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M A T H E M A T I C A L M O D E L L I N G OF A F I S H P O N D E C O S Y S T E M
Yu.M. SVIREZHEV, V.P. KRYSANOVA and A.A. VOINOV
Computer Centre of the U.S.S.R. Academy of Sciences, Moscow (U.S.S.R.)
(Accepted for publication 19 October 1983)
ABSTRACT Svirezhev, Yu.M., Krysanova, V.P. and Voinov, A.A., 1984. Mathematical modelling of a fish pond ecosystem. Ecol. Modelling, 21: 315-337. A mathematical model is constructed for a fish breeding pond for carp, silver carp and bighead. The model is a system of ordinary differential equations describing the material transformations in the ecosystem. It allows a choice of optimal regimes of the aeration, feeding and fertilization of a pond for different climatic conditions in order to maximize the yield. 1. INTRODUCTION
Fish ponds have long been used by man to meet his alimentary and--first of all--protein requirements. Fish breeding was highly developed in ancient China. A high efficiency is achieved only with optimum values of the control parameters, such as input of fodder and fertilizers, and re-aeration of the water body, and with an optimum choice of the seed piece characteristics. The management affects the entire fish pond ecosystem, resulting in unpredictable and, even more, not always desirable changes in the ecodynamics of the reservoir. A mathematical model permits an assessment to be made of the consequences of different control strategies and an estimation of all possible transformations in the whole complex of cause-effect relations in the ecosystem. Attempts to model a fish pond have been made by Vinberg and Anisimov (1966), Borshev (1977), and others. However, the energy approach used by Vinberg and his successors, although providing an adequate qualitative ecosystem description, gives little insight into the management of an ecosystem. Therefore our model was based on the models of lake ecosystems (Jorgensen, 1980; Voinov et al., 1981). This paper presents a fish pond simulation model. The food chain structure and the set of model phase variables are fixed. It is assumed a
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316 priori that the ecosystem in question already includes the biological species and that a management mode is envisaged that can result in the optimum yield. The model was identified for literature data and gives a quite reasonable response to changes in climatic factors and control parameters, thereby serving as a simulator for the analysis of all kinds of possible developments in a fish pond ecosystem. Suitable optimization techniques, derived on the basis of this model, allow one to define optimum control strategies for an ecosystem. On the other hand, fish ponds may serve as useful models for the analysis of ecosystem properties in general, due to their relatively simple trophic structure and the high intensity of the biotic material and energy transformations. From this point of view the mathematical modelling of fish ponds is of some general ecological and theoretical significance. 2. MATERIALCYCLES IN A FISH POND When analyzing a concrete problem, it is necessary to choose a degree of generality that is ample for meeting the purposes of modelling. In our case, to model an optimum fish pond, we have to choose variables that would fully reflect the specificity of fish ponds giving stable high yields over long periods of time. From experience, the joint breeding of carp and herbivorous fish (Bighead, Silver Carp, White Amur, etc.) is very effective. They complement each other well enough, as they occupy almost non-overlapping ecological niches. Although these species may compete for food, they prefer different natural feeds: benthos for Carp, phytoplankton for Silver Carp, zooplankton for Bighead, and macrophytes for White Amur. The last species has not been included into the model due to its relative independence from the other ecosystem components. In describing the natural enrichment of the fodder supplies, it will be logical to take into account the concentrations of the two most usually limiting nutrients, i.e. nitrogen and phosphorus. They are supplemented by the bacterial destruction of dead organics--detritus--and also from the input of artificial fertilizers. Finally, the dissolved oxygen concentration is quite an-important, and sometimes determining factor of the fish pond ecosystem. Hence, the model includes the following phase variables: phytoplankton (F), zooplankton (Z), benthos (B), Carp (C), Bighead (H), Silver Carp (S), dissolved mineral phosphorus (P), dissolved inorganic nitrogen (N), dissolved oxygen (O), artificial fodder (A), detritus combined with bacteria(D). It is assumed that a concrete ecosystem can roughly be described by substituting the complex multi-species community structure with
. etc.). In this case a separate block (for instance.g. The interaction between the phase variables is described according to the scheme of the material cycle presented in Fig. it is regarded as a special /• ELEMENTS(P.317 a simplified block pattern. Such a substitution may be considered correct if all the species within one block have close values of their prime ecological parameters (maximum growth rates. Since the dissolved oxygen (DO) has a controlling. new variables can be added into the model (e. certain variables can be disaggregated (e. 1. Later. macrophytes.g. BIGHEAD ~ PHYTOPLANKTON CARP II ZOOPLANKTON BOTTOM FAUNA II DETRITUS + BACTERIA Fig. respiration coefficients.N)~ BIOGENIC ~I MINERAL FERTILIZERS FORAGE I~ SILVER CARP • . Material cycle in the fish pond. F or Z) may contain dozens of species. It is assumed that such a scheme comprehensively reflects the material transformation processes in the pond. B. F. D). 1. other fish species). Z. regulating effect upon different chemical and ecological processes.
