Aquacultural Engineering 5 (1986) 161 - 170

Unstructured Food Webs: a Model for Aquaculture Edward A. Laws

Department of Oceanography, Universityof Hawaii, Honolulu, Hawaii 96822, USA

ABSTRACT Predictions derived from the unstntctured food web model developed by lsaacs (1972, 1973) and later modified by Lange and Hurley (1975) can be derived straightforwardly from a simple box model involving plant biomass, heterotroph biomass, and detritus. Differential equations rather than matrices are used to describe the transfer of materials between these boxes. By recasting these equations, several restrictions present in the original model as well as in the Lange and Hurley reformulation can be relaxed. The model was therefore generalized to allow for (1) a distinction between organic inputs due to plant photosynthesis and to applied feeds, (2) a difference in the turnover rates of the plants, heterotrophs, and detritus, and (3) the effect of harvesting a cash crop. This more general model is used to predict the potential biomass of heterotrophs in a fresh water prawn pond, and to predict the outcome of simple managerial manipulations on the system.

INTRODUCTION In a qualitative sense, our understanding of aquatic food webs has changed dramatically in the past 20 years. Simple linear food chain models have given way to the concept of a complex food web. It is now generally recognized that bacteria and protozoans play a critical role in recycling organics, and that the feeding relationships of many aquatic organisms are such as to preclude assigning particular organisms to particular trophic levels. While some simple systems may be reasonably well described within the context of plantherbivore-carnivore models, a more general theory is needed to describe many aquatic food webs. 161 Aquaculmral Engineering 0144-8609/86/S03.50 - Elsevier Applied Science Publishers Ltd, England, 1986. Printed in Great Britain

162

E . A . Laws

In 1972 John Isaacs proposed the use of a so-called "unstructured food web' model to describe the fluxes of organic material in marine food webs. Isaacs (1973) contrasted his model with the classical structured food web models in which heterotrophs were considered to occupy relatively well-defined positions within a finite sequence of trophic steps. In Isaacs' unstructured food web model, each moiety of food was considered to pass through an infinite number of steps and conversions, and at each such step/conversion was considered to be transformed into living biomass, dead biomass, or non-recoverable material (e.g. CO,). This approach completely avoided the assignment of particular organisms to particular trophic levels. In fact, since a given heterotroph might consist of an assemblage of organic moieties which had passed through a number of transformations ranging from some small integer to infinity, the position of a given organism in Isaacs' food web would in general be spread out over an infinite number of food web elements. Isaacs was able to show that his unstructured food web could account for the distribution of certain trace elements in marine food webs, whereas a simple food chain model could not (Isaacs, 1972). Because of the complex flow of food and energy in many aquaculture ponds, it seems useful to consider Isaacs' unstructured food web model as a means of understanding some of the transformations that occur within these ponds, and for predicting the effects of managerial manipulations on the systems.

THE BASIC M O D E L In his original model Isaacs (1972) assumed that food was converted to living tissue with an efficiency Kl, to irretrievable forms (e.g. CO2) with an efficiency K2, and to non-living but retrievable forms (i.e. detritus) with an efficiency K 3. If these are expressed in fractional form, then Ki + K 2 + K 3 = 1. Figure 1 is a diagrammatic representation of Isaacs' unstructured food web. New food is assumed to be introduced at the point (0, 0). The flow of food is either to the right or down. Flow to the right produces living biomass; flow down produces non-living but retrievable material. A critical assumption of the original model was that the time interval between steps in the matrix was the same for all steps. Later I will show that this assumption can be relaxed. If the system is in steady state and if the biomass at point (0, 0) is unity, then

UnstrucR~redfood webs: a model for aquaculture

LIVING STEPS K1 n ~
n

163

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FOR CONSTRUCTION O F M A r R I X recoverable

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as the ~umber oF ~orhs leodmg to the po,nt,

Fig. I. Isaacs"unstructured food web matrix. In the original version, living biomass in the form of plant biomass was envisioned as being introduced at point (0, 0). Conversions to living biomass correspond to moving to the right. Conversions to detritus correspond to moving down. Each diagonal line passes through points representing biomass which has undergone an equal number of transformations since being introduced into the system. The number of transformations associated with each diagonal line is indicated by the number at both ends of the diagonal. The expressions along the top of the matrix are the total biomass of living organisms along each diagonal in the steady state. The expressions at the left have a similar meaning for the detritus. K/ is the fraction of biomass converted into living biomass at each transformation. K_~ is the fraction of biomass similarly leading to detritus. The number of possible pathways leading to living biomass at a particular lattice point is indicated by a number below and to the right of the lattice point. The number of pathways leading to detritus at a particular lattice point is indicated by a number above and to the left of the lattice point.

