 8 and 12° C, and a group reared under 'temperature steps', (T‐step) i.e . with temperature reduced successively from 12 to 9 and 6° C were investigated. Growth rate and feed efficiency ration was significantly influenced by temperature and fish size. Overall growth rate was highest at 6° C (0·68% day⁻¹) and lowest at 12° C (0·48% day⁻¹), while the 4 and 8° C, and the T‐step groups had similar overall growth rates, i.e . 0·59, 0·62 and 0·51% day⁻¹ respectively. Optimal temperature for growth ( T _{opt G} ) and feed efficiency ratio (T_{opt FCE}) decreased as fish size increased, indicating an ontogenetic reduction in T _{opt G} and T _{opt FCE}. The results suggest a T _{opt G} of juvenile spotted wolffish in the size range 135–380 g, dropping from 7·9° C for 130–135 g to 6·6° C for 360–380 g juveniles. The T _{opt FCE} dropped from 7·4° C for 120–150 g to 6·5° C for 300–380 g juveniles. A wider parabolic regression curve between growth, feed efficiency ratio and temperature as fish size increased, may indicate increased temperature tolerance with size. Individual growth rates varied greatly at all time periods within the experimental temperatures, but at the same time significant size rank correlations were maintained and this may indicate stable size hierarchies in juvenile spotted wolffish.     

Thermoregulation and metabolism in the Cape golden mole (Insectivora: Chrysochloris asiatica)

The Cape golden mole, Chrysochloris asiatica is an insectivore which excavates superficial foraging burrows as it searches for its food. It has a mean (±S.D.) resting metabolic rate (RMR) when newly captured of 1–17±0.17 cm³ O₂g^-1 h^-1 ( n = 14), within the thermoneutral zone (TNZ) of 30–32°C.
The body temperature (Tb) of the mole in the TNZ is low 32.9 ± 0.36 ( n = 14) and remains stable at ambient temperatures (Tas) from 28–32°C. Above 32°C (range 34–37°C), Tb increases albeit slightly to 36 ± 1.75°C ( n = 14). The conductance is high 0.27 ± 006cm³ O₂g^-1 h^-l°C^-1 ( n = 46) at the lower limit of thermoneutrality. The mean RMR at 9°C (the lowest Ta tested) was 4.82±11 cm³ O₂g^-1h^-1, which is 4.1 times that of the RMR in the TNZ.
At an ambient temperature of 9°C, three of the golden moles entered a state of torpor where the RMR was reduced from 5.9±0.56 to 10 1.0 ± 0.69cm³O₂g^-1h^-1.     

Lough Ennell: laboratory studies on sediment phosphorus release under varying mixing, aerobic and anaerobic conditions

SUMMARY. 1. The exchange of phosphorus between the epilimnetic (shallow zone) sediment and water column in Lough Ennell was investigated in laboratory experiments using five intact cores.
2. Variations in water mixing, sediment suspension and aerobic–anaerobic oxygen status in the water column and its effects on sediment phosphorus release rates were determined.
3. Experimental results indicated that phosphorus release is possible under both aerobic and anaerobic conditions. Aerobic release (0.025 mg P l⁻¹ over 5 days) was possible up to the point when mass resuspension of sediment occurred. Anaerobic release for the same period and mixing conditions was 0.183 mg Pl⁻¹.
4. The release rate under aerobic conditions at 10°C equates to an internal areal loading of 0.134 g P m⁻² yr^−1, which is approximately 17% and 30% of the average total phosphorus and orthophosphate loadings respectively for the period 1974–79.
5. The results clearly implicate aerobic inorganic phosphorus release from the epilimnetic sediments as a significant source of this nutrient to the overlying water column and is likely a major factor in the continuing eutrophic status in the lake.     

Influence of different rations and water temperatures on the growth rates of shortfinned eels and longfinned eels

Juvenile (12–152 g) shortfinned eels Anguilla australis and longfinned eels A. dieffenbachia caught in New Zealand streams were fed squid mantle Nototodarus spp. 4 days per week in laboratory experiments. A linear multiple regression equation showed the amount of food eaten (0–2·7% w day⁻¹) explained 77·7% of the variation in specific growth rates (–0·60 to +1·07% w day⁻¹) among individual eels, while previous growth rates, water temperature (10·0–20·6°C), and eel weight (12–152 g) explained a further 5·6, 1·4 and 0·8%, respectively. Growth in length ranged from –0·3 to +0·9 mm day⁻¹. Eels which were starved and then given high rations grew substantially faster than expected. Once growth rates were adjusted for differences in ration and other factors, there were no significant differences in growth rates between species or individual fish. Growth of shortfinned eels fed maximum rations of commercial eel food depended on fish size and water temperatures and ceased below 9·0°C. Growth rates in the wild were substantially less than the maximum possible, after seasonal changes in water temperatures were taken into account, indicating that food supplies and not low water temperatures were controlling growth rates in the wild.     

An in situ estimate of daily food consumption and alimentary canal evacuation rates of common carp, Cyprinus carpio L.