is determined by the intensity of the solar radiation. Inflows and outflows of oxygenin the fish pond. taking into account that the intensity of photosynthesis. For instance. unlike the latter. Figure 2 shows the consumption and replenishment of DO in the ecosystem.. the main source of DO in the ecosystem. 2. Therefore. should be measured in hours rather than in days. i.e. It should be noted that the specific times inherent in the main DO transformations are shorter than the times of other ecosystem processes and. This is quite natural. many scientists stress that fish kills in ponds are most common in the morning.?. variable. ~ DETRITUS n Fig. When modelling a shallow pond with depths of about 1 m and a small area (less than 1 ha) we may neglect the effects of the spatial distribution of . the DO concentration is analysed separately in the model with its own time step.318 reaeration photosynthesi~ OXYGEN r. the DO concentration definitely depends upon the hour of the day.
KNF half-saturation constant for nitrogen uptake. and m = stoichiometric ratio. a point model. 6 x FT(1) = exp ( .e. i. [D]) [N]" )X[F] K~FT[-P]"' m K~VF+[N]n = where g ~ ' = phytoplankton maximum growth rate. accounting for the effect of external factors. 3): exp ( . All the variables are regarded as concentrations and the unit of measurement is mg/1. The external forcing functions in the model are the climatic factors--water temperatures and the total solar radiation--as well as the control elements. i. 02(1) . The relationship between the phytoplankton growth rate and temperature is described by the modified Lehman function (Jorgensen. BASIC MODEL EQUATIONS (a) Nutrients uptake by phytoplankton The phytoplankton growth is an important process. The cycles of these elements are closely related in the ecosystem. the N / P ratio in the living organic matter.e. r< tO(l) T>~ t O ( l ) (T-TO(l)) 4) . 1980) (Fig. K p F = half-saturation constant for phosphorus uptake. The uptake rates of the two elements may be viewed as synchronised according to the stoichiometric ratio. we can represent the phytoplankton growth rate as follows: = t~T"x × FT(1) x xmin( [P]" FF(L. Thus. By the concentration of living organisms we mean the ratio of their total biomass to the volume of the whole reservoir.4 . The nutrients limiting the phytoplankton growth in a fish pond may be nitrogen or phosphorus. the square brackets [ ] will stand for the concentration of the respective ecosystem variables. as we are constructing a local. 6 x (TO(i)-r)') Fff . For different estimates this ratio can be taken from N / P = 10/1 to N / P = 5/1. which depends on the presence of nutrients in the water and also on the external factors such as temperature (T) and illumination (L).4 . The phytoplankton growth is limited according to the stoichiometric ratio. 1 [ F ] . 3. Further. such as the input of artificial feed. The fish-breeding pond is modelled for five months (from April 15 to September 15). mineral fertilizers and the intensity of artificial water aeration.319 organisms and material.
we represent the light limitation function for the FF 1 o ' 2 o b o ' 4o'oo' ~'oo LIGHT ~5oo'Io6oo Fig. o 0 10 20 TEMP 30 \ 4b Fig.320 FT 1. is the difference between the optimum and minimum temperatures. Following Steele (1962). Q I ( 1 ) = To~t . Temperature limitation function for growth.T1. where TO(l) is the optimum temperature for the phytoplankton development.Tolpt is the difference between maximum 1 and optimum temperatures. . Light limitation function for phytoplankton growth. 4. Q2(1)= Tm'~x. 3.
