Towards a Theory of Evolution as Multilevel Learning

We apply the theory of learning to physically renormalizable systems in an attempt to develop a theory of biological evolution, including the origin of life, as multilevel learning. We formulate seven fundamental principles of evolution that appear to be necessary and sufficient to render a universe observable and show that they entail the major features of biological evolution, including replication and natural selection. These principles also follow naturally from the theory of learning. We formulate the theory of evolution using the mathematical framework of neural networks, which provides for detailed analysis of evolutionary phenomena. To demonstrate the potential of the proposed theoretical framework, we derive a generalized version of the Central Dogma of molecular biology by analyzing the flow of information during learning (back-propagation) and predicting (forward-propagation) the environment by evolving organisms. The more complex evolutionary phenomena, such as major transitions in evolution, in particular, the origin of life, have to be analyzed in the thermodynamic limit, which is described in detail in the accompanying paper.

Spontaneous emergence of multicellular heritability

The Major Transitions in evolution include events and processes that result in the emergence of new levels of biological individuality. For collectives to undergo Darwinian evolution, their traits must be heritable, but the emergence of higher-level heritability is poorly understood and has long been considered a stumbling block for nascent evolutionary transitions. A change in the means by which genetic information is utilized and transmitted has been presumed necessary. Using analytical models, synthetic biology, and biologically-informed simulations, we explored the emergence of trait heritability during the evolution of multicellularity. Contrary to existing theory, we show that no additional layer of genetic regulation is necessary for traits of nascent multicellular organisms to become heritable; rather, heritability and the capacity to respond to natural selection on multicellular-level traits can arise ''for free.'' In fact, we find that a key emergent multicellular trait, organism size at reproduction, is usually more heritable than the underlying cell-level trait upon which it is based, given reasonable assumptions.

4. Levels of selection

‘Levels of selection’ examines the levels-of-selection question, which asks whether natural selection acts on individuals, genes, or groups. This question is one of the most fundamental in evolutionary biology, and the subject of much controversy. Traditionally, biologists have mostly been concerned with selection and adaptation at the individual level. But, in theory, there are other possibilities, including selection on sub-individual units such as genes and cells, and on supra-individual units such as groups and colonies. Group selection, altruistic behaviour, kin selection, the gene-centric view of evolution, and the major transitions in evolution are all discussed.

Evolution of multicellularity: cheating done right

For decades Darwinian processes were framed in the form of the Lewontin conditions: reproduction, variation and differential reproductive success were taken to be sufficient and necessary. Since Buss (The evolution of individuality, Princeton University Press, Princeton, 1987) and the work of Maynard Smith and Szathmary (The major transitions in evolution, Oxford University Press, Oxford, 1995) biologists were eager to explain the major transitions from individuals to groups forming new individuals subject to Darwinian mechanisms themselves. Explanations that seek to explain the emergence of a new level of selection, however, cannot employ properties that would already have to exist on that level for selection to take place. Recently, Hammerschmidt et al. (Nature 515:75–79, 2014) provided a ‘bottom-up’ experiment corroborating much of the theoretical work Paul Rainey has done since 2003 on how cheats can play an important role in the emergence of new Darwinian individuals on a multicellular level. The aims of this paper are twofold. First, I argue for a conceptual shift in perspective from seeing cheats as (1) a ‘problem’ that needs to be solved for multi-cellularity to evolve to (2) the very ‘key’ for the evolution of multicellularity. Secondly, I illustrate the consequences of this shift for both theoretical and experimental work, arguing for a more prominent role of ecology and the multi-level selection framework within the debate then they currently occupy.

Role of stellar physics in regulating the critical steps for life

We use the critical step model to study the major transitions in evolution on Earth. We find that a total of five steps represents the most plausible estimate, in agreement with previous studies, and use the fossil record to identify the potential candidates. We apply the model to Earth-analogs around stars of different masses by incorporating the constraints on habitability set by stellar physics including the habitable zone lifetime, availability of ultraviolet radiation for prebiotic chemistry, and atmospheric escape. The critical step model suggests that the habitability of Earth-analogs around M-dwarfs is significantly suppressed. The total number of stars with planets containing detectable biosignatures of microbial life is expected to be highest for K-dwarfs. In contrast, we find that the corresponding value for intelligent life (technosignatures) should be highest for solar-mass stars. Thus, our work may assist in the identification of suitable targets in the search for biosignatures and technosignatures.

The levels of selection and evolution of individuality

In this article I will analyze the transfer of fitness during the major transitions in evolution and its place in the multilevel selection models. The aim of the analysis is to show how social evolution can explain the evolutionary transitions in individuality.

Stress and early experience underlie dominance status and division of labour in a clonal insect

Cooperation and division of labour are fundamental in the ‘major transitions’ in evolution. While the factors regulating cell differentiation in multi-cellular organisms are quite well understood, we are just beginning to unveil the mechanisms underlying individual specialization in cooperative groups of animals. Clonal ants allow the study of which factors influence task allocation without confounding variation in genotype and morphology. Here, we subjected larvae and freshly hatched workers of the clonal ant Platythyrea punctata to different rearing conditions and investigated how these manipulations affected division of labour among pairs of oppositely treated, same-aged clonemates. High rearing temperature, physical stress, injury and malnutrition increased the propensity of individuals to become subordinate foragers rather than dominant reproductives. This is reflected in changed gene regulation: early stages of division of labour were associated with different expression of genes involved in nutrient signalling pathways, metabolism and the phenotypic response to environmental stimuli. Many of these genes appear to be capable of responding to a broad range of stressors. They might link environmental stimuli to behavioural and phenotypic changes and could therefore be more broadly involved in caste differentiation in social insects. Our experiments also shed light on the causes of behavioural variation among genetically identical individuals.

Wieloraka realizacja warunków doboru i jej konsekwencje

Tekst analizuje problem wielorakiej realizacji warunków doboru naturalnego: zmienności, reprodukcji i dziedziczności, przy czym główny nacisk położony zostaje na trzecią z wymienionych cech. Głównym celem prowadzonych analiz jest chęć pokazania, że możliwość wielorakiej realizacji doboru istniałaby nawet wówczas, gdyby pominąć argumenty z zakresu astrobiologii i sztucznego życia i skupić się wyłącznie na analizie genetyki znanych nam form życia. Tekst omawia również konsekwencje powyższej konkluzji, wskazując jak teza o wielorakiej realizacji warunków doboru wpływa na rozważania o takich zagadnieniach, jak: redukcjonizm w biologii, definiowanie życia, badania dotyczące epigenetyki oraz autonomia biologii. Zaznaczone zostaje także jej znaczenie dla badań nad poziomami selekcji i wielkimi przełomami ewolucji (major transitions in evolution).