Sample collection and examination
Water samples were collected for six months in both rainy (March–May 2020) and the dry season (June–August 2020). Samples from the inflow and the outflow rivers were collected from Rivers Kagoma and Heissesero (Fig. 1), respectively. Samples were collected into 1L plastic sampling bottles between 9:00 and 11:00 am prepared and then transported in an icebox with ice to the National Water and Sewerage Corporation (NWSC) Central Laboratories in Kampala for analysis within 24 h. While in the laboratory, samples were stored in the refrigerator at 4 °C before analysis. Water temperature, DO levels and pH were measured on-site during sampling. DO and water temperature were measured using the DO meter (DO 5510 M.R.C model). The pH was measured using a water-resistant hand-held pH meter (HI8314 HANNA instruments) following APHA (2017) standards. In the laboratory, samples were analysed for the determination of NH3-N, NO3-N and Org-N following the APHA (2017) standard procedures as described by Saturday et al. (2021). Besides, climatic secondary data such as air temperatures, sunshine, relative humidity, rainfall for a period of six months (March–May 2020) and June–August 2020 were collected from Uganda National Meteorological Authority, Kabale centre. The water inflow into the lake through the Kagoma River and outflow through the Heissesero River were measured onsite using the float-area method as described by Dahal and Dorji (2019).
Model development
Conceptual model of nutrient balance
A conceptual schematic model that encompasses the forcing variables, state variables and the activities governing nitrogen transformation processes is used (Fig. 2). The forcing variables considered in this model are the volume of water in the lake (Q), water temperature (T), pH, volume of water inflow (in) and outflow (out), and DO. The state variables are the major forms of nitrogen; Organic Nitrogen (Org-N), Ammonia Nitrogen (NH3-N) and Nitrate Nitrogen (NO3-N) as indicated by boxes connected by respective processes. The nitrogen transformation considered includes nitrification, denitrification, volatilization, mineralization, microbes’ ammonia and nitrates uptake, algae uptake, sedimentation and decaying processes. A complete materials balance includes lake inflow and outflow, nutrients contained in the inflow and outflow of the lake, precipitation, evaporation, solar radiations, and air temperature; all of which influence nitrogen dynamic processes in the lake system are considered in the model. The nutrient mass balance around state variables was done based on the simplified Eq. (1) (Jørgensen and Bendoricchio 2001).
$${\text{Accumulation }} = {\text{Input}} - {\text{Output}} \pm {\text{Reaction}}$$
(1)
Here, Input = Nutrient load that enter the lake from diverse sources and through different ways; Output = Nutrient concentration that leaves the lake through different ways; Reaction = the way nutrients leave the lake system by chemical transformation into other substances.
As illustrated in the conceptual model diagram (Fig. 2), nitrogen enters the Lake Bunyonyi system majorly by inflow stream and runoff from agricultural land use activities. Besides nutrient inflow via stream discharges and agricultural runoff, nutrients enter into the lake system through direct rainfall on the lake. While in the lake, nutrients can be taken up by aquatic plants, animals and microbes but are released back into the lake system through excretion and decomposition after death. This precisely means that nutrients are temporarily stored in the biomass. It is generally expected that nutrient fixation is larger than release because organic matter cannot all be decomposed and therefore some nutrients remain fixed in organic matter. Besides, fish catches and water abstraction remove nutrients from the lake and some nutrients are adsorbed to the sediment and may be released when the sediments are suspended by waves.
The following assumptions were considered for the development of the Model.
-
i.
The inlet of water to the lake considered in the model is Kagoma River, while the other non-sources like runoff and precipitation are not included in the Model.
-
ii.
The outlet which has been considered in this model is the Heissesero River; other water loss ways like underground seepage and evaporation are not included in the model.
-
iii.
Similar microorganisms within the lake take up both NO3-N and NH3-N.
-
iv.
The rivers Kagoma (inlet) and Heissesero (outlet) are not seasonal; they continue to bring water into the lake and drain water from the lake throughout the year, respectively.
Based on these assumptions, the following equations for the mass balance of state variables and nitrogen transformation processes were derived.