2 or 3 approximate the experimental results with practically the same accuracy. In our model n = 2. The uptake of nutrients is presented by s-shaped trophic functions: V([X]) = [X]" K" + [ XI " where [ X ] = is the substrate concentration. KD = the shading parameter for suspended detritus. which is very important for the stability of the computer realisation of the model. exp(1 - L ) where L = L 0 e x p ( . The latter depends on the concentrations of phytoplankton and detritus in the water: k=KW+ KFX[F I+KDX[DIXKPD where K W = the light extinction coefficient for the water.321 phytoplankton growth as (Fig. and KPD = a fraction of detritus suspended in water. KF = the phytoplankton self-shading parameter. k.k × h) is the illumination at a depth of h calculated by the Bur-Lambert empirical formula in terms of the total solar radiation L 0 and the extinction coefficient.[F] ' [DI)=_L Lop. Note that the experimental data cannot provide an objective criterion for the choice of n: near zero the s-shaped functions with n = 1. V 2~ J n----3 n=2 I ---- 0 10 20 30 40 X 5b Fig. 4): FF(L. 5. 5). n = a dimensionless quantity characterizing the steepness of the function (see Fig. . Trophic functions. In this case the function vanishes with zero derivative.
and n -. [ F ] ) X [ Z ] . feeding with switching.~'.e. Oxygen limitation function for growth. KFZ. the  value in which FO(2) = 1/2. The trophic function is determined by an s-shaped curve: /zrm~ [ F ] " xV(ZF .2. g v z . for instance. 6) accounts for the relationship between the zooplankton growth and the presence of DO in the water: FO(2) -1 1 + exp(-X(2) ×([01 . 6. ~ ~ ~ ~ 02 6 ~ & ~ ~b Fig. The temperature function FT(2) is similar to FT(1). X(2) is the parameter characterizing the steepness of the curve.m ( 2 ) ) ) where m(2) is the oxygen half-maintenance coefficient. The FO(2) function of the logistic type (Fig. i. Carp. [ F ] ) + where /~vmz is the zooplankton maximum growth rate when grazing phyto~' plankton. .. prefers 1 FO "' o .322 (b) Feeding Let us describe the feeding of higher trophic level organisms on organisms of lower levels with an example of zooplankton grazing on phytoplankton: qFz = FT(2) X FO(2) X V(~t~. (e) Feeding with switching It follows from the literature that some fish are characterized by the selectivity of feeding.e. Kvz is the half-saturation constant for the phytoplankton uptake by zooplankton. i.
The feed items are listed according to the preference. where qi is the concentration of the ith type prey. 7.. . )~. x i. However. The interpretation of the experimental data is impeded by the fact that model studies usually incorporate notions that give insight into the essence of the process.323 to feed on benthic organisms.. have an inverted s-shaped form (Fig. Thus the inverted s-shaped function in the model is used to characterize the feeding of carp with switching: ~/([B]. Their choice is arbitrary. lim x~(Q) = 0. will be more appropriate. Then the probability x a = 1 for any set q = (qa. .i=aqi. qa is the favourite feed. the functions x2(ql). the dependence of x i on Q with large Q values seems doubtful. Tan has found a relationship between the probablility x i of the uptake of the ith type feed item and the total concentration of feed Q . Nevertheless. q2. etc. rnB) = exp(-X. to a great extent.~ This means that the transient regime is not clearly defined but covers an interval of Q-values. in the case of a deficit in benthos. q2). U p t a k e probabilities. 7).-. Tan who analyzed the experimental data of Ivlev (1955) on the relationship between feeding and concentration of feed items for carp. it can switch to feeding on zooplankton. . with x j < x i for j > i and lim x ~ ( Q ) = 1. . of the i t h type of feed. .qn).3 . x3(ql. but. Apparently.)) 1 + exp(-)~.m.S. Q-~0 Q--. Q = total c o n c e n t r a t i o n of feed.(IB]-ms) ) " Here the Xs and m s parameters have the same meaning as for the s-shaped sw o o Q 10 2b Fig. To describe feeding with switching we used the findings of W. . whereas the curves x i ( Q ) for i = 2. whereas an experimentor uses only the empirically available values.([B] .
