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PE EIE[R-Rg RESEARCH ON - HJ Andrews Experimental Forest

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which may be of importance in model selection<br />

or approach . Some nonlinear systems of<br />

differential equations are extremely difficult<br />

to solve and their solution may consume a<br />

great deal of computer time . Parameter estimation<br />

is often a significant problem in man y<br />

models. Some models require highly precis e<br />

measurement of input variables, reducin g<br />

their practical applications, but their theoretical<br />

contributions may be important . On the<br />

other hand, we sometimes settle for use of a<br />

linear regression model when the system is<br />

poorly understood and it is much easier t o<br />

measure the variables than to determine their<br />

exact interrelations . Such models are useful in<br />

science, but have some built-in dangers, e .g . ,<br />

the hazard of extrapolation of linear regression<br />

models .<br />

One danger of reliance upon regressio n<br />

models is that the confidence in the model i s<br />

greatest at the mean and diminishes at th e<br />

extremes. In fact, it is possible to fit a curvilinear<br />

function to data of a given range, only<br />

to have the model become totally inadequate<br />

at the extremes .<br />

For example, net photosynthesis as a function<br />

of temperature can be described as a<br />

symmetrical quadratic (Pisek and Winkle r<br />

1958, Webb 1972) (fig. 3) .<br />

values. Thus equation 6 is valid only in th e<br />

range of temperature within which the<br />

parameters were estimated .<br />

The failure of a model to predict syste m<br />

behavior at extremes is not necessarily fatal ;<br />

such models are common in physics, e .g., the<br />

ideal gas law and Newtonian physics . It i s<br />

necessary to understand the limitations of<br />

one 's models to know when the system deviates<br />

from the model .<br />

In choosing a model, it is important t o<br />

understand the assumptions and limitations of<br />

the model . For example, it is questionable to<br />

describe a biological system with a model tha t<br />

assumes a closed reversible system . Th e<br />

assumptions implicit in the model should be<br />

compatible with present knowledge of th e<br />

system of interest .<br />

This is not to say that models of different<br />

systems cannot be used to model another<br />

system. The idea of isomorphism of system s<br />

models is central to general systems theory<br />

(von Bertalanffy 1969) . Thus thermodynami c<br />

models, for example, may be of great utility<br />

in certain biological systems models . It re -<br />

mains for the researcher to be sure that the<br />

systems are isomorphic so that such model s<br />

are applicable .<br />

Pn = X30 + R1 T - 32 T2 ( 6)<br />

But extrapolation of the curve in either direction<br />

will lead to progressively more negativ e<br />

G(T)<br />

Examination of<br />

Some Models<br />

of Photosynthesis<br />

The discussion above dealt with criteria fo r<br />

selection of models . It would be useful to discuss<br />

some models described in the literature .<br />

Leaf Environment Mode l<br />

a<br />

Figure 3 . Net photosynthesis, G(T), as a quadrati c<br />

function of temperature .<br />

T<br />

Botkin (1969) simulated photosynthesis in<br />

an open oak-pine forest near Brookhave n<br />

National Laboratory . He developed a linea r<br />

model of net photosynthesis :<br />

Pn=c 0 + 01 T+ (3 2 ln S<br />

+ Q3 (ln s) 2 + Q4 T2 + (3 5 T 1n S (7)<br />

231

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