Abstract
Contemporary systems ecology has long been occupied with mechanical explanations of behavior; however, the physical theory that undergirds such explanations has certain limits. It’s not that the physical force laws are ever violated, but with heterogeneous, irreversible relationships subject to aleatoric influences, physical laws can only constrain, but not determine, outcomes. Such complex systems are better treated in the framework of quantified networks of interrelations. The application of simple information theory to networks reveals that ecosystems cannot achieve maximal efficiency without growing vulnerable to novel disturbances. A degree of inefficient redundancies is always necessary to sustain ecological and social functioning; and, if they are to function reliably, such non-optimal features become requisite in the design of infrastructures like power grids, water distribution networks, traffic patterns, and supply chains. In particular, the postulate that economic market efficiency should never be compromised must be re-examined if society is to remain sustainable. Furthermore, the capability of networks to represent distributed causalities allows one to rationalize behaviors like endogenous selection, centripetal acquisition of resources, and the precedence of indirect mutualism over competition in living systems–all phenomena that challenge conventional evolutionary dogma.
Introduction
Over 55 years ago, the late media sage, Marshall , observed how, whenever a new tool emerged within an endeavor, practitioners tended to use it in the context of previous habits and remained blind for a while to its full potential. His example was IBM, which saw its purpose as the manufacture of business machines. It wasn’t until its leaders realized they were in the business of processing information that the enterprise began to take off.
McLuhan’s example bears analogy to what happened in the scientific literature at the turn of the century, when an interest in networks exploded upon the scene (). Everyone was enamored with the new tool, but the preponderance of research was devoted to interpreting networks in mechanical terms. “What are the mechanisms that give rise to small-world or scale-free networks?,” etc. Few bothered with any possibilities that networks might afford beyond the conventional confines of physics. But there is a multitude of new phenomena and system behaviors that can be illuminated using network analysis. This essay is an attempt to encourage network investigators to step out of the mechanical mindset and entertain an “out-of-the-box” perspective on ecological and social dynamics.
It is rarely pointed out, for example, that physics has shortcomings that render its laws inadequate to determine outcomes in complex and living systems. It is not that such limitations haven’t been known for a long while now. For example, it was demonstrated over a century ago that all the four laws1 of physics can operate on only homogeneous variables (Whitehead and Russell, 1913). That is, physical laws treat only collections of objects that are identical and interchangeable. In order to treat combinations of different objects, one must either ignore distinctions or link dissimilar collections of objects via boundary relationships. Physicist Walter concluded, therefore, that any laws that might govern living systems would have to be qualitatively unlike the fundamental laws of physics.
In addition, physics focuses upon definable objects that are governed by the laws via the notion of fields. Thus, the crux of physics is “objects moving according to immutable laws.” Relationships are implicitly assigned a secondary status, so that connections among objects are assumed to be caused by the objects and not vice-versa.
Both assumptions limit the application of conventional physics toward understanding ecological or social ensembles. The most poignant features of these complex systems are their inherent (and sometimes overwhelming) heterogeneity and the relationships that coalesce the different entities into a functioning whole. To further complicate matters, the relationships among the categories often take the form of a process. A process is a temporal series of irreversible transformations leading from one state to another. The very temporality of processes is incongruent with the reversibility of the fundamental laws of physics. Furthermore, as the heterogeneity of a system increases, the number of possible combinations among the categories grows exponentially, eventually defeating any effort to formulate the closed set of boundary constraints necessary to apply the laws of physics (). It’s not that the fundamental laws of physics are ever violated; it’s that under heterogeneity, irreversibility, and contingency, the laws can at most constrain, but not determine particular system outcomes (Ulanowicz, 2016).
A Fundamentally New Tool
Because physics is demonstrably insufficient to encompass the behaviors of complex living systems, a new metaphysics is required (). A promising tool with which to begin building this reformed worldview is the network. Heterogeneity is virtually definitional to what constitutes a network; and processes, while not the only type of relationship among categories, are nonetheless quite acceptable as connections.