mB)])X[C]. The consumption of benthos by carp is represented as follows: qBC = FT(4) × FO(4) × v ( ~ r ~ '. Therefore. we describe the feeding with switching for bighead. FO(4) is the carp growth rate as a function of DO concentration. With . when switching to a new type of feed. BcR ) -- x. KBC. In this case we have a two-step switching. Thus. and detritus is the constrained feed. Ksc. whose favourite feed according to the literature is zooplankton. Phytoplankton is the substituting feed. BcR is the critical value of the benthos concentration at which carp switches to feeding on zooplankton. [B])]. [Z])Xn([B]. Moreover. By analogy.. in the abundant feeding of silver carp. the relationship between food assimilation and the ration values is taken into account for fish. The difference between the two trophic functions under the minimum sign ensures that the growth rate of carp is no more than the one attained for BcR.X. (d) Metabolism The excretion of the products of metabolism by the living organisms of the ecosytem may be considered to be approximately in proportion to the total uptake of food. [B]) × [ C ] . the food is continuously swallowed and passes through the intestine so quickly that only 30-40% of it can be assimilated.324 function for oxygen. (qFz + qDZ) is the ration of zooplankton. Here FT(4) is the carp growth rate as a function of temperature. the excretion of the products of metabolism and their transformation into detritus is represented in the following way (for zooplankton): q(1) = M B z × ( qFZ + qDZ) ZD where M B z is the metabolism parameter for zooplankton. Energy losses are taken into account by the outflow qzE = M B O z × [ Z] where M B O z is the zooplankton respiration coefficient. the uptake of zooplankton by carp under a deficit of benthos can be written in the form: qzc = FT(4) × FO(4) × rain([ KB c. qDz is the function of detritus uptake by zooplankton.and zooplankton. The FT(4) and FO(4) functions have forms similar to those for phyto. qFZ is the uptake function of the phytoplankton by zooplankton. For instance.
depending on the DO concentration in the water. Probable fish kills from oxygen deficit are described by another mechanism.[ 0 ] ) 1 ) .and zooplankton. F O X () = 1 + K A / [ 0 ] . There should be no natural mortality of fish in the optimum conditions of a fish-breeding pond. 1975). 8): E2() = exp[ C O P ( M . almost twice as much food can be assimilated (Pushchina. M is the threshold between the aerobic and anaerobic conditions. where COP is the parameter of the steepness of the oxygen curve. This model accounts for mortality only for phyto. the zooplankton mortality can be described in the following way: q(2) =FOX([O]) X M z X [ Z ] ZD Here F O X is the function of the mortality dependence on the DO concentration.[ 0 ] ) ] / ( 1 + exp[ C O P ( M . say. Therefore.325 moderate feeding. mineral phosphorus as a result of detritus destruction and dissolution can be described in the following way: qop= UDP X E l ( T ) XE2() X [ D ] . Artificial aeration is provided when oxygen concentration passes the threshold anaerobic value. Thus. the formation of. KA is the coefficient of mortality increase under oxygen deficit. Here the dependence of the destruction rate on temperature is given by the Vant-Hoff function E1 ( T ) = 2 (r-2°)/a° The oxygen function for this process is (Fig. (e) Mortality As we know from the literature. mortality is in the first approximation proportional to the biomass or concentration of living organisms. the mortality of living organisms depends on the DO concentration in the water. (d) Destruction The destruction process of the dead organic material. If this external factor is constant. M z is the mortality coefficient. which produces the basic nutrients--phosphorus and nitrogen. and benthos. .--depends on the thermal conditions and on the presence of DO in the water.
Oxygen function for detritus destruction process.COD' < M >~M Here C O D = M . where /0. in the main. is worth considering.326 E2 1 o o ~ O2 ~b Fig. . only the inflow of phosphates from detritus. decomposed in the water body. (h) O x y g e n flows The oxygen content in the fish pond water depends on the enrichment of water with oxygen and the rate of its consumption. 8. E3([01) = -M [ [01 . the process takes the opposite tendency: dissolved phosphates turn to insoluble forms and sink to the bottom. Thus. and S E D P is the maximum sinking rate of phosphorus.C K . mineral phosphorus begins to flow intensively from sediments into the water. where P H O T is the assimilation coefficient. qpD = S E D P X E3() X [ P ] . where C K is the steepness of the oxygen function. under anoxic conditions. The inflow of oxygen due to photosynthesis is proportional to the phytoplankton production and can be described by the function. qFo = P H O T x It. Under aerobic conditions.
