The network concept is increasingly used for the description of complex systems. Here we summarize key aspects of the evolvability and robustness of the hierarchical network-set of macromolecules, cells, organisms, and ecosystems. Listing the costs and benefits of cooperation as a necessary behaviour to build this network hierarchy, we outline the major hypothesis of the paper: the emergence of hierarchical complexity needs cooperation leading to the ageing (i.e. gradual deterioration) of the constituent networks. A stable environment develops cooperation leading to over-optimization, and forming an 'always-old' network, which accumulates damage, and dies in an apoptosis-like process. A rapidly changing environment develops competition forming a 'forever-young' network, which may suffer an occasional overperturbation exhausting system-resources, and causing death in a necrosis-like process. Giving a number of examples we demonstrate how cooperation evokes the gradual accumulation of damage typical to ageing. Finally, we show how various forms of cooperation and consequent ageing emerge as key elements in all major steps of evolution from the formation of protocells to the establishment of the globalized, modern human society.
Introduction: Evolvability, robustness and ageing of hierarchical networksThe network approach proved to be a highly efficient tool to describe various levels of the hierarchical organization of complex systems from macromolecules to the currently emerging world-wide social networks. The network description needs the identification of separable subsets of the system as network elements, and a catalogue of their interactions as network contacts or links. In most of the cases network elements themselves can be perceived as networks. Thus, elements of social networks, human individuals are networks of organs and cells, cells are networks of proteins, and proteins are networks of amino acids. Networks display a lot of rather general properties, such as (a) small-worldness, meaning the existence of short pathways between most network elements; (b) the existence of hubs, which have a much higher number of neighbours than the average; (c) a modular structure, which organizes networks to various overlapping groups; (d) the co-existence of strong and weak links, where the link-strength is usually defined as the real, physical strength of the connection, or as the probability of interactions and (e) the existence of a network skeleton, which is the subset of most important pathways in the network. Regretfully, the scope of the current paper does not allow us to give exact definitions and a detailed description of all these network properties, therefore, the reader is referred to a number of recent reviews for more details. (1)(2)(3)(4)