164

E. A. Laws

it is straightforward to show that the biomass at (0, 1) is K 1 and the biomass at ( 1, 0) is K 3. Proceeding with the analysis, we note that the diagonal lines in Fig. 1 pass through points, each of which represent biomass which has gone through the same number of steps since being introduced at point (0, 0). With the exception of points lying along the (0, n) and (n, 0) lines, each point in the matrix represents a combination of both living biomass and detritus. For example, living biomass at point ( 1, 1) is produced by a transformation of biomass from point ( 1, 0); detritus at p o i n t ( l , 1) is produced by transforming biomass from point (0, 1). The total number of pathways leading to the production of living biomass and detritus at a ~ven point is easily counted, and the method of calculation is indicated in Fig. 1. If the numbers of pathways leading to detritus and living biomass at point ( n - 1, m) are a and b, respectively, then the number of pathways leading to detritus at point (n, m) is a + b. Similarly, if the numbers of pathways leading to detritus and living biomass at point (n, m - 1) are c and d, respectively, then the number of pathways leading to living biomass at point (n, m) is c + d. Given the assumption of steady state, it is straightforward to calculate the amount of living biomass and detritus at each point in the matrix relative to the amount at point (0, 0) by remembering that the efficiency of living biomass production is K~, and the efficiency of detritus production is K 3. The sum of all the living biomass and of all the detritus along each diagonal line forms a simple geometric series, as indicated in Fig. 1, and one can therefore easily calculate the amount of living biomass and detritus in the system. If one assigns to the biomass at point (0, 0) the symbol M 0, then the total living biomass MI in the system in addition to that contained in M0 is
Mr-

MI) K I - -

K,

(1)

and the total amount of detritus contained in Mo is

Md

in the system in addition to that

Mj =

l~o K 3

K,

(2)

Although Isaacs' model does not require the assignment of particular organisms to particular trophic levels, it is nevertheless possible to

Unstn~cmredfood webs: a model for aquaculture

165

derive expressions for the potential biomass of certain types of organisms, but these expressions provide useful predictive tools only to the extent that such organisms exist in and dominate the real ecosystem. Isaacs (1972) provides the following expressions: Strict Predators (Mp)
Mp = KIMl =

K,

(3)

Particle Feeders (),.Ipf)

mpf = K~(Mj + Mo)=

Omnivores (M,,)

II/IoKI( K2 +

K3)

K,

(4)

MoK~ Mv = KI(M,, + M, + M d ) = - -

K,

(5)

Isaacs (1972) derived additional expressions on the assumption that M 0 consisted entirely of plants, but that assumption clearly need not apply in a pond receiving applied feeds. I will show later how to distinguish between inputs from applied feeds and from photosynthesis. Expressions such as eqns (3)-(5) may prove useful for predicting the effect of various managerial manipulations on the system. A potentially revealing application of Isaacs' equations is the study of elemental ratios in organisms. Let us assume for example that the coefficients K~, K2, and g 3 apply to carbon C, and that the coefficients K I, K~, and K; apply to some other element, E. Then according to eqn (5) the ratio E/C in the omnivores is related to the ratio E/C in M0 by the expression KIK2/(K;K~). Similar expressions can be derived for E/C ratios in other groups of organisms. Subsequent to Isaacs' two publications on the unstructured food web model, Lange and Hurley (1975) published a paper in which they showed that the model could be generalized and the mathematics simplified by the use of an alternative matrix formulation. They generalized the model by dropping the condition that all conversions to living biomass and all conversions to detritus occur with the same efficiency. They allowed for a total of nine separate conversion factors

166

E. A. Laws
TABLE 1

Definition of the Nine Conversion Factors Used in the Lange and Hurley (1975) Model
Symbol
K I

Meaning

K, K~ Ka Ks Ks K7 Ks K,~

Conversion factor from source to living Conversion factor from source to dead retrievable Conversion factor from source to irretrievable Conversion factor from living to living Conversion factor from livingto dead retrievable Conversion factor from livingto irretrievable Conversion factor from dead to living Conversion factor from dead to dead retrievable Conversion factor from dead to irretrievable

as indicated in Table 1. T h e y defined W~ as the amount of biomass in the plants, W, as the a m o u n t of biomass in the heterotrophs and W3 as the a m o u n t of detritus. T h e matrix representation of their model then takes the form 1 AI7V= Kt K2 0 K4 K5 0 K7 Ks
WI

