From the numerical model to the educational software: Lake Life
Transcription
From the numerical model to the educational software: Lake Life
Annls Limnol. 28 (2) 1992 : 175-189 From the numerical model to the educational software : Lake Life J . M . Thébaulti M.J. Salençon 2 Keywords : Ecosystem model, educational software, hydroelectric reservoir. « Lake Life » is a software program designed to introduce the lay person to lacustrian ecology and to the basic concepts of hydraulic management. Having become familiar with the dynamics of the trophic system as well as the mechanisms leading to eutrophication, the user may experiment with managing the reservoir of his choice. Change over time in the principal elements in the ecosystem is calculated by a mathematical model. This paper first presents the successive steps in development of the model : — choice of variables : three plankton groups, fish, nutrients, oxygen ; — representation of complex mechanisms : for example, the vertical structure simulated in two layers ; — transcription into equations. This recapitulation will initiate the reader into the problems of modeling an ecosystem. We shall then analyse a few simulated situations : variations in plankton groups in three situations with increasing trophism, and one example of the impact of turbining on the fish and planktonic populations and on oxygenation of the hypolimnion. The quite realistic behavior of the simulations makes this software an excellent teaching tool. Du m o d è l e numérique au logiciel didactique : la vie d u lac Mots clés : Modèle d'écosystème, logiciel didactique, retenue hydroélectrique. « La Vie du lac » est un logiciel destiné à initier un public non averti à l'écologie lacustre et aux notions fondamentales de la gestion hydraulique. Après avoir pris connaissance du fonctionnement de la chaîne trophique ainsi que des mécanismes conduisant à l'eutrophisation, l'utilisateur a la possibilité de gérer une retenue de son choix. L'évolution dans le temps des principales composantes de l'écosystème est calculée par un modèle mathématique. Nous présentons ici les étapes successives qui ont permis l'élaboration du modèle : — le choix des variables : trois groupes planctoniques, poissons, nutriments, oxygène ; — la schématisation de mécanismes complexes : par exemple la structure verticale simulée en deux couches ; — la mise en équations. Cette démarche constitue une bonne initiation à la modélisation d'un écosystème. Nous analysons ensuite quelques simulations : l'évolution des communautés planctoniques dans trois situations de trophie croissante, puis un exemple de l'impact du turbînage sur les populations et sur l'oxygénation de l'hypolimnion. Le comportement suffisamment réaliste des simulations permet d'utiliser ce logiciel comme support de cours pour l'enseignement. 1. Introduction « L a k e Life » is a s o f t w a r e p r o g r a m designed to describe for a lay public the m a n n e r in which a lacustrian ecosystem reacts t o disturbances, whether natural or related to h u m a n activities. M a k i n g use of 1. L a b o r a t o i r e d ' H y d r o b i o l o g i e , U R A 6 9 5 d u C N R S , U n i v e r sité P a u l S a b a t i e r , 118, R o u t e d e N a r b o n n e , 31062 T o u l o u s e Cedex, France. 2 . E l e c t r i c i t é d e F r a n c e , D i r e c t i o n d e s E t u d e s et R e c h e r c h e s , D é p a r t e m e n t E n v i r o n n e m e n t , 6, Q u a i Watier, 78401 C h a t o u Cedex, France. a n i m a t e d images, a choice of information level, a n d a user-friendly m o u s e , it gives a n attractive present a t i o n of the v a r i o u s c o m p o n e n t s in a lacustrian ecosystem : nutrients, phyto- and zooplankton, fish, etc. It also describes the principal mechanisms at work : t h e r m a l stratification, interaction of v a r i o u s kinds a m o n g t h e links in t h e trophic chain a n d t h e d y n a m i c s of e u t r o p h i c a t i o n . Once these fundamental concepts have been review e d , the user is a s k e d to choose a lake type c h a r a c terized b y t h e period required for renewal of w a t e r , Article available at http://www.limnology-journal.org or http://dx.doi.org/10.1051/limn/1992015 176 J.M. THEBAULT, M.J. a n d t o assume c o n t r o l over its future by m a n a g i n g n u t r i e n t c o n c e n t r a t i o n s , p a r a m e t e r s of hydraulic m a n a g e m e n t ( w i t h d r a w a l levels a n d p e r i o d ) as well as a n y s t o c k i n g with y o u n g fish. Critical p e r i o d s ( d u r i n g which water is unfit for s w i m m i n g , fish sur vival, etc.) a r e indicated by a l a r m s . T h e u s e r ' s g o a l is, of c o u r s e , t o fight e u t r o p h i c a t i o n while integra t i n g t h o s e m e c h a n i s m s which c o n t r i b u t e t o m a i n t a i ning water quality. In o r d e r for a simulation closely to a p p r o a c h rea lity, it is necessary t o use a numerical m o d e l which s i m u l a t e s the ecosystem d y n a m i c s . T h e great n u m ber of possible responses the model may give, depen ding o n the user's choice and the history of the reser v o i r , precludes use of pre-established scenarii. T h e m o d e l performs « on-line » calculation of variations in the m a i n ecosystem c o m p o n e n t s , before the user's eyes, enabling h i m to visualize continually the reper cussions of his system m a n a g e m e n t . This a p p r o a c h closely parallels t h a t m a d e of decision-support m o d e l s , a n d is, for t h a t r e a s o n , a g o o d i n t r o d u c t i o n t o o n e of t h e key functions of m o d e l i n g . T h e « L a k