A Universal Definition of Life: Autonomy and Open-Ended Evolution

Life is a complex phenomenon that not only requires individual self-producing and self-sustaining systems but also a historical-collective organization of those individual systems, which brings about characteristic evolutionary dynamics. On these lines, we propose to define universally living beings as autonomous systems with open-ended evolution capacities, and we claim that all such systems must have a semi-permeable active boundary (membrane), an energy transduction apparatus (set of energy currencies) and, at least, two types of functionally interdependent macromolecular components (catalysts and records). The latter is required to articulate a 'phenotype-genotype' decoupling that leads to a scenario where the global network of autonomous systems allows for an open-ended increase in the complexity of the individual agents. Thus, the basic-individual organization of biological systems depends critically on being instructed by patterns (informational records) whose generation and reliable transmission cannot be explained but take into account the complete historical network of relationships among those systems. We conclude that a proper definition of life should consider both levels, individual and collective: living systems cannot be fully constituted without being part of the evolutionary process of a whole ecosystem. Finally, we also discuss a few practical implications of the definition for different programs of research.     

The Evolution of Life Histories in Holo-anhydrobiotic Animals: A First Approach

The life histories of holo-anhydrobiotic animals differ fromthose of all other organisms by a regular or irregular entranceinto an ametabolic state induced by desiccation. Such ametabolicperiods will arrest growth and reproduction completely and thusaffect primary life history parameters dramatically. The selectiveforces and the genetic and physiological trade-offs acting onanhydrobiotic animals are to a large extent unknown. Assuminglow growth rates and low juvenile to adult survival, generaltheoretical models on life history responses to stress predictthat anhydrobiotic animals will be selected for a high degreeof iteroparity, with low fecundity, large egg size, and lowtotal reproductive investment. A high degree of variabilityin growth and reproduction should create a selective force inthe same direction. Although basic empirical data on life historyparameters are very scarce, available observations seem to beconsistent with this prediction.     

Engine of Life and Big Bang of Evolution: A Personal Perspective

Photosystem II (PS II) is the engine for essentially all life on our planet and its beginning 2.5 billion years ago was the 'big bang of evolution.' It produces reducing equivalents for making organic compounds on an enormous scale and at the same time provides us with an oxygenic atmosphere and protection against UV radiation (in the form of the ozone layer). In 1967, when I began my career in photosynthesis research, little was known about PS II. The Z-scheme had been formulated [Hill and Bendall (1960) Nature 186: 136–137] and Boardman and Anderson [(1964) Nature 203: 166–167] had isolated PS II as a discrete biochemical entity. PS II was known not only to be the source of oxygen but of variable chlorophyll fluorescence [Duysens and Sweers (1963) In: Studies on Microalgae and Photosynthetic Bacteria, pp. 353–372. University of Tokyo Press, Tokyo] and delayed chlorophyll fluorescence [Arnold and Davidson (1954) J Gen Physiol 37: 677–684]. P680 had just been discovered [Döring et al. (1967) Z Naturforsch 22b: 639–644]. No wonder the 'black box of PS II' was described at that time by Bessel Kok and George Cheniae [Current Topics in Bioenergetics 1: 1–47 (1966)] as the 'inner sanctum of photosynthesis.' What a change in our level of understanding of PS II since then! The contributions of many talented scientists have unraveled the mechanisms and structural basis of PS II function and we are now very close to revealing the molecular details of the remarkable and thermodynamically demanding reaction which it catalyzes, namely the splitting of water into its elemental constituents. It has been a privilege to be involved in this journey.     

Evolution of Rotifer Life Histories

When compared to most other multicellular animals, rotifers are all relatively small, short-lived and fast-reproducing organisms. However among and within different rotifer species there is a large variation in life history patterns. This review accounts for such variation in rotifers, with a strong focus on monogonont rotifers. As the life cycle of monogonont rotifers involves both asexual and sexual reproduction, life history patterns can be examined on the level of the genetic individual, which includes all asexual females, sexual females and males that originated from one resting egg. This concept has been applied successfully in many areas, for example in predicting optimal levels of mictic reproduction or sex allocation theory. The benefits and implications of the view of the genetic individual are discussed in detail. Rotifer life histories can also be viewed on the level of physiological individuals. A large part of this review deals with the life histories of individual amictic females and addresses life history traits like body size, egg size and resource allocation patterns. It asks which trade-offs exist among those traits, how these traits change under the influence of environmental factors like food availability or temperature, and whether these changes can be interpreted as adaptive.     