qD~ . Inflow of oxygen with artificial aeration is also taken into account.14.qzc -.q o B - qoc -- qos - qOH -- qoD dA d~ = A U( t )- q A c .q F E dZ dt = qFz + qoz -..qDz q D B . level 1 dF d-~ = q a F .q F H .qsE = qzH + qFn + qDH -.qoz -. for example: qoz = R E S P z × [ Z] for zooplankton.q F Z .40420 × T + 0...S E D × D ..qzn -.qce = q~+ qos - qsD . Os -.~D dD d~-t = q F D + q Z D + q B D + q C D + q S D + q n D + q A D qoe .. TABLE I Differential equations..00009 × T 3. The consumption of oxygen for the respiration of aqueous organisms and plants is proportional to their biomass..qzD -.0.q B D .q F $ -.q F B ..0.327 The exchange processes with atmospheric oxygen are characterized by the formula R=RE×(Os--[OI).qZE dB d--7 = q~m + q z B + q D B ..qno -.q B E dC d---7 = q A c + q B c + q z c dS d~ dH d t - qcD .q H E dP d---~ = q o e dN -~ + PU(t)- qeF - qeD = qoN + NU(t)- qNF d[O] dt = qFo + RE > ( REA - )+ OU( t )- qoF -. Following Wang et al.. where R E is the reaeration coefficient that is dependent on the wind velocity in the general case.q D s -.q D u .q. Os is the oxygen saturation concentration.00842 × T 2 .q n c .q F D .qzB -.61996 . (1978).
m F ) ] ) M B F X qcl:. K a o a ) x C qBc = FT(4) x FO(4) × V(I. KZH. Y2)× H qDn = FT(6)× FO( 6) ×rrfin([min( Y1.n fi n ( Y1. KzB. Knc. qos = FT(5) X FO(5) X V ( # n ~ . level 2 qGF = FT(1) × F F ( L . O )× Z __ 5 qpn = FT(3)× V(Fn~. F ) × B qzn = FT(3)X V(/. Xz. FcR)× ~I( Z. D)X S q z n = FT(6) X FO(6)X V ( # ~ . K z o Z)× 7q(B. Y2 )]. F. XF. F ) × S ~x. qan = MBB X(qFB + qzB + qoB) qzo = F < Fo F > Fo M B z X ( q v z + qDz) + FOX([O])X M z X Z qcD=(MBc+ MBBcX RCm~. Z ) Y2 = V ( # ~ . K z n . m z ) Y3 = V ( # ~ ~. K r z . K a o [ V ( # ~ x. D)× B qAc = FT(4)X FO(4)X V(#'~. ZcR ) . K z n . D) × # ~ x × min( __ 1 Kp F + 2 p2 1 N2 p 2 ' 5 K ~ r + N2 ) X F qPr -. Kos. K o z . )~z. Z)X B qos = FT(3)× V ( # ~ . RS etOmax RC )XRC qsD = ( MBs + MBBs × ~--~ff---)× R S . Y3 ) . Kns. qro = ( MB r x qcF + MF X F.t~x.T6g X qGr qFZ = FT(2)X FO(2) × V(p.~. g o n . Z)X H B)]. F ) X ~(Z.328 TABLE II Flows.~x.V(Ix'~. B)X C qrs = FT(5) X FO(5) × V(#~-s K FS.Trig)< qGF qNF -. m z ) qeH = FT(6)× FO(6)×min(Y1. F ) × Z qoz = FT(2)× FO(2)× V ( # ~ .t~". KFB . Knc. )~z. qzc = F T ( 4 ) × FO(4)×min([V(#~. An. KFr. D)X r/(Z. KFr. mz)X rt(F. [ V ( # ~ . ran)l)× C Y1 = V ( I t ~ . BcR)--V(t*~.
329 TABLE II (continued) RH )X RH qno = (MBn + MBBn × ~ R C = qac + qBc + q z c R S = qrs + qDs R H = qZH + qFn + q o n qFE = MBOv X F qZE = M B O z × Z qnE = M B O o × B qce = MBOc X C qse = MBOs X S qHE = MBOn × H qDP = U D P × E l ( T ) × E2()× D qPD = S E D P × E3()× P qDN = U D N X E I ( T ) X qFO = P H O T X qCF qOF = R E S P F X F qoz = RESPz X Z qoB = R E S P n X B qoc = RESPc X C qos = R E S P s × S qon = RESPH x H qAD = A L P H A X A E2()× D where R E S P z is the  consumption coefficient for the zooplankton respiration. we have presented the main flows of the material necessary for the description of the fish pond ecosystem. where O K is the oxidation coefficient. The oxygen consumption for oxidation of dissolved and suspended organic material in water is proportional to the detritus amount involved in the cycle qoo = O K × [ D ] . It should just be added that an input of feed for fish and an input of mineral fertilizers are given by external . Thus.