Ammonia–nitrogen
Based on the mass balance, NH3-N was added in the lake through Kagoma River, and within the lake by mineralization, regeneration from the sediments, while, volatilization, nitrification, uptake by plants and microorganism processes were reducing NH3-N concentration within the lake and some were drained from the lake by Heissesero River as shown in Eq. (2).
$$\frac{{\partial \left[ {{\text{NH}}_{3} - {\text{N}}} \right]}}{\partial t} = \left( {\frac{{\partial Q_{IF} }}{\partial t} \frac{{\partial \left[ {{\text{NH}}_{3} - {\text{N}}_{{{\text{in}}}} } \right]}}{\partial A}} \right) + \left( {r_{m} + r_{{{\text{reg}}}} } \right) - \left( {\frac{{\partial Q_{OF} }}{\partial t} \frac{{\partial \left[ { {\text{NH}}_{3} - {\text{N}}_{{{\text{out}}}} } \right]}}{\partial A} + r_{g1} + r_{v} + r_{n} } \right).$$
(2)
where, \(\frac{{\partial Q_{IF} }}{\partial t}\). = Water inflow rate (m3/d), \(\frac{{\partial Q_{OF} }}{\partial t}\) = Water outflow rate (m3/d), \(\frac{{\partial \left[ {{\text{NH}}_{3} - {\text{N}}_{{{\text{in}}}} } \right]}}{\partial A}\) = Ammonia–nitrogen loading in water inflow (g/d m−2), \(r_{m}\) = Rate of mineralization of organic nitrogen (g/d m−2), rreg = Rate of regeneration (g/d m−2), \(\frac{{\partial \left[ {{\text{NH}}_{3} - {\text{N}}_{{{\text{out}}}} } \right]}}{\partial A}\). = Ammonia–nitrogen loading in water outflow (g/d m−2), \(r_{g1}\). = Ammonia uptake by microorganisms, \(r_{v}\) = Rate of volatilization (g/d m−2), \(r_{u1 }\). = Uptake by the plant (g/d m−2), rn = Rate of nitrification of ammonia (g/d m−2).
Nitrate–nitrogen
Based on the mass balance, NO3-N was added to the lake through the Kagoma River. The nitrification, denitrification, and nitrate uptake by microorganisms and algae were the major processes reducing NO3-N concentration within the lake and some were dined through Heissesero River as shown in Eq. (3).
$$\frac{{\partial \left[ {{\text{NO}}_{3} - {\text{N}}} \right]}}{\partial t} = \left( {\frac{{\partial Q_{IF} }}{\partial t} \frac{{\partial \left[ {{\text{NO}}_{3} - {\text{N}}_{{{\text{in}}}} } \right]}}{\partial A} + r_{n} } \right)\left( {\frac{{\partial Q_{OF} }}{\partial t}\frac{{\partial \left[ {{\text{NO}}_{3} - {\text{N}}_{{{\text{out}}}} } \right]}}{\partial A} + r_{g3} + r_{dm} } \right)$$
(3)
\(\frac{{\partial Q_{{{\text{in}}}} }}{\partial t}\). = Water inflow rate (m3/d), \(\frac{{\partial Q_{{{\text{out}}}} }}{\partial t}\) = Water outflow (m3/sec), \(\frac{{\partial \left[ {{\text{NO}}_{3} - {\text{N}}_{IF} } \right]}}{\partial A} + r_{n}\). = Nitrate-nitrogen inflow (g/d m−2), rg2 = Rate of nitrate uptake by microorganisms, \(\frac{{\partial \left[ {{\text{NO}}_{3} - {\text{N}}_{OF} } \right]}}{\partial A}\) = Nitrate–nitrogen outflow (g/d m−2), \(r_{n}\) = Rate of nitrification (g/d m−2), \(r_{dn}\) = Rate of denitrification (g/d m−2).
Organic-nitrogen
Like NH3-N and NO3-N, Org-N was added into the lake through Kagoma River and within it by decomposition. On the other hand, mineralization, nitrification, denitrification, and nitrate uptake by microorganisms and algae were responsible for Org-N reduction within the lake and some were drained off it via Heissesero River as shown in Eq. (4).
$$\frac{{\partial \left[ {{\text{Org}} - N} \right]}}{\partial t} = \left( {\frac{{\partial Q_{{{\text{in}}}} }}{\partial t} \frac{{\partial \left[ {{\text{Org}} - N_{{{\text{in}}}} } \right]}}{\partial A}} \right) + \left( {r_{g1} + r_{g2} } \right) - \left( {\frac{{Q_{{{\text{out}}}} }}{\partial t} \frac{{\partial \left[ {{\text{Org}} - N_{{{\text{out}}}} } \right]}}{\partial A} + r_{m} + r_{s} } \right)$$
(4)
where, \(\frac{{\partial Q_{{{\text{in}}}} }}{\partial t}\) = Water inflow rate (m3/d), \(\frac{{\partial Q_{{{\text{out}}}} }}{\partial t}\) = Water outflow (m3/d), \(\frac{{\partial \left[ {{\text{Org}} - N_{{{\text{in}}}} } \right]}}{\partial A}\) = Organic-nitrogen inflow (g/d m−2), \(\frac{{\partial \left[ {{\text{Org}} - N_{{{\text{out}}}} } \right]}}{\partial A}\) = Organic-nitrogen outflow (g/d m−2), \(r_{g1}\) = Rate of ammonia uptake by microorganisms, \(r_{g2}\) = Rate of nitrate uptake by microorganisms (g/d m−2), \(r_{m}\) = Rate of mineralization (g/m2/day), \(r_{s}\) = Rate of sedimentation (g/d m−2).