e Life » model is a simplified version o f t h e c o m p u t e r c o d e perfected for study of P a r e l o u p L a k e ( T h é b a u l t & Salençon 1992). T h i s p a p e r presents the c o n c e p t s b e h i n d the m o d e l , t h e choice of t r o p h i c n e t w o r k a n d variables, as well as t h e simplifying h y p o t h e s e s it was neces s a r y t o m a k e . O n e of the m a j o r p r o b l e m s c o n f r o n t i n g us w a s t o find a c o m p r o m i s e b e t w e e n the m o d e l ' s complex form of representation, integrating t o a high degree present hydrobiologjcal k n o w - h o w in t h e field of reservoir ecology, a n d a calculating t i m e which seemed reasonable for a m i c r o c o m p u t e r . T h e p u r p o s e of t h e model is a b o v e all p e d a g o g i cal ; for this r e a s o n , it is essential t h a t t h e simula tions be characteristic — even caricatures, in a sense — so as t o c o n s t i t u t e « teaching cases » w h i c h can easily b e assimilated b y t h e user. O n l y t h e p r e d o m i n a n t system m e c h a n i s m s have t h e r e f o r e been selected. W i t h apologies t o the specialists, we have simplified t o t h e extreme the m o r e complex mechanisms. F o r all these r e a s o n s , this m o d e l can absolutely n o t b e used as a m a n a g e m e n t t o o l , a n d even less as a research t o o l . T h e s o f t w a r e r u n s on A t a r i or P C - c o m p a t i b l e c o m p u t e r s ; simulation of o n e year requires approxi m a t e l y t w o m i n u t e s ' calculating t i m e . T h i s is SALENÇON (2) between 1000 a n d 5000 times faster than the time it would take similar c o m p u t e r s t o run an actual m a n a g e m e n t m o d e l of the P a r e l o u p type. 2. Description of the model 2.1. Physical structure T h e physical structure of the system is o n e of the elements in the p r o g r a m which required the grea test simplification, essentially for reasons of com puter memory. T h e water mass is divided vertically into two h o m o g e n e o u s layers which represent sea sonal thermal stratification. T h e surface layer, the epilimnion, is separated from the b o t t o m layer, the hypolimnion, by a t h e r m o c l i n e . T h e b o t t o m layer is in contact with sediment. Each layer is characterized by its t e m p e r a t u r e , volume, depth and m e a n illumination. The dyna mics of the two water masses are simulated in two phases : o n e phase of homogeneity in winter, when only the epilimnion exists, a n d o n e phase of strati fication in s u m m e r , when the thermocline level is constant. It was impossible t o introduce the tran sient spring thermoclines, variations in depth of the thermocline or the g r a d u a l deepening of the ther mocline in a u t u m n . T h u s , the onset of stratification and the a u t u m n mixing are instantaneous ! Inflow and outflow of water are represented very schematically : inflow is always localized in the sur face layer ; outflow due to turbining may be at the surface or on the b o t t o m , with the user free to deter mine the outlet level for each season. The program proposes a choice among three types of reservoir, characterized by the period required for water renewal : a short.residence time of from two weeks to a m o n t h ( G r a n g e n t , for example), from o n e to three m o n t h s (as for C h a m b o n , for exam ple), a n d on the order of one year for reservoirs with a p r o t r a c t e d residence t i m e (such as P a r e l o u p and Sainte-Croix). T h e length of the period of stratifi cation a n d exchange between the two layers differ according to the type of reservoir : in the first case, no stratification is introduced ; in the second, there is stratification, but t h e r e is nonetheless a low but constant a m o u n t of exchange between the two layers ; in the third case, stratification lasts from late spring t o m i d - a u t u m n , and there is no exchange except in the event of t u r b i n i n g . F o r the last two (3) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE LIFE 177 reservoir types, t u r b i n i n g t h r o u g h the lower outlet triggers m o v e m e n t from t h e epilimnion t o w a r d the hypolimnion. T h e fish prey o n z o o p l a n k t o n . We a s s u m e here that the population is homogeneous over time, which permits introduction of a single v a r i a b l e . There are two possible sources of phosphorus and nitrogen : c o n c e n t r a t i o n s from upstream and any discharge of water containing some concentrations of these same elements. T h e residence time deter mines the annual inflow-outflow rate. All variables, with the exception of fish, are withdrawn by turbining. Detritus is composed of algae a n d dead a n i m a l s , p h y t o p l a n k t o n which has settled, and m a t t e r n o t assimilated by animals. Bacteria a r e not simulated in this m o d e l . We assume that they are c o n s t a n t l y effective in mineralizing organic matter. 