Degeneracy: a link between evolvability, robustness and complexity in biological systems

A full accounting of biological robustness remains elusive; both in terms of the mechanisms by which robustness is achieved and the forces that have caused robustness to grow over evolutionary time. Although its importance to topics such as ecosystem services and resilience is well recognized, the broader relationship between robustness and evolution is only starting to be fully appreciated. A renewed interest in this relationship has been prompted by evidence that mutational robustness can play a positive role in the discovery of adaptive innovations (evolvability) and evidence of an intimate relationship between robustness and complexity in biology. This paper offers a new perspective on the mechanics of evolution and the origins of complexity, robustness, and evolvability. Here we explore the hypothesis that degeneracy, a partial overlap in the functioning of multi-functional components, plays a central role in the evolution and robustness of complex forms. In support of this hypothesis, we present evidence that degeneracy is a fundamental source of robustness, it is intimately tied to multi-scaled complexity, and it establishes conditions that are necessary for system evolvability

Networked buffering: a basic mechanism for distributed robustness in complex adaptive systems

A generic mechanism - networked buffering - is proposed for the generation of robust traits in complex systems. It requires two basic conditions to be satisfied: 1) agents are versatile enough to perform more than one single functional role within a system and 2) agents are degenerate, i.e. there exists partial overlap in the functional capabilities of agents. Given these prerequisites, degenerate systems can readily produce a distributed systemic response to local perturbations. Reciprocally, excess resources related to a single function can indirectly support multiple unrelated functions within a degenerate system. In models of genome:proteome mappings for which localized decision-making and modularity of genetic functions are assumed, we verify that such distributed compensatory effects cause enhanced robustness of system traits. The conditions needed for networked buffering to occur are neither demanding nor rare, supporting the conjecture that degeneracy may fundamentally underpin distributed robustness within several biotic and abiotic systems. For instance, networked buffering offers new insights into systems engineering and planning activities that occur under high uncertainty. It may also help explain recent developments in understanding the origins of resilience within complex ecosystems. \ud \u

Invasions and Extinctions Reshape Coastal Marine Food Webs

The biodiversity of ecosystems worldwide is changing because of species loss due to human-caused extinctions and species gain through intentional and accidental introductions. Here we show that the combined effect of these two processes is altering the trophic structure of food webs in coastal marine systems. This is because most extinctions (∼70%) occur at high trophic levels (top predators and other carnivores), while most invasions are by species from lower trophic levels (70% macroplanktivores, deposit feeders, and detritivores). These opposing changes thus alter the shape of marine food webs from a trophic pyramid capped by a diverse array of predators and consumers to a shorter, squatter configuration dominated by filter feeders and scavengers. The consequences of the simultaneous loss of diversity at top trophic levels and gain at lower trophic levels is largely unknown. However, current research suggests that a better understanding of how such simultaneous changes in diversity can impact ecosystem function will be required to manage coastal ecosystems and forecast future changes

The Effect of Consumers and Mutualists of Vaccinium membranaceum at Mount St. Helens: Dependence on Successional Context

In contrast to secondary succession, studies of terrestrial primary succession largely ignore the role of biotic interactions, other than plant facilitation and competition, despite the expectation that simplified interaction webs and propagule-dependent demographics may amplify the effects of consumers and mutualists. We investigated whether successional context determined the impact of consumers and mutualists by quantifying their effects on reproduction by the shrub Vaccinium membranaceum in primary and secondary successional sites at Mount St. Helens (Washington, USA), and used simulations to explore the effects of these interactions on colonization. Species interactions differed substantially between sites, and the combined effect of consumers and mutualists was much more strongly negative for primary successional plants. Because greater local control of propagule pressure is expected to increase successional rates, we evaluated the role of dispersal in the context of these interactions. Our simulations showed that even a small local seed source greatly increases population growth rates, thereby balancing strong consumer pressure. The prevalence of strong negative interactions in the primary successional site is a reminder that successional communities will not exhibit the distribution of interaction strengths characteristic of stable communities, and suggests the potential utility of modeling succession as the consequence of interaction strengths

Soil resource supply influences faunal size–specific distributions in natural food webs

The large range of body-mass values of soil organisms provides a tool to assess the ecological organization of soil communities. The goal of this paper is to identify graphical and quantitative indicators of soil community composition and ecosystem functioning, and to illustrate their application to real soil food webs. The relationships between log-transformed mass and abundance of soil organisms in 20 Dutch meadows and heathlands were investigated. Using principles of allometry, maximal use can be made of ecological theory to build and explain food webs. The aggregate contribution of small invertebrates such as nematodes to the entire community is high under low soil phosphorus content and causes shifts in the mass–abundance relationships and in the trophic structures. We show for the first time that the average of the trophic link lengths is a reliable predictor for assessing soil fertility responses. Ordered trophic link pairs suggest a self-organizing structure of food webs according to resource availability and can predict environmental shifts in ecologically meaningful ways

Single gene locus changes perturb complex microbial communities as much as apex predator loss

Many bacterial species are highly social, adaptively shaping their local environment through the production of secreted molecules. This can, in turn, alter interaction strengths among species and modify community composition. However, the relative importance of such behaviours in determining the structure of complex communities is unknown. Here we show that single-locus changes affecting biofilm formation phenotypes in Bacillus subtilis modify community structure to the same extent as loss of an apex predator and even to a greater extent than loss of B. subtilis itself. These results, from experimentally manipulated multitrophic microcosm assemblages, demonstrate that bacterial social traits are key modulators of the structure of their communities. Moreover, they show that intraspecific genetic variability can be as important as strong trophic interactions in determining community dynamics. Microevolution may therefore be as important as species extinctions in shaping the response of microbial communities to environmental change

The stability of multitrophic communities under habitat loss

Habitat loss (HL) affects species and their interactions, ultimately altering community dynamics. Yet, a challenge for community ecology is to understand how communities with multiple interaction types—hybrid communities—respond to HL prior to species extinctions. To this end, we develop a model to investigate the response of hybrid terrestrial communities to two types of HL: random and contiguous. Our model reveals changes in stability—temporal variability in population abundances—that are dependent on the spatial configuration of HL. Our findings highlight that habitat area determines the variability of populations via changes in the distribution of species interaction strengths. The divergent responses of communities to random and contiguous HL result from different constraints imposed on individuals’ mobility, impacting diversity and network structure in the random case, and destabilising communities by increasing interaction strength in the contiguous case. Analysis of intermediate HL suggests a gradual transition between the two extreme cases