Equations for nutrient dynamic processes
Mineralization
The mass balance for mineralization of organic nitrogen in the lake system was modelled using first-order kinetics as presented in Eq. (5).
$$\frac{\partial m}{{\partial t}} = r_{m} = \left[ {{\text{Org}} - N } \right] \times K$$
(5)
where, K = Mineralization rate of constant nitrogen (day−1).
Nitrification
According to Fritz et al. (1979), the nitrification process model is based on the assumption that nitrite formation by Nitrosomonas is a rate-limiting step and is inhibited by dissolved oxygen, temperature, and pH. The nitrification process was modelled using Eq. (6). The temperature and pH influence on Nitrosomonas bacteria were explained by the empirical relationship presented by Eqs. (7) and (8), respectively.
$$r_{n} = \left[ {\left( {\frac{{u_{n} }}{{Y_{n} }}} \right) \times \left( {\frac{{{\text{NH}}_{3} - {\text{N}}}}{{K_{1} + {\text{NH}}_{3} - {\text{N}}}}} \right) \times \left( {\frac{{{\text{DO}}}}{{K_{2} + {\text{DO}}}}} \right) \times \left( {C_{T} } \right) \times \left( {C_{{{\text{pH}}}} } \right)} \right] \times \left[ {{\text{Org}} - {\text{N}} } \right]$$
(6)
where, \(\mu_{n}\) = maximum Nitrosomonas growth rate (d−1), \(Y_{n}\) = Yield coefficient for Nitrosomonas bacteria (mg VSS/mg N), \(K_{1}\) = Ammonia Nitrosomonas half-saturation constant (g/d m−2), \(K_{2}\) = Oxygen Nitrosomonas half-saturation constant (g/d m.−2), \(C_{T }\) = Temperature-dependent factor, \(C_{{{\text{pH}}}}\) = Nitrosomonas growth-limiting factor for pH
$$C_{T} = {\text{exp}}\left[ {0.0098\left( {T - T_{O} } \right)} \right]$$
(7)
where, \(C_{T}\) = Temperature dependence factor, \(T\) = Temperature in °C, \(T_{O}\) = Reference temperature (°C) = 15 °C
$$C_{{{\text{pH}}}} = \left\{ {\begin{array}{*{20}l} {1 - 0.83\left( {7.2 - {\text{pH}}} \right) \ldots } \hfill & {{\text{for}}} \hfill & {{\text{pH}} \ldots < 7.2} \hfill \\ {10 \ldots \ldots \ldots \ldots \ldots \ldots \ldots .} \hfill & {{\text{for}} \ldots } \hfill & {{\text{pH}} \ge 7.2} \hfill \\ \end{array} } \right..$$
(8)
Denitrification
To model the rate of denitrification, a combination of denitrifying bacterial activities is applied. The model expression is presented by Eq. (9).
$$r_{dm} = \left\lfloor {(DR_{ - 20 \times } \theta^{T - 20} } \right\rfloor \times \left[ {{\text{NO}}_{3 - } {\text{N}}} \right]$$
(9)
where, \(DR_{ - 20}\) = Denitrification rate constant at 20 °C (d−1), \(\theta\) = Arrhenius constant microorganism growth temperature coefficient.