There is yet a third attribute that networks portray that is virtually missing from most problems in physics–indeterminacy. True, stochasticity can be imposed upon a physical problem via a boundary condition or as an attribute of a parameter, but it is not inherent in the governing laws themselves. By contrast, almost all realistic networks exhibit an intrinsic degree of indeterminacy (). The nodes in most networks of interest usually connect to more than one other node. For example, in Figure 1 there are five different compartments that receive output from the Zooplankton (compartment 4). Which node will receive a given organism is not determined beforehand. Therefore, there exists in most cases a proliferation of alternative pathways to get from one specific node to another, just as there is a combination of possible routes to get from one point of a city to another. While some routes may be more efficient than others, the network itself does not determine the one which a particular unit of exchange will transverse. Over time, observation can assign probabilities to each of the possible pathways.
FIGURE 1
Indeterminacy being acknowledged, the network still exerts constraints upon inter-nodal transfers. In directed networks it is often impossible to get from some points to all others. In Figure 1 there are no direct pathways from the Mullet (#17) to the Gold spotted Killifish (#10). As
Quantifying Agonistic Dynamics
Fortunately, information theory provides a convenient method to quantify the degrees of order vs. disorganization in directed networks (
where Tij/T.. is an estimate of the joint probability that a quantum of medium is currently flowing from i to j. Now, H is conventionally referred to as the “entropy” of the system, but this is usually a misnomer, because H represents entropy only when elements i and j are completely independent of each other, which is never the case in social or ecosystems.
Whenever i and j are related to (partially constrained by) one another, H can be decomposed into two non-negative components (Ulanowicz and Norden, 1990). Noting that the logarithm of the quotient of two variables is equal to the difference between their respective logarithms, Equation (1) can be rewritten as,
The magnitude of the terms in braces is not changed by adding and then subtracting the terms logTij and log(T.jTi.), or,
Grouping the first three terms and then the second three yields,
The reason for these particular algebraic manipulations is that the first summation is a non-negative variable in information theory called the “average mutual information,” I, (≥0) (
The average mutual information, I, quantifies the degree of constraint or coordination among all variables i and j. On the other hand, Φ is properly called the conditional entropy and quantifies the lack of constraint [or freedom (to assume different functionalities)] among the elements. Alternatively, it is said to represent the freedom among the i and j. In networks, it also quantifies the redundancy of pathways within the network. In simple terms, it measures the degree on average to which the i and j are independent of each other,
The key to understanding the nature of entropy is in the word “lack,” because entropy represents something that is rarely encountered in physics–an apophasis, or something that does not exist (
Well before physicists discovered networks, there existed a tradition in ecology to represent trophic transfers in ecosystems as networks of flows among species (
In particular, it is informative to focus on the normalized mutual information, a = I/H, which can be referred to as the “degree of order” in the network, or the fraction of its activity that is constrained. This ratio can be considered a surrogate for system efficiency, or the ability of the system to process material. When a is close to one, the system is very streamlined, with most flow concentrated along the most efficient pathways. A high value of Φ/H, by contrast, reveals the presence of many small redundant pathways of lesser efficiency.
If one plots the magnitudes of a for different ecosystems in various habitats, it is somewhat surprising to find that their values cluster around the relatively low figure of 0.40 (Figure 2, Ulanowicz, 2009c). It is evident that ecosystems are performing nowhere near maximal efficiency. That so many less efficient processes should persist counters most thinking in evolutionary biology that only the fittest (most efficient) survive. One notices from Eq. 2b that efficiency and disorder (including redundancy) are mathematically complementary terms–more of one implies less of the other.
FIGURE 2

The degrees of order, a, plotted against corresponding magnitudes of fitness [–a log(a)] for a set of 17 ecosystem flow networks having a number of components, n > 12 (Ulanowicz, 2009c).
The Primacy of Efficiency?