2.2. The variables Oxygen is a n i m p o r t a n t indicator of w a t e r q u a lity, a n d determines the possibility for survival of m o s t fish species. This variable is therefore inclu ded in the model. Forcing variables : These variables are directly entered into the model a n d are therefore not calculated : — respective volumes of the two layers a n d of the sediment, — inflow a n d outflow rates, — c o n c e n t r a t i o n of N a n d P in inflow, — m e a n light energy in the epilimnion, represen ted schematically by a sine curve with a period of 365 days. It is supposed that light energy in the hypo limnion is t o o low t o permit algal growth ; — the t e m p e r a t u r e of each layer, also represen ted by a sine curve. D u r i n g stratification, the h y p o limnion t e m p e r a t u r e remains low and stable. State variables in t h e m o d e l : T h e model is based o n a p h o s p h o r u s budget, which is to say t h a t all variables, with the exception of nitrogen a n d dissolved oxygen, are expressed in p h o s p h o r u s units. Phytoplankton are divided into two groups : algae which may be c o n s u m e d by the z o o p l a n k t o n , and Cyanobacteria, capable of fixing atmospheric nitro gen a n d not c o n s u m e d by t h e z o o p l a n k t o n . In order t o limit the n u m b e r of variables, we do not simulate silica a n d therefore d o not distinguish a diatom group. Nutrients : b o t h g r o u p s of algae need p h o s p h o rus. We have introduced nitrogen as it may be a limi ting factor. It is c o n s u m e d by both groups of algae, but is n o t indispensable to Cyanobacteria. Z o o p l a n k t o n : o u r experience at P a r e l o u p led us t o distinguish a single g r o u p of herbivorous zoo p l a n k t o n , as the i m p a c t of c a r n i v o r o u s z o o p l a n k ton is limited in c o m p a r i s o n with prédation by plankton-grazing fish. 2.3. Modeled processes T h e d i a g r a m of the model dynamics is s u m m a r i zed in Figure 1 ; all equations and variables in Tables 1, 2, 3, 4 and 5. Phytoplankton T h e factors for phytoplankton production include light, t e m p e r a t u r e and nutrient concentration : all factors which can curb the p h y t o p l a n k t o n g r o w t h rate, especially in ranges of values far off o p t i m u m values. We have used M o n o d ' s equation (1942), a hyper bolic function which is well suited t o description of the relationship between phytoplankton growth a n d nutrient concentration. We have chosen a similar e q u a t i o n to represent the effect of light and t e m p e r a t u r e , introducing a threshold value below which t h e r e is n o g r o w t h . These functions are less sophis ticated t h a n those used in m o r e complete m o d e l s (Steele 1962, Parker 1974, Eilers & Peters 1988, Tal bot et al. 1991) : in particular, they d o not simulate t h e d r o p in growth rate at high t e m p e r a t u r e s or u n d e r conditions of photo-inhibition, which is less i m p o r t a n t in this model. They are, on t h e other h a n d , far simpler t o calculate. Real g r o w t h is defined as the product of a maxi m u m g r o w t h rate and the three limiting factors. T h e t w o groups of algae need p h o s p h o r u s a n d nitrogen for development. T h e first group is limi ted b y the least a b u n d a n t element (Leibig law : D r o o p 1974, 1975, Rhee 1978). C y a n o b a c t e r i a are limited only by a p h o s p h o r u s deficit ; they are capable of fixing atmospheric nitro gen w h e n mineral nitrogen is lacking. 178 (4) J.M. THEBAULT, M.J. SALENÇON Factors in disappearance : prédation by zooplankt o n a n d sinking a r e the factors for d i s a p p e a r a n c e o f t h e first g r o u p of algae. A t least two causes m a y b e f o u n d for m a j o r variations in sinking speed. O n e is physical, related to turbulence in the m e d i u m , and c a n n o t b e simulated here ; t h e other d e p e n d s on n u m e r o u s factors such as cell size, t e m p e r a t u r e , age o f the p o p u l a t i o n or lack of n u t r i m e n t . W e chose this latter factor for o u r model, linking sinking speed t o g r o w t h limitation d u e t o n u t r i e n t s . N a t u r a l m o r t a l i t y is considered to be a negligible f a c t o r in c o m p a r i s o n to p r é d a t i o n a n d sinking. C y a n o b a c t e r i a c a n generally not develop in a turb u l e n t m e d i u m a n d at low t e m p e r a t u r e s (Steinberg & H a r t m a n 1988, Z o h a r y & Breen 1989). A s we d o n o t s i m u l a t e t u r b u l e n c e , we have linked m o r t a l i t y in this g r o u p of algae to t e m p e r a t u r e ; field o b s e r v a t i o n s at P a r e l o u p indicate t h a t , generally speaking, agitation in the w a t e r mass is at its maxim u m level in winter a n d spring, when the temperature is the lowest. Zooplankton T h e rate of ingestion depends primarily in the short term on t e m p e r a t u r e a n d nutrient concentration. W e have simplified the model p r o p o s e d by Thébault (1985) which permits simulation of their c o m p o u n d e d effect. T h e function linking ingestion to the quantity of available prey has been replaced by a hyperbolic function (Mullin et al. 1975). T o this, we have added a threshold below which feeding is nil. Only the « m a x i m u m feeding » coefficient is a function of the t e m p e r a t u r e . This simplification permits a considerable reduction in the number of coefficients needed. Rates of assimilation a n d n a t u r a l mortality are constant. P r é d a t i o n by fish, the principal cause of Phytoplankton growth Grazing F i g . 1. D i a g r a m o f t h e d y n a m i c s o f t h e b i o l o g i c a l m o d e l . F i g . 1. D i a g r a m m e d e f o n c t i o n n e m e n t d u m o d è l e b i o l o g i q u e . (5) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE z o o p l a n k t o n disappearance, is directly proportional t o the fish p o p u l a t i o n . 