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Quantitative Agrobiology: 111.

The Mitscherlich Equation and Its Constants


0. W. Willcox2
HE criticisms aimed by Black, Kempthorne and White against willcoxs agrabiology in the current issue of Agronomy Journal are directed against three principal targets. First, they undertake to show that Willcoxs derivation of the agrobiologic nitrogen constant 318 is mathematically specious, and that attribution of biological significance to the numbzr 318 i s therefore illusory. As if this were not enough to discredit the so-called agrobiology, they go on with an effort to prove that there is na such thing as an inverse yield-nitrogen law. In the preceding Part I1 hereof it was shown that Black and Kempthornes case against the constant 318 is vitiated by their inattention to the mass action law, well known to chemists, physicists, and agrobiologists who have to deal with dynamic systems that approach equilibria, whereto the Mitscherlich equation or its ,equivalent is applied to evaluate one variable in terms of an associated variable, both being dependent on the same independent variable. In Part I it was shown that their case against the inverse yield-nitrogen law is demolished by their own experiment that was expressly designed to test the validity of this law. But even if their assaults on the constant 318 and on the inverse yield-nitrogen law were admitted to be failures, Black and Kempthorne still have a point which, if it could be made to stick, would ruin the foundation of quantitative agrobiology. This point is that the Mitscherlich equation itself is invalid and that its supposed constants are in fact variables. They open with the remark:
Willcoxs use of the Mitscherlich equation as the base for the derivation of the constant 318 presupposes that the equation is suitable for the purpose. Willcox accepts the Mitscherlich equation as correct. In fact, he elevates it to the level of a Law. tors of plant growth that are available to them. When the supply of these factors is relatively small, the yield will be relatively large; as the quantities of growth factors are increased the yields also increase, but at a continually diminishing rate until finally the plants make no further response to increased application of growth factors. This statement of the law of diminishing increments and of its mode of action is so thoroughly grounded in the experience of farmers through all the ages and in all countries that no responsible authority in the upper ranks of agricultural instruction and research is likely to dispute it.

Origin of the Mitscherlich Equation


What has the Mitscherlich equation to do with the law of diminishing increments ? This equation arose from an attempt to find a mathematical expression for the law of diminishing increments. The deductive approach to such an expression is relatively simple. The known facts are: ( l ) , in all cases, as the quantity of growth factor is increased, the quantity of yield is increased until a limit-yield, designated as A, is reached; and (2), throughouc this process the increments of yield become smaller the nearer the obtained yield approaches A. At any intermediate stage of the operation the actual yield, designated as y, will be smaller than A ; the difference between A and y, that is, A - y, represents the additional quantity of yield that might be obtained if the plants were supplied with additional quantities of growth factor. So the situation at any point is given by A - y; it is around this nuclear expression that a formula for the law of diminishing increments has to be built. Since the increase of yield diminishes at a continually decreasing rate, it is necessary to adjoin another factor that measures the rate of increase, so we write (A-y) c , the quantity c being a factor of proportionality that determines the slope of the yield curve. Now (A-y) c is a function of the quantity of growth factor or combination of growth factors, designated as x, that is acting on the plants, so the situation at any stage is measured by dy/dx = (A - y) c ( 1I which on integration becomes log (A-y) or, if x is measured in baules: log (A-y) =10gA-0.301.~

Here, undoubtedly, is the main issue of the whoIe dispute. Quantitative agrobiology has been put forward as a mathematical-biological discipline that embraces the whole world of rooted and green-leaved plants. If this discipline is found not to apply to all kinds of such plants and all essential factors of plant growth, it could hardly be called a science. It is therefore pertinent to determine whether the Mitscherlich equation is actually the authentic expression of a general law of the kingdom of plants.

= log A - C X

(2)

(3)

LAW OF DIMINISHING INCREMENTS


TO start on a search for a universal law of the plant world, one need look no farther than the law of diminishing increments of yield in agriculture. N o matter how well any kind of plant is watered and fertilized, or how thoroughly it is cultivated, or how adequately it is protected from insects, disease or other abnormal circumstances, in all human experience that plant is incapable of yielding an unlimited amount of vegetable substance from a limited area of soil in one cycle of plant growth. Here, surely, is a law that meets the test of universality. The characteristic of the law of diminishing increments of yield in agriculture is that the quantity of vegetable substance produced by plants is an asymtotic function of the quantities of the facReply to Willcoxs Agrobiology series, pp. 303 to 315, this issue Agronomy Journal. Rec. for publication April 2, 1954. Consulting Agrobiologist, 197 Union St., Ridgewood, N. J.

(The baule of a growth factor is that quantity of it that is necessary and sufficient to produce half of the maximum possible yield A. When x is measured in baules the first baule is found to produce 50% of A, two baules 7533, three baules 87.5%, four baules 93.7S%, and so on in the same diminishing order, each successive b a d e producing half as much as the preceding one until ten baules produce 99.9% of A, which for all practical purposes may be regarded as the limit of possibility). Such is the equation deduced by Mitscherlich, with the help of the mathematician Baule, for the ubiquitous law of diminishing increments of yield in agriculture. There can be no question that this equation takes account of the dominant characteristics of the indisputable law of diminishing increments, namely: the existence of a definite limit of yield fixed by nature for every kind of plant; the approach to this limit by continually decreasing increments of yield ; and, since the equation is logarithmic, the approach to the limit is by the rule of halved increments which excludes all other mathematical, expressions for this natural law. That is to say, any mathematical expression for the law of diminishing increments of yield in agriculture must be an expression of the rule of halved increments. There remains only one essential point not subsumed in the Mitscherlich thesis: the actual values of the constants c. These constants, whether physical or chemical, are specific for each growth factor. There is no theoretical way to ascertain the values of these constants; each has to be determined by experiment, just as the atomic weights of the chemical elements, the gravity

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