Daily food consumption and alimentary canal evacuation rates for common carp, Cyprinus carpio L., were estimated at 4-week intervals from April to October 1979. Both the rate of food evacuation and its relationship to temperature were described by an exponential function. Gut evacuation rates ranged from 6.0% of gut contents per hour at 9°C to 32.7% h⁻¹ at 26.5°C. Temperature accounted for 72–91% of the variation in gut evacuation rate among months. Daily food consumption varied from a low of 3.87 mg g⁻¹ of fish wet wt per day in April at 14°C (0.39% of wet body wt) to a high of 40.75 mg g⁻¹ (4.08%) in August at 26.5°C. Although a greater quantity of food appeared to be consumed during the morning in spring and early summer and a greater amount in the evening after July, there was not a statistically significant time of day for peak feeding activity. The field method described here seemed to provide reasonable estimates of food consumption compared with other field and laboratory studies.     

Simulation of nitrogen assimilation by heterotrophic soil microbial biomass

Assimilation of N by heterotrophic soil microbial biomass is associated with decomposition of organic matter in the soil. The form of N assimilated can be either low molecular weight organic N released from the breakdown of organic matter (direct assimilation), or NH⁺₄ and NO⁻₃ from the soil inorganic N pool, into which mineralized organic N is released (mineralization immobilization turnover). The kinetics of C and N turnover in soil is quantifiable by means of computer simulation models. NCSOIL was constructed to represent the two assimilation schemes. The rate of N assimilation depends on the rate of C assimilation and microbial C/N ratio, thereby rendering it independent of the assimilation scheme. However, if any of the N forms is labeled, a different amount of labeled N assimilation will be simulated by the different schemes. Experimental data on inorganic N and ¹⁵N and on organic ¹⁵N dynamics in soils incubated with ¹⁵N added as NH⁺₄ or organic N were compared with data simulated by different model schemes. Direct assimilation could not account for the amount of ¹⁵N assimilated in any of the experimental treatments. The best fit of the model to experimental data was obtained for the mineralization immobilization turnover scheme when both NH⁺₄ and NO⁻₃ were assimilated, in proportion to their concentration in the soil.     

Temperature-induced changes of early development and yolk utilization in the African catfish Clarias gariepinus

Artificially fertilized eggs and yolk-sac larvae of a freshwater tropical/subtropical fish Clarias gariepinus receiving no external food were incubated at 22, 25 and 28° C until full yolk resorption. Developmental time, size and matter composition (CHNS-O Analyzer and ashing) were assessed at egg fertilization, hatching and yolk resorption; respiration was measured every 4–5 h. The course of acceleration of C. gariepinus embryonic developmental rate with temperature ( Q_{10 dev}) was compared over the temperature range to those of Cyprinus carpio and Oncorhynchus mykiss ; they differed greatly, but were similar when compared on the basis of effective temperatures specific to each fish. Specific growth rates for energy (88, 150 and 183% per day at 22,25 and 28° C, respectively) as well as the conversion efficiencies of egg energy (64, 71 and 68%, respectively) and protein (71, 78 and 76%, respectively) in C. gariepinus larval tissues were higher than those known for the endogenous feeding period of coldwater and temperate fish species. In C. gariepinus at the end of yolk resorption, the carbon percentage and caloric values of dry weight, size (in terms of dry matter, minerals, protein and energy per larva) and transformation efficiencies were lowest at 22° C, highest at 25° C and had slightly decreased at 28° C. A tentative mechanism which leads to the positive or negative response of body size to temperature over the viable temperature range is defined.     

Mineralization budgets in sediment microcosms: Effect of the infauna and anoxic conditions

Abstract A number of sediment incubations were set up to reproduce some of the conditions used by Kristensen and Blackburn [1] and to make a comparison with their results. There were three types of microcosm: aerobic (OX), anaerobic (AN) and aerobic with Nephtys (NOX). In addition to other measurements, dissolved organic nitrogen (DON) pools and fluxes, were measured. The sediment in this experiment contained more particulate organic matter (POM). Nephtys (NOX) had the same effect as Nereis in increasing the rate of mineralization of POC and PON, compared with the OX-cores (2.1 and 2.6 times, respectively). Again, the AN-cores had a higher mineralization rate (loss of POM) than that of the OX-cores, but in addition, mineralization in NOX-cores was not significantly different from AN-cores. It was thus confirmed that anoxic mineralization could be as high, or higher, than the oxic process. Both the temporal patterns of O₂-and and CO₂-fluxes and their magnitudes were very similar to those reported earlier. This contrasts with the higher loss of POM in the present experiment. However, the loss of C in DOC (associated with the measured DON) can account for the extra POM loss. The pore-water profiles of σCO₂ and NH₄⁺ were similar to those in the earlier report, and the fluxes of σCO₂, O₂, NH₄⁺ and NO₃⁻ followed the same temporal pattern.     

