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Law of the Minimum Paradoxes - WWW.OLOSCIENCE.COM

Law of the Minimum Paradoxes - WWW.OLOSCIENCE.COM

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Published by Fausto Intilla
Summary. The “law of the minimum” states that growth is controlled by the
scarcest resource (limiting factor) (Justus von Liebig (1840), [1]). This concept was
originally applied to plant or crop growth and quantitatively supported by many experiments.
Some generalizations based on more complicated “dose-response” curves
were proposed. Violations of this law in natural and experimental ecosystems were
also reported. We study models of adaptation in ensembles of similar organisms under
load of environmental factors and prove that violation of the Liebig law follows
from adaptation effects. If the fitness of an organism in fixed environment satisfies
the law of the minimum then adaptation equalizes the pressure of essential factors
and therefore acts against the law. This is the the law of the minimum paradox: if for
a randomly chosen pair “organism–environment” the law of the minimum typically
holds, then, in a well-adapted system, we have to expect violations of this law.
For the opposite interaction of factors (a synergistic system of factors which
amplify each other) adaptation leads from factor equivalence to limitations by a
smaller number of factors.
For analysis of adaptation we develop a system of models based on Selye’s idea
of the universal adaptation resource (adaptation energy). These models predict that
under the load of an environmental factor a population separates into two groups
(phases): a less correlated, well adapted group and a highly correlated group with
a larger variance of attributes, which experiences problems with adaptation. Some
empirical data are presented and some evidences of interdisciplinary applications to
econometrics are discussed.
Summary. The “law of the minimum” states that growth is controlled by the
scarcest resource (limiting factor) (Justus von Liebig (1840), [1]). This concept was
originally applied to plant or crop growth and quantitatively supported by many experiments.
Some generalizations based on more complicated “dose-response” curves
were proposed. Violations of this law in natural and experimental ecosystems were
also reported. We study models of adaptation in ensembles of similar organisms under
load of environmental factors and prove that violation of the Liebig law follows
from adaptation effects. If the fitness of an organism in fixed environment satisfies
the law of the minimum then adaptation equalizes the pressure of essential factors
and therefore acts against the law. This is the the law of the minimum paradox: if for
a randomly chosen pair “organism–environment” the law of the minimum typically
holds, then, in a well-adapted system, we have to expect violations of this law.
For the opposite interaction of factors (a synergistic system of factors which
amplify each other) adaptation leads from factor equivalence to limitations by a
smaller number of factors.
For analysis of adaptation we develop a system of models based on Selye’s idea
of the universal adaptation resource (adaptation energy). These models predict that
under the load of an environmental factor a population separates into two groups
(phases): a less correlated, well adapted group and a highly correlated group with
a larger variance of attributes, which experiences problems with adaptation. Some
empirical data are presented and some evidences of interdisciplinary applications to
econometrics are discussed.

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Published by: Fausto Intilla on Jul 18, 2009
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05/11/2014

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Law of the Minimum Paradoxes
Alexander N. Gorban
1
, Elena V. Smirnova
2
, and Tatiana A. Tyukina
1
1
Centre for Mathematical Modelling, University of Leicester, Leicester, LE1 7RH,UK,
{
ag153,tt51
}
@le.ac.uk
2
Siberian Federal University, Krasnoyarsk, 660041, Russia,
seleval2008@yandex.ru
Summary.
The “law of the minimum” states that growth is controlled by thescarcest resource (limiting factor) (Justus von Liebig (1840), [1]). This concept wasoriginally applied to plant or crop growth and quantitatively supported by many ex-periments. Some generalizations based on more complicated “dose-response” curveswere proposed. Violations of this law in natural and experimental ecosystems werealso reported. We study models of adaptation in ensembles of similar organisms un-der load of environmental factors and prove that violation of the Liebig law followsfrom adaptation effects. If the fitness of an organism in fixed environment satisfiesthe law of the minimum then adaptation equalizes the pressure of essential factorsand therefore acts against the law. This is the
the law of the minimum paradox 
: if fora randomly chosen pair “organism–environment” the law of the minimum typicallyholds, then, in a well-adapted system, we have to expect violations of this law.For the opposite interaction of factors (a synergistic system of factors whichamplify each other) adaptation leads from factor equivalence to limitations by asmaller number of factors.For analysis of adaptation we develop a system of models based on Selye’s ideaof the universal adaptation resource (adaptation energy). These models predict thatunder the load of an environmental factor a population separates into two groups(phases): a less correlated, well adapted group and a highly correlated group witha larger variance of attributes, which experiences problems with adaptation. Someempirical data are presented and some evidences of interdisciplinary applications toeconometrics are discussed.
Keywords: Liebigs Law, Adaptation, Fitness, Stress
Introduction
The “law of the minimum” states that growth is controlled by the scarcestresource (limiting factor) [1]. This concept was originally applied to plant orcrop growth. Many times it was criticized, rejected, and then returned to anddemonstrated quantitative agreement with experiments [1], [2], [3]. Liebig’s
 