)l) O.150). TO(I). and the regime of artificial aeration. E3() =  .K.k x h)xexp[1 . of feed. Some of the parameters were determined from previous works . level 3 [exo(46( 141 QI(I) ] 1' ( 4"6X( T-TO(I)]4] exp T< TO(I) T>~ TO(I) FT(I)=FT(T. The system of 11 ordinary differential equations is solved by the R u n g e . L L FF(L. F.tmax. To solve the system we had to specify 112 parameters. D) = ~'optexp(.~optmexp(-k × h)] k=KW+ K F x F + K D x D x K P D FO(1) = 1/(1 + exp( .M ~ [O]-COD' [01 < M [O]>/M inflows. parameter estimation is quite a problem.X(I)(. the inflows of nitrogen and phosphorus fertilizers.)]/(1 + exp[COP(M . X) #max × X 2 K 1 + X2 FOX([O])= 1 + KA/[0l El(T) = 2(r-z°)/l° E2() = exp[COP(M . the arrays of temperature and illumination. SIMULATION EXPERIMENTS The computer runs of the Fortran program for the model were carried out on BESM-6. while the sedimentation process of detritus and the settling of phosphorus (losses from the material cycle) are characterized by outflows.Q2(I))= 11. m) = e-X(x-'n)/(1 + e -x(x-m)) V(/.QI(I).K u t t a technique with an automatic choice of the step in the interval (0. As with all simulation models of this kind. )~. The flows they include and the functional relationships are listed in Tables II and III respectively. All the model equations are given in Table I. 4.330 TABLE III Functions.m(I)))) • q(X.
More difficulties arise because certain parameters characterize extremely aggregated processes.0 0. Wang et al.02 1.0 0.06 5. 1978.2 0. Grygierek. TABLE IV Ecological parameters Notation Ecological meaning Maximum growth rate of phytoplankton Maximum uptake rate of phytoplankton by zooplankton The corresponding half-saturation parameter Maximum uptake rate of detritus by zooplankton The corresponding half-saturation parameter Maximum uptake rate of phytoplankton by benthos The corresponding half saturation constant Maximum uptake rate of zooplankton by benthos The corresponding half-saturation constant Maximum uptake rate of detritus by benthos The corresponding half saturation constant Maximum uptake rate of zooplankton by carp The corresponding half saturation constant Maximum uptake rate of benthos by carp The corresponding half-saturation constant Maximum uptake rate of artificial feed by carp The corresponding half saturation constant Maximum uptake rate of phytoplankton by silver carp The corresponding half-saturation constant 3.0 0. and barley was used as the fodder for carp. it is rather difficult to find unique quantitative values for them.2 15.4 1. The parameters resulting from the model identification are presented in Table IV.0 0.0 Units 1/day 1/day mg/l 1/day mg/l 1/day mg/1 1/day mg/1 1/day mg/1 1/day mg/l 1/day mg/l 1/day mg/1 1/day mg/l (continued) ~.2 60. 1978)..).) and on three groups of ponds with the addition of silver carp (4.000.5 60.4 15. 8. 1978. For model calibration we used the data of the Polish scientists (Opuszynski.0 0.000 fish/ha.000 and 1.~ax max ~FZ KFz IXDZ KD~ I~FB KFB I~ZB KZB I~DB KDB I~ZC Kzc max l~BC KBC max I~Ac max max max K~ K FS .0 0.0 0. Wasilewska.03 0. etc. 1978.200 fish/ha. and others should be identified according to the variable dynamics. some can be estimated on the basis of the literature data. The fish ponds were regularly fertilized (by urea and superphosphate)..0 1.).331 (Jorgensen et al.1 20. 1978. who carried out complex studies on a group of test ponds with carp as a monoculture (4.