Growth, diet and metabolism of common wolf–fish in the North Sea, a fast–growing population

The von Bertalanffy growth parameters for common wolf–fish Anarhichas lupus in the North Sea were: male: L ∞=111·2 cm, t ₀=–0·43 and K =0·12; and female: L ∞=115·1 cm, t ₀=–0·39 and K =0·11, making this the fastest growing stock reported. Resting metabolic rates (RMR±S.E.) and maximum metabolic rates (MMR±S.E.) for six adult common wolf–fish (mean weight, 1·39 kg) at 5° C were 12·18±1·6 mg O₂ kg^–1 h^–1 and 70·65±7·63 mg O₂ kg^–1 h^–1 respectively, and at 10° C were 25·43±1·31 mg O₂ kg^–1 h^–1 and 113·84±16·26 mg O₂ kg^–1 h^–1. Absolute metabolic scope was 53% greater at 10° C than at 5° C. The diet was dominated by Decapoda (39% overall by relative occurrence), Bivalvia (20%) and Gastropoda (12%). Sea urchins, typically of low energy value, occupied only 7% of the diet. The fast growth probably resulted from summer temperatures approximating to the optimum for food processing and growth, but may have been influenced by diet, and reduced competition following high fishing intensity.     

Seasonal changes in vertical distribution, biomass and faecal production of tubificids in the profundal region of a shallow Japanese lake

Two tubificid species Limnodrilus hoffmeisteri and L. claparedeianus formed more than 93% of the total number of oligochaetes in the profundal. Limnodrilus spp. worms were found down to 33 cm in the sediment but in great numbers in the upper zone in June and October. Worms confined to the top 15 cm of sediment accounted for 53-92% of the total number. There were two annual maxima in population density and biomass, one in late spring (66000 inds m⁻², 17 g wet wt m⁻²) and the other in mid autumn (97000 inds m⁻², 176 g wet wt m⁻²). Two regression lines describing the effect of temperature on faecal production rate were obtained; Log F = 0.0604 T (°C) −0.7660 (below 15°C), Log F = 0.0266 T – 0.2170 (above 15°C). In total 26.8 kg dry wt m⁻² of sediment was defecated annually by Limnodrilus spp. The sediment in the 0–10 cm stratum may pass through the guts of the worms 2.3 times a year. Sedimentation rates in profundal region were very low with respect to the faecal production rates of the tubificids.     

A new type of metabolism chamber for the determination of active and postprandial metabolism offish, and consideration of results for coregonid and salmon juveniles

The optomotor reaction of juvenile Coregonus schinzipalea Val. et Cuv. and Salmo salar L. was utilized to develop a circular tube metabolism chamber to measure oxygen consumption and ammonia excretion as a function of swimming speed. The metabolism chamber with a constant water flow assured the maintenance of stable conditions. The unidirectional movement of fish was measured in a circular tube with a single narrowing. The relationships between the swimming speed and oxygen consumption or ammonia excretion described by exponential equations allowed the extrapolation towards the standard metabolism, i.e., zero swimming speed. For a juvenile coregonid (0.1–0.15 g individual weight, 2.6–2.8 cm total length) standard metabolism at 14° C was estimated as 0.65 mg0₂ g⁻¹ h⁻¹ and 17.3 μg N(NH₃)g⁻¹ h⁻¹, whereas for juvenile salmon (136mg individual weight) respective values at 22° C were 0.047mg0₂g⁻¹h⁻¹ and 0.61 μg N(NH₃)g⁻¹ h⁻¹. The feeding test with juvenile salmon was also performed in this circular chamber, and in both energy and nitrogen budgets after a meal the partitioning could be precisely attributed to standard metabolism, active metabolism and specific dynamic action (in the case of oxygen consumption) or postprandial nitrogen increase.
The new metabolism chamber allowed the relationship between metabolism and swimming velocity of juvenile fish with developed rheotactic response. It could be used with adult fish for similar purposes.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

A reaction diffusion model of C-N-S-O species in some arctic sediments

Abstract A reaction diffusion model was used to simulate the mineralization processes in an Arctic sediment. The simulation and the actual sediment were compared in relation to profiles of O₂, NO₃ and NH₄⁺. The site of particulate organic matter (POM) degradation was the single most important factor in fitting the simulation profiles to those of the sediment. It was deduced that most POM degradation occurred close to the sediment surface. When a reasonably good simulation had been obtained, the sensitivity of the model to changes in other parameters was investigated. Increases in POM degradation in the upper sediment resulted in increases in concentration of NH₄⁺ and NO₃⁻, but further increases in POM degradation created anoxic conditions below 3 mm, resulting in decreases in NO₃⁻ concentrations. The model was relatively intensive to changes in POM degradation in the lower sediment layers; increases led to more anoxic conditions and to less NO₃⁻. Increases in the C/N ratio of the POM in the lower sediment layers had little effect; increases in C/N in the upper layers led to a decrease in NH₄⁺ and NO₃⁻. The model was sensitive to changes in the first order rate constant for nitrification, but not for denitrification. Decreases in the K _m for O₂ of the nitrifying bacteria had no effect on the profiles.