Wl
= K, W I + K ~ W . _ + K T W 3 K2 WI + Ks IV,_ + Ks W3

(6)

T h e matrix A operating on the column vector W from the left projects the values of W~, W2, and W3 from time, t, to time, t + A t . Lange and Hurley (1975) required that K t + K 2 + K 3 = K4 + K5 + K6 = K7 + K8 + K9 = 1. T h e s e conditions in effect require that the turnover rates of Wt, W2, and W3 be identical, a condition similar to Isaacs' (1972) requirement that the time intervals between all steps in the food web matrix be the same. Using the Lange and Hurley (1975) approach, it is straightforward to derive many relationships similar to those obtained by Isaacs (1972, 1973), but with nine transfer coefficients rather than three. For example, Lange and Hurley (1975) show that the steady-state potential biomass of strict predators is given by the expression
- K 4 K I ( K 8 - 1 ) + K4K,_K7 (Ka - 1 )(Ks - 1 ) - Ks K7

Mp = Mo

(7)

Unstr~wtured food webs." a model for aquaculture

167

This model seems potentially useful for studying food webs in aquaculture, but I feel several modifications in the model would be desirable. First, it would be useful to distinguish b e ~ ' e e n organic inputs due to plant photosynthesis and organic inputs due to applied feeds. Second, one would like to relax the restriction that the turnover rates of W l, ~ , and B'~ be identical. Finally, one should allow for the effect of harvesting the cash crop. Actually the unstructured food web described with the use of matrLx equations by Lange and Hurley (1975) can be equally well described with the use of differential equations, and in order to generalize the model I propose the following equations
dW~ dt d }~q dt 1 r 1 r

(M,-(C,

+ C, + (73) W,)

s)
(9)

-=-(CaW2+C,W~+C, W3-(C,+Cs+C6) W:)

1

dW~

- = - ( M:, + G W, + C2 w~ + C5 ~ - ( C7 + G + C,) W; ) dt r

(10)

Here M~/r and M~/r are the rates at which plant biomass and feed (non-living but retrievable organic biomass) are introduced into the system via photosynthesis and feed applications, respectively. The Ci are defined in Table 2, and Wt, W_~,and I~ are as previously defined. Note from Table 2 that there is no constraint on the C, except that they be positive numbers. Now let

Ki = Ci/( C, + C, + C3)
m, = M ~ / ( G + C~ + C3)
m 3 = M 3 / ( C I + C, + (73)

(1!) (12)
(13)

T = r/( CI + C, + C3)

(14)

Equations (8)-(10) now become d Wt

dt

1 = 7..(rn, - ( K , + K z + K:,) WI)

1

(15)

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E. ,4. Laws
TABLE 2

Expressions which Define the C~

Expression

31eaning
Fraction of plant biomass converted to heterotroph biomass per unit time Fraction of plant biomass converted to detritus per unit time Fraction of plant biomass converted to irretrievable form per unit time Fraction of heterotroph biomass converted to heterotroph biomass per unit time Fraction of heterotroph biomass converted to detritus per unit time Fraction of heterotroph biomass converted to irretrievable form per unit time Fraction of detritus converted to heterotroph biomass per unit time Fraction of detritus converted to detritus per unit time Fraction of detritus converted to irretrievable form per unit time

Ct/r
C2/r C~/r CJ r

C~/r
C./r
C7/r

c,/r C~ r

dW, dt
dW3 _

1 T (K~W'+K~W~+KvW'-(K~+KS+K~)W)

(16)

dt

(tn3+KsW3+K2W~+K~W'-(KT+K~+Kg)

W3)

(17)

It is obvious from the definition of the K~ that K~ + K , + K 3 -- 1, and that m ~ is the a m o u n t of new plant biomass introduced into the system in time T. H e n c e rnt has the same meaning as Isaacs' M,, and K~, K2, and K 3 have meanings identical to the same symbols as defined by Lange and H u r l e y (1975). However, there is no reason to assume that K 4 + K s + K 6 = K 7 + K s + K 9 = 1 unless one believes that the biomasses of h e t e r o t r o p h s and of detritus are turning over at the same rate as the plant biomass. A c c o r d i n g to the model the t u r n o v e r times of the plants (rp), h e t e r o t r o p h s (rh) , and detritus (rd) are given by the following expressions rp = T (18)