The Origin of Life: A Problem of History,Chemistry, and Evolution

The origin of life is a field full of controversies, not only because of our vague understanding concerning the relevant issues, but also, perhaps more often, owing to our dim conceptual framework throughout the whole field. To improve this situation, an in‐depth conceptual dissection is presented here. It is elucidated that, at its core, the origin of life has three aspects. The facts involved in the process are taken as the historical aspect, which is destined to be uncertain and often irrelevant to debate regarding details. The rules involved include two distinct aspects: chemical mechanisms operated in the whole process, while evolutionary mechanisms joined in only after the emergence of the first Darwinian entities – and then accounted for the subsequent buildup of complexity (this cannot be explained solely by natural selection). Basically, we can ask about the possibility of any assumed event in the origin of life: ‘Is it evolutionarily plausible, chemically feasible, and historically likely?’ Clues from any of the three aspects may be quite valuable in directing our explorations on the other two. This conceptual dissection provides a clearer context for the field, which may even be more useful than any sort of specific research.     

Lemur Biorhythms and Life History Evolution

Skeletal histology supports the hypothesis that primate life histories are regulated by a neuroendocrine rhythm, the Havers-Halberg Oscillation (HHO). Interestingly, subfossil lemurs are outliers in HHO scaling relationships that have been discovered for haplorhine primates and other mammals. We present new data to determine whether these species represent the general lemur or strepsirrhine condition and to inform models about neuroendocrine-mediated life history evolution. We gathered the largest sample to date of HHO data from histological sections of primate teeth (including the subfossil lemurs) to assess the relationship of these chronobiological measures with life history-related variables including body mass, brain size, age at first female reproduction, and activity level. For anthropoids, these variables show strong correlations with HHO conforming to predictions, though body mass and endocranial volume are strongly correlated with HHO periodicity in this group. However, lemurs (possibly excepting Daubentonia) do not follow this pattern and show markedly less variability in HHO periodicity and lower correlation coefficients and slopes. Moreover, body mass is uncorrelated, and brain size and activity levels are more strongly correlated with HHO periodicity in these animals. We argue that lemurs evolved this pattern due to selection for risk-averse life histories driven by the unpredictability of the environment in Madagascar. These results reinforce the idea that HHO influences life history evolution differently in response to specific ecological selection regimes.     

Life History Evolution in Amphicarpic Plants

Abstract Plants with dimorphic flowers or seeds provide excellent material for the study of life history evolution because the dimorphism often involves measurable differences in morphology, size, number, or genetic relatedness. For amphicarpic plants, the proportion of aerial: subterranean morphs produced is highly variable (from 0 to > 100) and related to both environmental and genetic factors. Plants from aerial seeds produce lower ratios of aerial: subterranean morphs than those from subterranean seeds. Despite substantial variation, subterranean seeds are generally heavier than aerial seeds (but fewer) and produce vigorous seedlings with high survivorship and high fitness. Adaptive advantages of subterranean seeds include retention of offspring in favorable parental microhabitats, protection of seeds from herbivory, predation, or fire, and avoidance of desiccating conditions on the soil surface; potential disadvantages include lack of gene exchange, high energy costs, limited dispersal, and sibling competition. For the few species studied, aerial reproduction is more plastic than subterranean reproduction and more likely to be affected by environmental conditions. Quantitative genetic analyses of a population of the annual grass Amphicarpum purshii have revealed lower heritabilites for subterranean relative to aerial reproductive traits. Subterranean seed number and mass show genetic correlations with shoot mass while aerial seed number and mass do not; seed set percentages of both seed types as well as percentage allocation to both reproductive morphs show negative genetic correlations with shoot mass. In this Amphicarpum population, directional selection on shoot mass may indirectly select for increased subterranean (but not aerial) seed output.