179 LIFE T a b l e 1. S e d i m e n t a t i o n p r o c e s s . T a b l e a u 1. L a s é d i m e n t a t i o n . Fish W e assume that the population is stable over time, neither increasing n o r decreasing under n o r m a l c o n d i t i o n s of o x y g e n a t i o n . Below a first threshold for concentration of dissolved oxygen, the most fra gile fish (the S a l m o n i d a e ) die. If concentrations of dissolved oxygen c o n t i n u e to d r o p , no fish survive. Detritus a n d n u t r i e n t s Detritus settles at a constant rate. P h o s p h o r u s and nitrogen are mineralized from detritus, in w h a t is n o m o r e t h a n t r a n s f o r m a t i o n from an o r g a n i c t o a mineral f o r m , the speed of which is t e m p e r a t u r e d e p e n d e n t . W e do n o t distinguish t h e different forms of nitrogen o r p h o s p h o r u s . A fraction of these elements is stored in sediment a n d later released into t h e h y p o l i m n i o n w h e n it is anoxic (insufficient oxygen). 3aX 3Xi x 3t Sedimentation ; ' 3z in top layer V i ^ At = -S aXi X l to bottom layer Sedimentation in bottom layer Dissolved oxygen Dissolved oxygen is p r o d u c e d as a result of p h o tosynthesis a t t r i b u t a b l e t o p h y t o p l a n k t o n a n d gaseous exchange at t h e air-water interface, o r rea e r a t i o n . Such e x c h a n g e is faster from a u t u m n to spring, the p e r i o d d u r i n g which wind speed is gene rally h i g h e r . V 2 ^ 2 At = S X l aXi-S from top layer Sedimentation X 2 aX 2 to sediment into sediment layer 2.4. Numerical considerations As certain g r o w t h rates are extremely high (a high value for the derivative), o n e needs to have a small integration time s t e p . W e usually use the Runge a n d K u t t a f o u r t h - o r d e r m e t h o d for numerical integra tion (Legras 1971) in solving differential e q u a t i o n s in biological m o d e l s . While this m e t h o d is highly simple, it calculates the totality of the equations a n d related s u b r o u t i n e s four times per time s t e p . T h e finite difference m e t h o d is quicker (as c a l c u l a t i o n is performed once only), b u t less precise for an iden tical t i m e step. E r r o r d u e t o numerical i n t e g r a t i o n (and to a lesser degree t o c o m p u t e r error) can be esti m a t e d by calculating a m a s s budget. A c o m p r o m i s e between acceptable e r r o r for this type of software a n d speed in calculation led us to solve e q u a t i o n s b y m e a n s of finite differences with a t i m e step of 1/10 of a d a y . V ^S-=S At s X l aX 2 for X = A , C, D a = mean area of the lake. In order to simplify expressions, sedimentation c o r r e s p o n d i n g to each layer i and variable X has been written in table 2 : O 3Xi 180 J.M. THEBAULT, M.J. (6) SALENÇON T a b l e 2. S y s t e m of d i f f e r e n t i a l equations. T a b l e a u 2. L e s y s t è m e d e s é q u a t i o n s différentielles. In water layers 3Ai -G R . = S I , . T dAi (GA-VJ-)A,-^Z-. - A S =(a rz-Hz-|4Z - P F z F = (ccfpf - N ) F F SÇl. 3t 3D „ , ,„ . „, , ,„ . S „ „_3D: m C , + « l - a ) r + | i ) Z + « l - a ) p + Hf)F - (m, + ^ - ) D , - V - g p c c 2 -gp = ^ . z 3Ai S A + V z z 3Di d F F ^ I A I 1 = r m,D np - (VN + - ^ ) N 2 a l 2 + <p N N p A c ^ S m In sediment ^ s 1 A = max ( 0 , . X + 0 ( g + g ) - i ^ m D dD F X ^ - + m D , n p - g A ] n p - gcCinp - ^JN 3D = \|/pP + m D 2 ^jp = ¥NN _ 2 r 2 - cppP.s s + ni D np - cp N r s 3D; 2 a = Xpi nwD -g A -gcC -ii-P dNi dt D . .„ ... . .r, , S ^ « ! - z ) r z + Uz)Z + ( ( L - a ) P + J1 )F - (m, + ^ ) D - V + + U F N s I l - ^ - O, F 2 D (7) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE 181 LIFE T a b l e 4 . P r o c e s s e s r e p r e s e n t e d in t h e m o d e l . T a b l e a u 4 . L e s p r o c e s s u s r e p r é s e n t é s d a n s le m o d è l e . Phytoplankton growth Effect of light if 1 1|(D = I X D , CT| " K] + I - 2GJT if I < Table 3. Model variables. Effect of 1,(1) = 0 13, température Tableau 3. Les variables du modèle. ifT>r&R RRFT) = — - ~ ^ — k + T - 2t5r ifT£ti3 1 (T)= 0 t Forcing variables I M e a n solar i r r a d i a n c e in the u p p e r layer T] (Tj) T e m p e r a t u r e of t h e u p p e r (lower) layer Effect of T T nutrients V[ (V2) V o l u m e of the u p p e r (lower) layer 1 M = l for C y a n o b a c t e r i a Vg V o l u m e of the s e d i m e n t layer 1 M = M i n ( l p . \ ) f o r o t h e r algae X^, X p C o n c e n t r a t i o n o f v a r i a b l e s N and P i n the i n f l o w E Inflow in t o p layer Sj O u t f l o w from l a y e r i, i - 1 , 2 Growth p N rate : g = g 1 , 1 1 T Cyanobacteria M mortality if T < D3 1 m m (T)=Mmaxfl-- c \ State 0m variables Aj P h y t o p l a n k t o n g r a z e d by h e r b i v o r o u s z o o p l a n k t o n , i = l Cj Cyanobacteria, i = l Z H e r b i v o r o u s z o o p l a n k ton ifT>ro m (T)=0 ifl <ras s (l ) = v c M F Fish Dj Detritus. i=1.2,S Pj Dissolved inorganic phosphorus. i=l,2,S Nj D i s s o l v e d i n o r g a n i c n i t r o g e n . i = l ,2.S Oj Dissolved oxygen, i=l,2 A M m i , 1 - M Zooplankton Effect of temperature ingestion on r,n.,(T) = t z Ingestion K-rz + f rate i = 1 : t o p layer i = 2 : b o t t o m layer i = S : sediment layer IF A > GJA. RZ(A)=R, ifA<0J r (A)=0 A 2 Pish Ingestion rate ifZ>0Jz pKZ) = ifZ£GJ p (Z)=0 z Xf- F Nutrient regeneration from m,(T) = a , e& T detritus rate 182 J.M. THEBAULT, M.J. SALENÇON (8) 3. Simulation results Table 5. Model parameters. T a b l e a u 5. Les paramètres d u m o d è l e . Phytoplankton k , Kp k . k l p parameters Half-sa lu radon coefficient for limitation by respectively light, temperature, phosphorus and nitrogen n OBj, Uij Light and tempera cure threshold for growth y Maximum growth rate for algae 0„ Temperature threshold for Cyanobacteria mortality jl .