001 0.15 1.00004 0.05 0.01 0.1 60.01 0.4 0.002 1.001 0.3 0.0 0.09 0.3 0.3 0.05 Units 1/day mg/1 1/day mg/1 1/day mg/1 1/day mg/l dimensionless dimensionless dimensionless dimensionless dimensionless dimensionless dimensionless dimensionless dimensionless rag/1 rag/1 rag/1 dimensionless dimensionless dimensionless dimensionless dimensionless dimensionless 1/day 1/day 1/day 1/day 1/day 1/day 1/day 1/day 1/day KDS max ~FH KFH max I~zH KZH max ~DH KDH MB r MBz MB s MB c MBs MBI~ MBB c MBB s MBB~ RCmax RSm~ RHm~.07 60.001 0.0 0.3 0.001 0.01 0.3 0.332 TABLE IV (continued) Notation tXns max Ecological meaning Maximum uptake rate of detritus by silver carp The corresponding half saturation constant Maximum uptake rate of phytoplankton by bighead The corresponding half-saturation constant Maximum uptake rate of zooplankton by bighead The corresponding half-saturation constant Maximum uptake rate of detritus by bighead The corresponding half-saturation constant Metabolism parameter for phytoplankton Metabolism parameter for zooplankton Metabolism parameter for benthos Minimum metabolism parameter for carp for silver carp for bighead Additional metabolism parameter for carp for silver carp for bighead Maximum ration for carp for silver carp for bighead Respiration coefficient for phytoplankton for zooplankton for benthos for carp for silver carp for bighead Mortaility coefficient for phytoplankton for zooplankton for benthos D O consumption parameter for F respiration for Z for B for carp for silver carp for bighead Phosphorus destruction parameter Nitrogen destruction parameter Assimilation coefficient Transformation coefficient of fodder into detritus Reaeration coefficient Detritus oxidation parameter Sedimentation parameter of detritus 0.4 13.0 10.1 20.3 0.0 0.005 0.0 0.11 0.001 0.3 0.001 0.0 0.01 0.085 0.2 0.0 0.0 10.4 0.001 0. MBOF MBO z MBO B MBO c MBO s MBO H MF Mz Me RESP F RESP z RESP B RESP c RESP s RESP H UDP UDN PHOT A LPHA RE OK SED .
03 0.0 26.0 3.0 26.0 3000.0 3.0 35.0 24.0 9.0 20.0 3.0 1.0 2.0 1.0 2.0 1.0 1. Tm3in T4.0 13.1 24.2 0.0 10.0 1.0 26.0 1.0 Units oC oC oC oC oC oC oC oC oC oC oC *C oC oC °C oC oC oC SEDP TO(l) TO(2) TO(3) TO(4) TO(5) TO(6) Tli.0 3.0 9.0 34.0 34. T~ax T2ax Tm3ax T4~x TmS~ Trax Lopt KW KF KD KPD h m(2) m(4) m(5) rn (6) X(2) X(4) A(5) A(6) BcR ZcR FcR rn B rn z mF As AZ AF KA m COP M COD .0 30.0 35. Tr.0 1.0 34.5 0. TmSi.0 10.0 35.0 5.4 0.0 9.1 3.0 5.333 TABLE IV (continued) Notation Ecological meaning Sedimentation parameter of phosphorus Optimum temperature for F growth for Z for B for C for S for H Minimum temperature for F growth for Z for B for C for S for H Maximum temperature for F growth for Z for B for C for S for H Optimum illumination for photosynthesis Light extinction coefficient in water Self-shading parameter for phytoplankton Shading parameter of detritus Fraction of detritus suspended in water Mean photosynthesis depth Oxygen half-maintainance parameter for Z for C Oxygen half-maintainance parameter for S for H Steepness of the oxygen curve for Z for C for S for H Critical value of benthos concentration Critical value of zooplankton concentration Critical value of phytoplankton concentration Parameter of the switch function for B for Z for F Steepness of the switch function for B for Z for F Parameter of mortality increase at DO deficit Stoichiometric ratio Parameter of the oxygen function Parameter of the oxygen function Threshold between oxic and anoxic conditions 0.0 0.0 1.0 13. TEi.0 24.0 2.0 15.
11 shows the succession of the maxima of phytoplankton. Under other conditions. N = 1 . this being quite natural for such a dense fish population. .0 .334 5. whereas bighead and silver carp begin to grow only in July. but its concentration may increase at the beginning of the season. 9.06 . zooplankton. the natural fodder reserves are better developed. since their growth is to a greater extent limited by temperature. when the phytoplankton growth is not limited by nutrients. Right after their I" E3 J J o ~ 02 lb Fig.2 mg/1 (mostly due to fertilizers). 12). The next variant shows the ecosystem development in the conditions of a warmer climate (Fig. The phytoplankton growth is limited by nutrients (mostly phosphorus). Fish consumes a lot of zooplankton. and benthos concentrations. it turns out that the phytoplankton growth is limited by the amount of phosphorus. 6 mg/1. A lot of benthos is consumed by carp. we have derived the dynamics of the variables that adequately reflect the real picture of the development of the ecosystem for one season. When the nutrient concentration is maintained at a level of P = 0. Oxygen function for phosphorus sinking process. and the phytoplankton dynamics clearly follows the dynamics of phosphates (in Fig. Carp begins to gain weight at the very beginning of the season. RESULTS. 10 the arrows indicate the moments of introducing the fertilizers). CONCLUSIONSAND DISCUSSION The following graphs show different variants of the ecosystem development. in this case the zooplankton and benthos concentrations once again drop sharply in the middle of the season. Fig. However. As a result of the simulations.