Systems engineers have long recognized that redundancy is a key contributor to system reliability. Ecologists, in turn, have theorized that redundant pathways facilitate alternative supplies and functionalities when major routes have been impacted (
The social ramifications of this alternative view of nature are revolutionary to a society that emphasizes efficiency. Most of what humanity designs is intended to be as efficient as possible, and in so many ways this goal is desirable. But in a complex world consisting of multitudinous processes, too much efficiency can be harmful, if not lead to catastrophe in the face of contingent events. This became abundantly clear during the recent COVID-19 pandemic, when global supply chains that had been honed to emphasize the most efficient and profitable pathways began to fail for lack of alternative connections (
Perhaps the most overarching example of the devotion to efficiency is the field of economics, where the sine qua non of economics is that no activity or constraint should be allowed to hinder the efficient operation of markets. In the financial sector such laissez-faire attitude is considered to be the cause of boom-bust cycles in banking and national economies (
Fortunately, the balance between efficiency and reliability observed in natural systems may provide ways of avoiding or mitigating crises due to overdevelopment. As mentioned above, information theory reveals that a natural balance between the organization (I) of a trophic network and its entropy (Φ) appears to exist in the ratio 40:60 (Ulanowicz, 2009c). If a system is far removed from this balance, simple calculus provides a way of reckoning how much and in which direction each Tij should be changed in order to move the whole system network toward the propitious balance (P1890, ibid.). Knowledge of such prescribed changes could be useful to managers engaged in ecosystem remediation.
This balancing protocol has been adopted by some engineers under the rubric of “biomimicry” whereby, if one has some knowledge of what an ideal balance should be, one can design a degree of redundancy and constraints that would allow the system to “fail-soft,” i.e., not collapse in the wake of major disturbance. Quantitative examples of implementing such reliability have been described in the literature for power grids (
Compound Causalities
Having considered the need for reliability in socio-ecological systems, one is now prompted to ask whether the network perspective might also cast light on the other side of the balance, i.e., on the causal roots of development in those ensembles? The chief advantage of considering patterns of relationships is that they can be examined for indirect propagation of simple binary interactions (Ulanowicz and Puccia, 1990).
Networks can be represented as matrices wherein each element represents the magnitude of the effect of row i upon column j. Multiplying such matrices together provides the magnitudes of all second–order (two-step) interactions, and further multiplications describe higher-order indirect influences.
An intriguing result of the last method was that direct interactions of a given sign (+ or −) can generate additive indirect effects of the opposite sign. Regarding second-order effects, this switch has been common knowledge for centuries (e.g., “The enemy of my enemy is my friend.”). The new method, however, allows one to ascertain the cumulative effect that any component has on any other over all possible pathways between them. Thus for example, when it was applied to a network of trophic interactions in the ecosystem of the Florida Everglades, the negative effect that predation by the American alligator has on several of its direct prey was more than compensated by the positive indirect effects it exerted on them along other pathways (
Some of the most important causal configurations in real systems appear as cycles among component elements. That is, the effect that a compartment has on another can be propagated further along various pathways, some of which might lead back to the original source (
An autocatalytic cycle is a circular configuration of causalities wherein the effects of any member upon the next in the cycle is always positive (Ulanowicz, 2016). For example, consider the simple three-member cycle in Figure 3. If process A facilitates process B, and B catalyzes C, which in turn benefits A, then the activity of A indirectly promotes itself. The same goes for B and C. In general, A, B, and C can be objects, processes or events. While the linkages can be deterministic (mechanical), the most interesting dynamics arise due to the intervention of some type of contingency (
FIGURE 3

A primitive 3-component autocatalytic cycle, where each element benefits the next.
A familiar ecological illustration of this simple triad is the community that establishes itself among the freshwater family of aquatic weeds known as Utricularia (
Indirect causalities comprising autocatalysis, when they interact with contingent events, can give rise to phenomena that are uncharacteristic of mechanical scenarios. For example, they can exert selection pressure upon the participating elements and relationships. Should there happen to be an aleoteric change in the nature of B that happens either to make it more sensitive to A or impart more benefit to C, then that change would receive more benefit from A. Whenever there is some form of memory [either analog (
The Driver of Evolution
Although they are not depicted in the Figure 3, the continuation of all component processes is dependent upon resources taken into the system (
FIGURE 4

Schematic of a six-element autocatalytic cycle, where the enhancement of each element on the next results over time in centripetality, the tendency of the loop to draw ever more resources into its orbit.