„ maximum mortality rate for Cyanobacteria np N : P ratio for all variables w Sedimentation V parameters Maximum sedimentation rale M ffl Nutrient threshold below which set! i men talion begins s V Constant sedimentation rate for detritus D yf V N Uptake of P and N by sediment <Pp, t p N Release of P and N from sediment r Animal tp parameters Maximum ingestion rale at optimal temperature for respectively zooplankton and fish Half-saturation coefficient for r ^ f T ) tD Threshold concentration of algae for zooplanlnon feeding A (Bg Threshold concentration of zooplankton for fish feeding * . Half-saturation coefficient for r ( A ) and p (Z) O j , Op Assimilation rate U^ U Mortab'ty rate A z F Nutrient regeneration and oxygen parameters p. 1 Coefficients for «1,07) Diffusion of oxygen at the lake surface •p Oxygen produced by photosynthesis •„ Oxygen used in organic decay of detritus 0 Saturated oxygen concentration — F T h e most interesting simulations from an educational point of view are those related to reservoirs where the residence time is seasonal o r a n n u a l . T h e management scenarii the user adopts here can significantly modify the future of the reservoir, unlike the case of the reservoir with a low residence time where it is impossible to exercise any influence o n eutrophy in t h e river. In o u r model, when there is no hydroelectric power generation, the flow left above the d a m is equal t o inflow from the watershed. W e can suppose t h a t , in this case, the lake behaves like a natural lake. W h e n the nutrient concentrations are low (Simulation 1, Fig. 2), we note a springtime increase in phytoplankton, followed by an increase in herbivorous zooplankton. In the summer, when the lake is stratified, nutrients become a growth-limiting factor in the surface layer for the phytoplankton, which decline rapidly d u e to active consumption by the z o o p l a n k t o n . In a u t u m n , cooling in the surface layers triggers mixing of the nutrient-deficient epilimnion with the nutrient-rich hypolimnion, leading to a new development of plankton communities. In the hypolimnion, dissolved oxygen is consumed in decay of settled detritus. The autumn mixing p r o m p t s the d r o p in mean oxygen values for the entire water mass. As the d e m a n d for oxygen in the hypolimnion is not very high in this simulation, we note n o significant hypoxia following the mixing. If we increase the phosphorus concentrations upstream (Simulation 2, Fig. 3), the behavior of the springtime plankton is identical to that in the previous simulation but the biomass is greater. As phosp h o r u s is highly concentrated in inflow, nitrogen becomes a limiting factor in the epilimnion before phosphorus. This situation, linked to the rise in temperature and the decreased turbulence, fosters the development of Cyanobacteria, which assimilate excess phosphorus and fix dissolved atmospheric nitrogen. During the a u t u m n mixing, the impact of Cyanobacteria is so great that other algae can n o longer develop. T h e Cyanobacteria decline when the medium is more turbulent a n d the temperatures lower. In this case, as the Cyanobacteria are not consumed, there is only one period of zooplankton development, in the spring. (9) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE LIFE 183 At the end of summer, the hypolimnion is anoxic, a n d the a u t u m n mixing triggers a serious d r o p in oxygen concentrations t h r o u g h o u t the water mass, which may be fata! to certain fish species. the first and second simulations, depending o n the year (Salençon & C a p b l a n c q 1987, Salençon et al. 1988, 1989, 1990a, 1990b). The third s i m u l a t i o n could apply to a highly eutrophicated reservoir. If high p h o s p h o r u s concentrations are maintained over several years (Simulation 3, Fig. 4), o n e reaches a m o r e p r o n o u n c e d situation of e u t r o p h y , with m a j o r declines in oxygen level, even in the epilimnion. Figure 5 shows the effect of two other possible types of hydraulic management : the initial conditions and inflow concentrations are identical to those in Simulation 2 ; summer turbining is through either the u p p e r or lower outlets. Here we note a m a j o r decrease in Cyanobacteria a n d a marked a u t u m n peak for consumable algae and z o o p l a n k t o n . T h e model results correspond closely t o observations o n a n u m b e r of sites. For example, the first simulation could well apply to Sainte-Croix, an oligotrophic lake ( P o u r c h e r & Salençon 1990). Variations in the p l a n k t o n communities at P a r e l o u p situate this particular reservoir somewhere between T h e level of the turbining outlet has n o m a j o r effect o n variations in the epilimnic p o p u l a t i o n s . Turbining t h r o u g h t h e lower outlet, on t h e o t h e r h a n d , does permit better reoxygenation of t h e F i g . 2 . S i m u l a t i o n 1. A n o l i g o t r o p h y l a k e w i t h n o h y d r o e l e c t r i c p o w e r g e n e r a t i o n . A n n u a l v a r i a t i o n i n p l a n k t o n c o m m u n i t i e s ( c o n s u m a b l e a l g a e , C y a n o b a c t e r i a a n d z o o p l a n k t o n ) [a] a n d i n d i s s o l v e d o x y g e n in t h e e p i l i m n i o n a n d h y p o l i m n i o n [ b ] . D i s s o l v e d o x y g e n is e x p r e s s e d a s a p e r c e n t a g e o f t h e c o n c e n t r a t i o n u n d e r c o n d i t i o n s o f Fig. 2. Simulation saturation. 