Ammonia uptake by microorganisms
In the model, it is assumed that ammonium uptake, i.e., MO (NH3-N) would take place as long as is available in the lake since autotrophic bacteria prefer it to nitrates. Equation 10 is used to model NH3-N uptake by microorganisms in the water phase.
$${\text{MO}} \left( {{\text{NH}}_{{3 - {\text{N}}}} } \right) {\text{Uptake}} = \left[ {\left( {u_{\max - 20} } \right){\text{Thita}}^{T - 20} \times \left( {\frac{{{\text{NH}}_{{3 - {\text{N}}}} }}{{K_{3} + {\text{NH}}_{{3 - {\text{N}}}} }}} \right)} \right] \times \left[ {\frac{{{\text{DO}}}}{{K_{2} + {\text{DO}}}}} \right] \times C_{{{\text{pH}}}} \times {\text{NH}}_{{3 - {\text{N}}}} \times P_{1}$$
(10)
where, \(\mu_{{{\text{max}} - 20 }}\) = Maximum growth rate of bacteria at 20 °C (d−1), \(P_{1}\) = Ammonia uptake preference factor, \(K_{3}\) = Ammonia uptake half-saturation constant (g/m3), \(Thita\) = Microorganisms growth temperature coefficient.
Nitrate uptake rate by microorganisms
The nitrate uptake is done by autotrophic bacteria but it is assumed that it takes place after all the NH3-N has been consumed and hence depleted from the system. Equation (11) is used to model the rate of nitrate uptake by microorganisms.
$${\text{MO}} \left( {{\text{NO}}3 - {\text{N}}} \right){\text{ Uptake }} = \left[ {\left( {U_{\max - 20} } \right) \times {\text{Thita}}^{T - 20} \times \left( {\frac{{{\text{NO}}_{3 - N} }}{{K_{4} + {\text{NO}}_{3 - N} }}} \right)} \right] \times \left[ {\frac{{{\text{DO}}}}{{K_{2} + {\text{DO}}}}} \right] \times C_{{{\text{pH}}}} \times P1 \times {\text{NO}}3 - {\text{N}}$$
(11)
\(\mu_{{{\text{max}} - 20 }}\) = Maximum growth rate of bacteria at 20 °C (d−1), \(P_{1}\) = Nitrate uptake preference factor, \(K_{4}\) = Nitrate uptake half-saturation constant (g/m3), \(Thita\) = Microorganisms growth temperature coefficient.
Sedimentation rate
The transformation processes considered in the modelling of nitrogen in sediments in the lake system are sedimentation and ammonia–nitrogen regeneration. The organic nitrogen in sediments is mineralized by microorganisms, which ultimately regenerate NH3-N to the water column. It is assumed that the rate of regeneration (rr) follows the first-order kinetics presented by Eqs. (12) and (13).
$$\frac{\partial S}{{\partial t}} = r_{s} - r_{r}$$
(12)
where, \(r_{s} =\) Settling rate of organic-nitrogen to the bed (g/d m−2), \(r_{r}\) = rate of ammonia regeneration (g/d m.−2)
$$r_{r} = r_{{{\text{reg}}}} \times N_{{{\text{sediments}}}}$$
(13)
where, \(r_{reg }\). = Regeneration rate constant of ammonia (g/d m−2).
Ammonia volatilization
To model the rate of ammonia volatilization, the mass balance Eq. (14) is used.
$$r_{v} = \frac{{\left[ {NH_{3} - N} \right]}}{h} \times KL$$
(14)
where, h = Water depth in m, T = Temperature in °C, KL = Mass transfer coefficient = 0.056 exp (0.13 (T−20).
Model calibration
To run this model, the STELLA software (version 9.0.1) was used. The software has the capabilities to simulate the behaviour of nitrogen transformation in the Lake Bunyonyi system using conservation of mass principles. The mathematical equations presented in subsection 7.5.1 for the mass balance of ammonia–nitrogen, nitrate-nitrogen and organic-nitrogen along with forcing functions were entered in the STELLA software. Data collected from samples obtained from Lake Bunyonyi were used as inputs for the model calibration. These data include monthly averages for NH3-N, NO3-N, Org-N, DO, temperature, and pH (of the le system, inflows and outflows), rainfall, solar radiation, water inflows and outflows.
Nutrient mass balance
Total nutrient inflow was computed as the sum of the total volume of water that flows into the lake system multiplied by the concentration of nutrients in water inflows (Eq. 15). Total nutrient outflow was computed as a sum of the total volume of water outflow multiplied by the concentration of nutrients in the water outflows (Eq. 16).
$$Q_{{{\text{in}}}} C_{{{\text{in}}}} = Q_{{{\text{STMS}}}} C_{{{\text{STMS}}}}$$
(15)
$$Q_{{{\text{out}}}} C_{{{\text{out}}}} = Q_{{{\text{STM}}\left( {{\text{out}}} \right)}} C_{{{\text{STM}}\left( {{\text{out}}} \right)}}$$
(16)
To obtain the best values of coefficients, model calibration was performed. The model efficiency criterion by Nash and Sutcliffe (1970) was used to establish the efficiency of the model.