Unstructured food webs: a model for aquaculture

169

rh = T/( K~ + K5 + K~ )
r d =

(19) (20)

T/(K7 + Ks + Ks)

Note that the parameter K~ may be considered to include the effects of both respiration and harvesting cash crops, since both processes transfer living biomass to an irretrievable form. The steady-state solutions to eqns (15)-(17) are as follows:
W I = rnl (21 )

W. = KIrtl'(Kv + Ko)+ Kv(K'-tn'

+m3)

(22)
(23)

KsK,~ + K6(K7 + K,,) W3= (Ks + K6)(K2ml + m3)+ KtKsm~ KsK~ + K6(K7+ Kg)

The potential biomass of various trophic levels can now be calculated in a manner similar to that employed by Isaacs (1972) and Lange and Hurley (1975). For example, the potential biomass of strict herbivores (mh)iS
rnh = K~ rn,/( K4 + K5 + K~)

(24)

The potential biomass of omnivores (my), which are assumed to feed on plants, living organisms, and detritus, is equal to the right-hand side of eqn (22); the potential biomass of strict predators (rnp) is equal to K4 Wz/(K~ + K5 + K6), where W, is given by eqn (22); the potential biomass of detrital feeders is equal to K7 W3/(K4 + Ks + Kc,), where W3 is given by eqn (23); and so forth. Table 3 is a simple application of eqns (21 )-(23) to a prawn pond in Hawaii. Carbon loading rates associated with photosynthesis and applied feeds as well as the turnover time of the phytoplankton are known or could be calculated from data in Costa-Pierce (1984). The turnover times of the heterotrophs and detritus were simply assumed. The values of K j, K2, and K 3 appear to be reasonable based on Isaacs' (1972) calculations. The values of K 4 - K 9 were chosen to have the same relative magnitude as K j, K 2, and K~ while satisfying eqns (19)-(20). Equations (22) and (23) then reduce to two simple linear equations involving rn~ and rn 3. The calculated biomass of heterotrophs in terms of carbon is 138 kg ha-~ compared to the known

170

E. A. Laws
TABLE 3

Application of Unstructured Food Web Model to Prawn Ponds; Biomass Expressed in Terms of Organic Carbon m~ = 22.5 kg ha- ~(Costa-Pierce, 1984, p. 58) rp = 3 days m 3= 12 kg ha- t (40 kg ha- ~day- l x 10% carbon x 3 days; Costa Pierce, 1984, p. 90) r h= 30 days r d = 60 days K l =0.2 K, =0-3 K3 =0.5 Ka = 0-02 K.~= 0-03 Ka=0.05 Kr =0-01 Ks = 0"015 K, = 0"025 W,=4rn~ +4m 3= 138 kgha~,"~= 12m I + 32m 3= 654 kg ha- 1 Standing crop of prawns= 112 kg ha -~ (900 kg ha -~ x 25% organic content x 50%c) Harvesting rate = 0.69 kg ha- ~day- ~= 1.9% of prawn standing crop every three days

s t a n d i n g c r o p of p r a w n c a r b o n of 1 12 kg ha -~. N o t e that the actual p r a w n h a r v e s t i n g rate of a b o u t 1.9% e v e r y t h r e e d a y s is a b o u t 4 0 % of the a s s u m e d value of K6. If the p r a w n h a r v e s t i n g rate w e r e increased, K 6 w o u l d also increase a n d the t u r n o v e r time of the p r a w n s at s t e a d y state w o u l d d e c r e a s e . T h e m o d e l o b v i o u s l y p r o v i d e s a m e c h a n i s m for p r e d i c t i n g the effects of m a n a g e r i a l m a n i p u l a t i o n s o n the p r o d u c t i v i t y of the system. For e x a m p l e , r e d u c i n g the rate of feed applications by a factor of two is p r e d i c t e d to r e d u c e the h e t e r o t r o p h s t a n d i n g c r o p by only 17%.

REFERENCES Costa-Pierce, B. (1984). Ecological studies of semi-intensive prawn aquaculture ponds. PhD dissertation, University of Hawaii, 208 pp. Isaacs, J. D. (1972). Unstructured marine food webs and 'pollutant analogues'. Fish. Bull., 70, 1053-9. Isaacs, J. D. (1973). Potential trophic biomasses and trace-substance concentration in unstructured marine food webs. Mar. Biol., 22, 97-104. Lange, G. D. & Hurley, A. C. (1975). A theoretical treatment of unstructured food webs. Fish. Bull., 7 3 , 3 7 8 - 8 1 .