1. C a s d ' u n l a c o l i g o t r o p h e s a n s p r o d u c t i o n h y d r o é l e c t r i q u e . E v o l u t i o n a n n u e l l e d e s c o m m u n a u t é s planctoniques ( a l g u e s c o n s o m m a b l e s , c y a n o p h y c é e s e t ^ o p l a n c t o n ) ( a ) e t d e l ' o x y g è n e d i s s o u s d a n s l ' é p i l i m n i o n et l ' h y p o l i m n i o n ( b ) . L ' o x y g è n e dissous est e x p r i m é en p o u r c e n t a g e de la c o n c e n t r a t i o n à saturation. 184 J.M. T H E B A U L T , M.J. (10) SALENÇON —PHYTO —CYANO — ZOO /'A J F M A M J J A S O N D N D 100 80 60 40 20 J J F F M M A A M M J J J J A S A S O O N D Fig. 3 . S i m u l a t i o n 2. A n n u a l variation in p l a n k t o n communities ( c o n s u m a b l e algae, Cyanobacteria a n d z o o p l a n k t o n ) [a], in dissolved o x y g e n i n t h e e p i l i m n i o n a n d h y p o l i m n i o n [ b ] , a n d i n n u t r i e n t s ( P a n d N ) i n t h e e p i l i m n i o n [ c ] . D i s s o l v e d o x y g e n is e x p r e s s e d a s a percentage of the concentration under conditions of saturation. T h e high nutrient concentrations a n d the absence of m a n a g e m e n t foster t h e d e v e l o p m e n t of hydraulic Cyanobacteria. F i g . 3 . S i m u l a t i o n 2. E v o l u t i o n a n n u e l l e d e s c o m m u n a u t é s p l a n c t o n j q u e s ( a l g u e s c o n s o m m a b l e s , c y a n o p h y c é e s e t z o o p l a n c t o n ) ( a ) , d e l ' o x y g è n e d i s s o u s d a n s l ' é p i l i m n i o n et l ' h y p o l i m n i o n (b) e t d e s n u t r i m e n t s ( P e t N ) d a n s l ' é p i l i m n i o n ( c ) . L ' o x y g è n e dissous e s t e x p r i m é e n p o u r c e n t a g e d e la c o n c e n t r a t i o n à s a t u r a t i o n . L e s a p p o r t s i m p o r t a n t s d e n u t r i m e n t s et l ' a b s e n c e d e g e s t i o n h y d r a u l i q u e f a v o r i s e n t le d é v e l o p p e m e n t d e s c y a n o p h y c é e s . (11) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE J F M A M J J A S O N D J F M A M J J A S O N D LIFE F i g . 4 . S i m u l a t i o n 3 . A n e u t r o p h i c l a k e . A n n u a l v a r i a t i o n in p l a n k t o n c o m m u n i t i e s ( c o n s u m a b l e a l g a e , C y a n o b a c t e r i a a n d 185 zooplank- t o n ) [a] a n d in d i s s o l v e d o x y g e n i n t h e e p i l i m n i o n a n d h y p o l i m n i o n [ b ] . D i s s o l v e d o x y g e n is e x p r e s s e d a s a p e r c e n t a g e o f t h e c o n c e n tration under conditions of saturation. Fig. 4. Simulation 3. C a s d ' u n lac e u t r o p h e . E v o l u t i o n annuelle des c o m m u n a u t é s planctoniques {algues c o n s o m m a b l e s , c y a n o p h y c é e s et z o o p l a n c t o n ) ( a ) e t d e l ' o x y g è n e d i s s o u s d a n s r e p i l i m n i o n e t P h y p o l i m m o n ( b ) . L ' o x y g è n e d i s s o u s e s t e x p r i m é e n p o u r c e n t a g e d e la c o n c e n t r a t i o n à saturation. hypolimnion than does turbining t h r o u g h the upper outlet, by fostering exchange between epilimnion and hypolimnion. It would b e unwise t o turbine t h r o u g h the lower outlet at the end of s u m m e r , when the hypolimnion is anoxic ; oxidation a n d precipitation of iron a n d m a n g a n e s e , together with the potential effects on d o w n s t r e a m river quality, are clearly pointed out to the novice hydraulic m a n a g e r . We must r e m e m b e r t h a t the model takes into account the a c c u m u l a t i o n of nutrients in the sedim e n t , where concentrations may become very high. W h e n the h y p o l i m n i o n is anoxic, p h o s p h o r u s (and t o a lesser degree, nitrogen, through de-nitrification) is released into t h e water m a s s , proportionally increasing the a m o u n t available t o algae at t h e t i m e of deepening of the thermocline. Therefore, when the lake is highly e u t r o p h i c a t e d , it would be a mistake to imagine t h a t the situation will rapidly improve if concentrations are reduced. T h e m o d e l represents well the inertia attribuable to t h e q u a n tity of nutriment stored in the sediment. T h e m o s t effective a n n u a l hydraulic management a p p r o a c h is to turbine at the surface in spring, to eliminate a m a x i m u m of algae a n d z o o p l a n k t o n so as t o decrease the fraction in the sediment, and at the b o t t o m in s u m m e r , to eliminate detritus which m a y have settled and oxygenate the hypolimnion as m u c h as possible. It m a y take several years to r e t u r n to acceptable conditions. 186 J . M . T H E B A U L T , M.J. J F M A M J J F M A M J A J (12) SALENÇON S O N D N D N D 100 80 60 40 20 0 J A S O 100 80 60 40 20 J F M A M J J A S O F i g . 5 V a r i a t i o n i n p l a n k t o n c o m m u n i t i e s in t h e c a s e o f t u r b i n i n g d u r i n g t h e s u m m e r [ a ] . I m p a c t o f t h e c h o i c e o f o u t l e t l e v e l o n o x y g e n a t i o n in t h e w a t e r m a s s : t u r b i n i n g t h r o u g h t h e l o w e r [b] or u p p e r o u t l e t [c]. F i g . 5 . E v o l u t i o n d e s c o m m u n a u t é s p l a n c t o n i q u e s d a n s l e c a s d ' u n t u r b i n a g e p e n d a n t l ' é t é ( a ) . E f f e t d u c h o i x d e la p r i s e d ' e a u sur l ' o x y g é n a t i o n d e la m a s s e d ' e a u : t u r b i n a g e p a r la p r i s e de f o n d (b) o u d e s u r f a c e (c). 