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

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Systems<br />

A system, as defined by Klir (1969), is imposed<br />

upon an object-a segment of nature<br />

(the earth, a community, a tree, an automobile,<br />

a computer)-by the observer from a distinct<br />

point of view. Everything that does not<br />

belong to the object is the environment . The<br />

boundary between the object and its environment<br />

cannot be clearly defined ; thus th e<br />

delimitation of the object is somewhat arbitrary<br />

and reflects the personal perspectives o f<br />

the modeler . Because we usually cannot stud y<br />

an object in its entirety (because of its complexity),<br />

we observe or measure values of certain<br />

quantities . The choice of quantities to<br />

measure depends on what we consider to b e<br />

of interest or important to the given purpos e<br />

(Klir 1969) .<br />

Most scientists are familiar with the concept<br />

of a system, especially as it is applied i n<br />

thermodynamics. For example, the laws of<br />

thermodynamics were derived by studyin g<br />

idealized closed systems in equilibrium . In<br />

dealing with such closed systems it is possible<br />

to study independent state variables such as<br />

pressure, temperature, and volume, which de -<br />

fine the state of the system. A closed thermodynamic<br />

system can exchange heat and work<br />

but not matter with its environment (Daniels<br />

and Alberty 1967) .<br />

As Ludvig von Bertalanffy (1969) pointe d<br />

out, biological systems are open system s<br />

whose structure is maintained by energy, in -<br />

formation, and matter flow through the systems.<br />

The behavior of the system is the resul t<br />

of the interactions of the elements of th e<br />

system and the flow through the system . Because<br />

the thermodynamic system is closed<br />

and idealized, and because the state variable s<br />

are independent (i .e., one can be varied with -<br />

out affecting another), it is convenient and<br />

correct to describe the system in terms o f<br />

total differentials; for example, the relation<br />

of internal free energy, E, of a system to<br />

temperature and volume, T and V, can be expressed<br />

(Daniels and Alberty 1967) :<br />

E = f(T, V)<br />

dE _<br />

(3T<br />

) V dT + () T dv (1)<br />

Equation 1 implies that the variables E, T,<br />

and V are independent ; one can be varied<br />

while the others are held constant, but mor e<br />

importantly, equation 1 implies summativity<br />

in the system where the behavior of a summa -<br />

tive system is the physical sum of the behavior<br />

of the parts . Because this approach is useful<br />

in thermodynamics and other special<br />

areas, biologists often attempt to explai n<br />

observed phenomena by assuming that the<br />

elements of all systems are independent (vo n<br />

Bertalanffy 1969) . We in plant physiology<br />

have accepted as a standard procedure th e<br />

isolation of a plant in a growth chamber, an d<br />

the study of the response of the plant to on e<br />

factor while holding the others constant . This<br />

approach is epitomized by Cleary (1970) who<br />

derived the following model of photosynthesis :<br />

the n<br />

P = f(M,T,L,N,Pr)<br />

dP= (3P) dM y {('r,) dT<br />

am T,L,N P dT M .L.N.P<br />

+ .<br />

. + dpr~ dP r<br />

1, 7'. L,N<br />

(2 )<br />

where P = photosynthesis, M,T,L,N, and Pr<br />

are moisture, temperature, light, nutrition an d<br />

preconditioning effect, respectively . This<br />

model assumes a summative system analogou s<br />

to a closed thermodynamic system .<br />

This assumption is not totally valid in a<br />

biological system like photosynthesis . Light ,<br />

for example, has an effect not only on the<br />

photochemical reactions within the leaf, but<br />

also is converted to heat, which influences<br />

leaf temperature. Likewise, leaf temperature<br />

is affected by transpiration rates, which is in<br />

turn affected by temperature, moisture status<br />

of the leaf and air, and so on, but equation 2<br />

does not account for these interactions. While<br />

this model is inadequate in that respect ,<br />

Cleary (1970) does recognize that photo -<br />

synthesis is not a simple light-and-temperatur e<br />

related phenomenon but represents a complex<br />

interaction of diverse factors, all of whic h<br />

should be taken into account in order to full y<br />

understand the process . Some of the factor s<br />

listed by Cleary (1970) probably are independent,<br />

namely, Pr and N.<br />

228

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