+ = phytoplankton. t~ = bighead. 10. Ecosystem development in phosphorus limited conditions. Ecosystem development in the conditions of nutrient abundancy. + + 0~"------~--~-------%--~-------W¢~--'--'--.335 '°°I lOC 50 O0 5~)--"~ ~ TIME 100 ~ 150 Fig._. t~ = silver carp..zooplankton.. + = phytoplankton..  = carp.~ 0 50 TIME 100 150 Fig... ~ = bethos. 11.-------.+ ~ + .. .. × --. 150 50 .
1977. the artificial aeration. Mathematical modelling experience in a simple fishpond ecosystem. fertilizers are quickly consumed. application. Ecosystem development in the conditions of a warmer climate. X = silver carp. 99: 81-92. x = zooplankton. Nauk rol.. (T= To +4°C).. Rocz. . The control parameters are the inflows of fertilizers and fodder.336 200- 150 100 50 0 50 TIME 100 150 Fig. H. The influence of the silver carp on eutrophication of carp ponds. Grygierek.N. IV. REFERENCES Borshev. In: Fish Breeding in Ponds. and the p h y t o p l a n k t o n concentration increases. T he a m o u n t of detritus has increased by the middle of the season. The oxygen control unit is so designed that the oxygen c o n ten t never drops below 3 mg/1. VNIIPRH. + = phytoplankton. Th e next stage envisages the search for the o p t i m u m regimes of the fish pond.  = carp. The artificial aeration is increased when this threshold is passed. 12. E. The mathematical model allows one to find the operating modes that maximize the yield. N = bighead. ~ = bethos. and then begins to diminish due to its uptake by bighead and silver carp. 1978. Zooplankton. and the density of implantation.. V. 18:260-281 (in Russian).
Pishchepromizdat.. The influence of the silver carp on eutrophication of carp ponds... A. Rocz. D. Pushchina.V. Vielkind. 1978. V. Luckyanov..S.. W. 1978.I. Januszko. and Wang. Lake Management... The influence of the silver carp on eutrophication of carp ponds. Vinberg. (in Russian). Mathematical model of an aquatic ecosystem..I. The influence of the silver carp on eutrophication of carp ponds. M. and Svirezhev... Denmark. Henriksen.G. J.E.G. M. 1978. Friis.. Recapitulation. Handbook of Environmental Data and Ecological Parameters. Experimental ecology of fish nourishment. 1980.H. (in Russian). Jorgensen. Mathematical models of dissolved oxygen concentration in fresh water. Voinov. (Editors). Moscow..A.. 99: 93-108. I. B. Wasilewska. Pergamon Press.. N.P. VII. Gidrometeoizdat. 23 pp.. 99: 127-151. 1981.. H. 1978. Yu.B. V~eerl~bse. H.A. V. Phytoplankton. H. Oxford. Rocz. IV. 99: 7-32.M.F. Steele. The influence of the silver carp on eutrophication of carp ponds. L. Vinberg. and Liachnovitch.E. Physico-chemical conditions. Nauk rol.K. III. Opuszynski. 1966. Phytoplankton of fishbreeding ponds in Krasnodar region under intensive management. Fertilization of the ponds. Linmol. Jorgensen. Leningrad. Ecol. Moscow. 167 pp. Rocz. and Mejer.K. 1975. Pishchepromizdat. 1962. In: Photosynthetic Systems of High Productivity. Leningrad (in Russian). 1978.. Moscow (in Russian). Oceanogr. Nauk rol. S. I.. Wang. G.S.. Environmental control of photosynthesis in the sea. 5: 115-123. 271 pp.H. 1978.E. Piotrowska. S. (in Russian).M... Lake ecosystems modelling. L. 99: 55-80. Voronkova. Nauka.. W. V. Nauk rol. Bottom fauna. G. M. 7: 137-150. J. 1965. 1955. 252 pp. S. O. Modelling.337 Ivlev.. Jorgensen. Nauk rol. L. H.. Rocz. . H. In: Ecological Monitoring Problems. and Anisimov.
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