4. Conclusion While « Lake Life » is a n educational software p r o g r a m permitting the user to e m b a r k , mouse in h a n d , on a discovery of lake ecosystems, it is at the s a m e t i m e a simplified version of an actual compu ter code developed in the course of scientific research. T h e a p p r o a c h followed in developing this pro gram is identical t o that followed in development of ecosystem models used in lacustrian ecology studies. T h e modeled processes are relatively simplified, though the m o d e l ' s response is on the whole most satisfying. (13) FROM THE NUMERICAL MODEL TO THE EDUCATIONAL SOFTWARE : LAKE 187 LIFE T h e r e are m a n y potential uses for this software : as a teaching a i d , in sensitizing hydraulic m a n a g e ment specialists, in informing the general public, etc. Gentil S. 1982. — A n a l y s e de s y s t è m e e n écologie. U n e A s the user retains control over pollutants discharged, hydraulic management and stocking with young fish, he can visualize the repercussions of his decisions o n the ecosystem as a w h o l e . In this way, he discovers the m e c h a n i s m s which lead t o e u t r o p h i cation as well as t h e difficulties o n e encounters in trying t o find remedies. J a r g e n s e n S . E . 1976. — A n e u t r o p h i c a t i o n model f o r a l a k e . d e c a s ( L a c d ' A i g u e b e l e t t e ) . I n Modélisation et simulation eau des systèmes et végétation. Modelling, Ecol. 2 : 147-165. J e r g e n s e n S.E., M e j e r H . & Friis M . 1978. — E x a m i n a t i o n a l a k e m o d e l . Ecol. Modelling. Kremer J.N. & Nixon S.W. tem. Simulation 1 9 7 8 . — A coastal and analysis. of 4 : 253-278. marine ecosys- E c o l o g i c a l Studies 2 4 , S p r i n g e r - Verlag, Heidelberg : 217 p . L e g r a s J . 1 9 7 1 . — M é t h o d e s et techniques que. de t'analyse numéri- D u n o d , Paris : 323 p. tériennes. In addition t o t h e authors mentioned in this text, a n u m b e r of w o r k s dealing with modeling aquatic ecosystems might be consulted : Bierman 1976, C a n a l e et al. 1976, Jorgensen 1976, Jorgensen et al. 1978, Kremer & Nixon 1978, G a r ç o n 1981, Spain 1982, Salençon et al. 1984, Thébault 1984, Riley & Stefan 1988, A n d e r s e n & Nival 1989. Dominante E d i t i o n s d u C N R S , Paris : 17-62. M o n o d J . 1 9 4 2 . — Recherches To find out more étude mathématique de l'environnement. sur la croissance des cultures bac- Hermann, Paris : 210 p. M u l l i n M . M . , S t e w a r t E . F . & F u g l i s t e r F . J . 1975. — Ingestion by p l a n k t o n i c g r a z e r s as a f u n c t i o n of c o n c e n t r a t i o n o f f o o d . Limnol. Oceanogr., 20 : 259-262. P a r k e r A . 1974. — E m p i r i c a l functions relating m e t a b o l i c pro- c e s s e s in a q u a t i c s y s t e m s t o e n v i r o n m e n t a l v a r i a b l e s . J. Res. Board Can., 31 : 1 5 5 0 - 1 5 5 2 . P o u r c h e r A . M . & Salencon M . J . 1990. — Modélisation d u p l a n c t o n d a n s u n e r e t e n u e o l i g o t r o p h e : S a i n t e - C r o i x s u r le V e r d o n . Hydroécol. Appt., 1 : 91-134. R h e e G - Y u l l 1978. — Effects of N : P . a t o m i c ratios a n d nitrate limitation Ack nowledgements o n algal g r o w t h , cell c o m p o s i t i o n , u p t a k e . Limnol. « L a k e Life » was developed for the E D F « Mission Environm e n t » , in c o l l a b o r a t i o n w i t h t h e M y r i a d firm, which contribu- t e d t h e g r a p h i c s a n d t h e u s e r i n t e r f a c e . It exists n o w in t h e F r e n c h v e r s i o n , a n d will s o o n b e a v a i l a b l e in E n g l i s h , f o r u s e o n PC- Oceanogr., and nitrate 23 : 10-25. Riley M . J . & S t e f a n H . G . 1988. — M 1 N L A K E : a d y n a m i c l a k e water quality simulation model. Ecol. Modelling, 43 : 155-182. Salençon M . J . , Merle G. & S a b a t o n C. 1984. — L e réseau de c o m p a t i b l e c o m p u t e r s (3 1 / 2 " a n d 5 1 / 4 " d i s k e t t e s ) a n d o n A t a r i m e s u r e s h y d r o b i o l o g i q u e s d e la r e t e n u e d e G r a n g e n t ( L o i r e ) ; S T . It m a y b e o r d e r e d f r o m t h e M i s s i o n E n v i r o n m e n t f o r F F 9 5 . a n a l y s e p a r t i e l l e d e s r é s u l t a t s . Cahiers E L E C T R I C I T E D E F R A N C E , Mission Environnement, r u e d e Ea B a u m e , 7 5 0 0 8 26, tereau, du laboratoire de Mon- 15 : 7 - 1 2 . S a l e n ç o n M . J. & C a p b l a n c q J. 1987. — E t u d e de la r e t e n u e d e Paris. M Y R I A D S.A.R.L., 4 rue de Bordeaux, 31200 Toulouse. P a r e l o u p . Bilan des t r a v a u x réalisés en 1986 d a n s l e c a d r e de la c o n v e n t i o n E D F - M i n i s t è r e d e l ' E n v i r o n n e m e n t . Rap- p o r t H E - 3 1 / 8 7 . 5 , Electricité d e F r a n c e , Paris : 26 p . References S a l e n ç o n M . J . , T h é b a u l t J . M . & C a p b l a n c q J. 1 9 8 8 . — A n d e r s e n V. & Nival P . 1989. — Modelling of phytoplankton p o p u l a t i o n d y n a m i c s i n a n e n c l o s e d w a t e r c o l u m n . J. biol. Ass. U.K., mar. B i e r m a n V . J . Jr. 1976. — M a t h e m a t i c a l m o d e l of t h e selective e n h a n c e m e n t of blue green algae by nutrient enrichment. In : tic ecosystems, biochemical processes in aqua- A n n A r b o r S c i e n c e s , M i c h i g a n : 1-31. C a n a l e R . P . , D e P a l m a L . M . & Vogel A . H . 1976. — A p l a n k t o n b a s e d food w e b m o d e l for L a k e Michigan. In : C a n a l e R . P . ( E d . ) : Modelling tems, biochemical processes in aquatic ecosys- D r o o p M . R . 1974. — T h e n u t r i e n t status o f algal cells in contibiol. Ass. U.K., 54 : 825-855. D r o o p M . R . 1975. — T h e n u t r i e n t s t a t u s o f algal cells in b a t c h c u l t u r e . J. mar. biol. Ass. Eilers P . H . C . & Peeters J . C . H . d a n s le c a d r e de l a C o n v e n t i o n E D F - M i n i s t è r e d e l ' E n v i r o n - U.K., 43 p . S a l e n ç o n M . J . , T h é b a u l t J . M . & C a p b l a n c q J. 1989. — Etude d e la r e t e n u e d e P a r e l o u p . Bilan d e s t r a v a u x réalisés e n 1 9 8 8 d a n s le c a d r e d e l a C o n v e n t i o n E D F - M i n i s t è r e d e l ' E n v i r o n n e m e n t . R a p p o r t H E - 3 1 / 8 9 . 