2 A.N. Gorban, E.V. Smirnova, T.A. Tyukina
law of the minimum was extended then to more a general conception of fac-tors, not only for the elementary physical description of available chemicalsubstances and energy. Any environmental factor essential for life that is be-low the critical minimum, or that exceeds the maximum tolerable level couldbe considered as a limiting one.The biological generalizations of Liebig’s Law were supported by the bio-chemical idea of limiting reaction steps (see, for example, [4] or recent review[5]). Some of the generalizations of the law went quite far from agricultureand ecology. The law of the minimum was applied to economics [6] and toeducation, for example [7].We consider also the opposite principle organization of factors. This issynergy, the superlinear mutual amplification of factors. Synergy is possiblefor interactions of various poisons, for example.The reaction of an organismto the load of a single factor may have plateaus(intervals of tolerance consideredin Shelford’s “lawof tolerance”,[22], Chapter5). The dose–response curves may be nonmonotonic [8] or even oscillating.Nevertheless, we start from a very simple abstract model that is close tothe usual factor analysis. We consider organisms that are under the influenceof several factors
1
,...F 
q
. Each factor has its intensity
i
(
i
= 1
,...q
). Forconvenience, we consider all these factors as negative or harmful (later, afterwe introduce fitness
, it will mean that all the partial derivatives are non-positive
∂W/∂f 
i
0). This is just a convention about the choice of axesdirections: a wholesome factor is just a “minus harmful” factor.We represent the organisms, which are adapting to stress, as the systemswhich optimize distribution of available amount of resource for neutralizationof different aggressive factors (we consider the deficit of anything needful as anegative factor too). These
factor–resource
models with optimization are veryconvenient for the modeling of adaptation. We use a class of models
many  factors – one resource
.Each organism has its adaptation systems, a “shield” that can decrease theinfluence of external factors. In the simplest case, it means that each systemhas an available adaptation resource,
R
, which can be distributed for theneutralization of factors: instead of factor intensities
i
the system is underpressure from factor values
i
a
i
r
i
(where
a
i
>
0 is the coefficient of efficiencyof factor
i
neutralization by the adaptation system and
r
i
is the share of theadaptation resource assigned for the neutralization of factor
i
,
i
r
i
R
).The zero value
i
a
i
r
i
= 0 is optimal (the fully compensated factor), andfurther compensation is impossible and senseless.Interaction of each system with a factor
i
is described by two quantities:the factor
i
pressure
ψ
i
=
i
a
i
r
i
and the resource
r
i
assigned to the factor
i
neutralization. The first quantity characterizes,how big the uncompensatedharm is from that factor, the second quantity measures, how intensive is theadaptation answer to the factor (or how far the system was modified to answerthe factor
i
pressure).
 
Law of the Minimum Paradoxes 3
For the modeling of adaptation we follow first the classical Selye ideasabout an adaptation resource (adaptation energy [9, 10]). Already one factor–one resource models of adaptation produce the tolerance law and give theseparation of groups of organism into two subgroups: the less correlated well-adapted organisms and highly correlated organisms with a deficit of the adap-tation resource. The variance is also higher in the highly correlated group of organisms with a deficit of the adaptation resource.The crucial question is: what is the
resource of adaptation 
? This questionarose for the first time when Selye published the concept of 
adaptation energy 
and experimental evidence supporting this idea [9, 10]. Selye found that theorganisms (rats) which demonstrate no differences in normal environment maydiffer significantlyin adaptation to an increasingload of environmentalfactors.Moreover, when he repeated the experiments, he found that adaptation abilitydecreases after stress. All the observations could be explained by existence of an universal adaptation resource that is being spent during all adaptationprocesses.Later the concept of adaptation energy was significantly improved [11],plenty of indirect evidence supporting this concept were found, but this elu-sive adaptation energy is still a theoretical concept, and in the modern “En-cyclopedia of Stress” we read: “As for adaptation energy, Selye was never ableto measure it...” [12]. Nevertheless, the notion of adaptation energy is veryuseful in the analysis of adaptation and is now in wide use (see, for example,[13, 14]).The idea of 
exchange
can help in the understanding of adaptation energy:there are many resources, but any resource can be exchanged for another one.To study such an exchange we have to specify, what is the
exchange rate
,how fast this exchange could be done (what is the
exchange time
), what isthe
margin 
, how the margin depends on the exchange time, and what is the
limit of that exchange
. Market economics seems closer to the idea of resourceuniversalization than biology is, but for biology this exchange idea also seemsattractive. Of course there exist some limits on the possible exchanges of dif-ferent resources. It is possible to include the exchange processes into models,but many questions appear immediately about unknown coefficients. Never-theless, we can follow Selye’s arguments and postulate the adaptation energyunder the assumption that this is not an “energy” or Aristotle’s “entelechy”(which definitely had an impact on Selye’s work), but just a pool of vari-ous exchangeable resources. When an organism achieves the limits of resourceexchangeability, the universal non-specific stress and adaptation syndrometransforms (disintegrates) into specific diseases. Near this limit we have toexpect the
critical retardation 
of exchange processes.Adaptation optimizes the state of the system for given available amountsof the adaptation resource. This idea seems very natural, but it may be a dif-ficult task to find the objective function that is hidden behind the adaptationprocess. Nevertheless, even an assumption about the existence of an objec-

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