The inclusion of indirect causality among heterogeneous networks that interact with contingent events has provided insight into a phenomenon that most investigators simply ignore, lest they be accused of trafficking in the transcendental. It is usually considered to be epi-phenomenal; but centripetality is real, palpable, and perhaps most importantly, primary. Consider, for example, a finite field of resources within which two autocatalytic cycles exist. Both will attract ever more of the resource into their respective orbits until the field starts to be depleted. That is, autocatalysis, comprised of forms of mutual benefaction, gives rise to competition at the next higher level. Effectively, no competition can exist unless it is driven by mutual beneficence at the next level down (
Recognizing that mutual beneficence is primary and competition is instead derivative, inverts the conventions of evolutionary theory. Evolution is not driven ultimately by competition, but rather has its origins in mutuality. Furthermore, the belief that the reversible laws of physics determine all events had led Hume to conclude that one cannot get an “ought” from an “is.” Autocatalysis, however, imparts an underlying directionality to ecological and social dynamics that could lead to an emergent ethos.
Not that all change is determined by positivist agency such as autocatalysis. It is necessary also to remember that the yan of centripetality must be balanced by the yin of dissipation and decay in order for a system to persist (Xu et al., 2018).
A New Metaphysics?
Conceiving of complex, heterogeneous systems in terms of networks broadens the scope of science to encompass more of what is familiar in human experience. Rather than picturing a world of only mechanical behaviors that are separated from the external forces that drive them, network systems can be motivated by endogenous agencies that struggle against, but sometimes can take advantage of, necessary dissipation, degradation, and apophasis. Investigators themselves are no longer required to separate themselves from the systems they are studying, but now become participants in a quasi-engineering endeavor to understand how the world around them develops.
On a more applied note, Gregory
Regarding whole-system ecology, indexes arising from networks are becoming useful tools for the comparison of the status of different ecosystems (Wulff and Ulanowicz, 1989;
Networks provide insights into how different categories interact and how causalities propagate beyond their origins and cooperate in ways that give rise to agency. No longer can concepts such as selfhood and intention (
Statements
Data availability statement
The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author.
Author contributions
RU takes full responsibility for all elements of this article, unless otherwise attributed.
Acknowledgments
The author is grateful to Antonio Bodini for wresting him out of retirement to undertake this plea for a change in scientific metaphysics. Publication Number 5965 of the University of Maryland Center for Environmental Science.
Conflict of interest
The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Footnotes
1.^The four physical forces are strong and weak nuclear, columbic, and gravitation. The three laws of thermodynamics emerged out of engineering, not physics. In the late 19thCentury physicists attempted to “sanitize” thermodynamics by confining it to the realm of statistical mechanics (
2.^For binary, or unweighted networks, all Tij could be assumed to possess equal magnitudes.
3.^Russell did not call the dynamic centripetality, preferring instead the negative connotation associated with “chemical imperialism.”
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Summary
Keywords
autocatalysis, centripetality, contingencies, ecosystems, indeterminacy, information theory, networks, sustainability
Citation
Ulanowicz RE (2021) Socio-Ecological Networks: A Lens That Focuses Beyond Physics. Front. Ecol. Evol. 9:643122. doi: 10.3389/fevo.2021.643122
Received
17 December 2020
Accepted
08 February 2021
Published
26 February 2021
Volume
9 - 2021
Edited by
Ferenc Jordan, Centre for Ecological Research, Hungarian Academy of Science, Hungary
Reviewed by
Marco Ortiz, University of Antofagasta, Chile; Stuart R. Borrett, University of North Carolina at Wilmington, United States
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© 2021 Ulanowicz.
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*Correspondence: Robert E. Ulanowicz, ulan@umces.edu
This article was submitted to Models in Ecology and Evolution, a section of the journal Frontiers in Ecology and Evolution
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