1 3 , E l e c t r i c i t é de F r a n c e , P a r i s : 77 p . S a l e n ç o n M . J . , T h é b a u l t J . M . & C a p b l a n c q J . 1990a. — Etude d e la r e t e n u e d e P a r e l o u p . Bilan d e s t r a v a u x réalisés e n 1 9 8 9 A n n A r b o r Sciences, Michigan : 33-74. n u o u s c u l t u r e . J. mar. de la r e t e n u e d e P a r e l o u p . B i l a n d e s t r a v a u x réalisés e n 1 9 8 7 n e m e n t . R a p p o r t H E - 3 1 / 8 8 . 2 3 , Electricité de F r a n c e , P a r i s : 69 : 6 2 5 - 6 4 6 . C a n a l e R . P . ( E d . ) : Modelling Etude d a n s le c a d r e d e la C o n v e n t i o n E D F - M i n i s t è r e d e l ' E n v i r o n n e m e n t . Rapport H E - 3 1 / 9 0 . 1 7 , Electricité de France, Paris : 67 p . S a l e n ç o n M . J., T h é b a u l t J . M . & C a p b l a n c q J . 1990b. — 55 : 5 4 1 - 5 5 5 . 1988. — A m o d e l for t h e rela- Etude d e la r e t e n u e d e P a r e l o u p . S y n t h è s e d e s t r a v a u x r é a l i s é s d a n s tionship between light intensity a n d the rate of p h o t o s y n t h e - le c a d r e d e l a C o n v e n t i o n E D F - M i n i s t è r e d e l ' E n v i r o n n e m e n t s i s i n p h y t o p l a n k t o n . Ecol. (mars 1 9 8 6 - mars 1990). R a p p o r t H E - 3 1 / 9 0 . 2 3 , Electricité Modelling, 42 : 199-215. G a r ç o n V. 1981. — M o d é l i s a t i o n n u m é r i q u e d ' u n écosystème a q u a t i q u e . A p p l i c a t i o n a u réservoir d e G r a n g e n t sur la Loire. T h è s e d e 3* cycle. U n i v e r s i t é P a r i s 7, P a r i s : 2 3 0 p . de France, Paris : 40 p. Spain J . D . 1 9 8 2 . — Basis microcomputer models in A d d i s o n - W e s l e y , R e a d i n g , M a s s a c h u s e t t s : 354 p . biology. 188 Steele J . H . J . M . T H E B A U L T , M.J. 1962. — E n v i r o n m e n t a l control of i n t h e s e a . Limnol. Oceanagr., photosynthesis forming Cyanobacteria and the eutrophication of lakes and Biol., A comparative study a n d mathematical modeling of temper a t u r e a n d light o n g r o w t h of three microalgae useful for wastewater treatment. Water Res., potentially 25 : 465-472. 1984. — Modélisation des premiers niveaux réseau trophique pélagique marin. Mise au point de les et s i m u l a t i o n d e séries e x p é r i m e n t a l e s . T h è s e d e 3 Université Paris 7, Paris : 94 p. stylifera D a n a ) . Effets de la t e m - p é r a t u r e e t d e l a c o n c e n t r a t i o n d e n o u r r i t u r e . J. Exp. Biol. EcoL, Mar. 93 : 223-234. Thébault J.M. & Salençon M.J. — Simulation mode! of a meso- 20 : 279-287. T a l b o t P . , T h é b a u l t J . M . , D a u t a A. & D e la N o u e J., 1991. — Thébault J.M. T h é b a u l t J . M . 1 9 8 5 . — E t u d e e x p é r i m e n t a l e d e la n u t r i t i o n d ' u n c o p é p o d e c o m m u n (Temora 7 : 137-150. Steinberg C E . W . & H a r t m a n n H . M . 1988. — Planktonic bloomr i v e r s . Fresh. (14) SALENÇON du modue cycle, trophic reservoir m o d e l . Ecol. (Lac Modelling de Pareloup, France) : Biological (in p r e s s ) . Z o h a r y T . & Breen C h . 1989. — Environmental factors r i n g t h e f o r m a t i o n o f Microcystis a h y p e r t r o p h i c l a k e . Hydrobiologia, aeruginosa favou- hyperscums in 178 : 1 7 9 - 1 9 2 . (15) FROM THE NUMERICAL MODEL LA VIE DU LAKE LAC Electricité d e France Mode TO THE EDUCATIONAL SOFTWARE : LAKE Myriad d'emploi LIFE Electricité de F r a n c e Myriad Instructions for use Allumez votre ordinateur avec une disquette système M S - D O S ( v e r s i o n 3 o u 5) d a n s l e l e c t e u r A . L e l o g i c i e l n e p e u t n e r a v e c la v e r s i o n 4 d e 189 LIFE fonction- MS-DOS. Start u p y o u r c o m p u t e r w i t h M S - D O S system disk in disk drive A . O n c e t h e s y s t e m is l o a d e d , eject t h e s y s t e m d i s k a n d it w i t h t h e Lake Life replace disk. T y p e L A K E , followed by « return » : U n e fois le s y s t è m e c h a r g é , e n l e v e z la d i s q u e t t e s y s t è m e et r e m p l a c e z - l a p a r l a d i s q u e t t e La vie du lac. T a p e z L A C s u i v i d e r e t o u r - c h a r i o t . U n e f o i s l a v e r s i o n d u j e u c h o i s i e et le l o g i c i e l c h a r g é , l a d i s q u e t t e p e u t ê t r e r e t i r é e d u l e c t e u r . La vie du lac p e u t également être copiée dans un répertoire du disque dur. t h e p r o g r a m w i l l l o a d a n d s t a r t u p . A s s o o n as y o u c h o o s e t h e Si v o u s p o s s é d e z u n e s o u r i s , celle-ci d o i t ê t r e i n s t a l l é e a v a n t d e l a n c e r La vie du lac ( s e r e p o r t e r a u m a n u e l d ' i n s t a l l a t i o n d e votre souris). r u n n i n g Lake L e logiciel n é c e s s i t e u n e m é m o i r e d ' a u m o i n s 512 K o p o u r f o n c tionner. Les cartes g r a p h i q u e s Hercules, C G A et E G A sont a u t o matiquement reconnues. g a m e level y o u w a n t a n d t h e disk d r i v e l i g h t goes o f f , e j e c t t h e d i s k a n d p u t it a w a y carefully. I f y o u h a v e a m o u s e a n d w o u l d l i k e t o u s e it, y o u m u s t i n s t a l l it ( f o l l o w t h e i n s t r u c t i o n s i n y o u r u s e r ' s m a n u a l ) Lake Life first before Life. calls for at least 512 K o of m e m o r y . Y o u r g r a p h i c s c a r d i s a u t o m a t i c a l l y d e t e c t e d i f it i s a H e r c u l e s , C G A o r E G A . .