Stasis is an Inevitable Consequence of Every Successful Evolution

Evolutionary stasis is discussed in light of the idea that the common output of every successful evolution is the creation of the entities that are increasingly resistant to further change. The moving force of evolution is entropy. This general aspiration for chaos is a cause of the mortality of organisms and extinction of species. However, being a prerequisite for any motion, entropy generates (by chance) novelties, which may happen to be (by chance) more resistant to further decay and thus survive. The entities that change rapidly disappear. All existing entities are endowed with an ability to resist further change. In simple organisms, the stasis is primarily achieved by means of the high fidelity of DNA reproduction. In organisms with a large genome and complex development, the achievable fidelity of genome reproduction fails to guarantee homeorhetic reproduction: there is more mutation than reproduction. Such species must be capable of surviving and remain phenotypically unchanged at continuous changes of their genes. This capability (canalization or robustness) reflects a global degeneracy of the link structure-function: there are more genotypes than phenotypes. Hence, function (i.e. meaning), not structure, is selected. The selection for successful ontogenesis in a varying environment creates developmental robustness to mutational and environmental perturbations and, consequently, to the halt of evolution. Evolution is resistance to entropy, the adaptation to environment being only one of the means of this resistance. Everything essential in biology is determined not by physical causality but by semantic rules and goal-directed programs. This principal operates on various levels of biological organization.     

Evolution of altruistic cooperation among nascent multicellular organisms

Cooperation is a classic solution to hostile environments that limit individual survival. In extreme cases this may lead to the evolution of new types of biological individuals (e.g., eusocial super‐organisms). We examined the potential for interindividual cooperation to evolve via experimental evolution, challenging nascent multicellular “snowflake yeast” with an environment in which solitary multicellular clusters experienced low survival. In response, snowflake yeast evolved to form cooperative groups composed of thousands of multicellular clusters that typically survive selection. Group formation occurred through the creation of protein aggregates, only arising in strains with high (>2%) rates of cell death. Nonetheless, it was adaptive and repeatable, although ultimately evolutionarily unstable. Extracellular protein aggregates act as a common good, as they can be exploited by cheats that do not contribute to aggregate production. These results highlight the importance of group formation as a mechanism for surviving environmental stress, and underscore the remarkable ease with which even simple multicellular entities may evolve—and lose—novel social traits.     

A thermodynamic theory of evolution

As a closed thermodynamic system subject to an essentially constant free energy gradient, the biosphere must evolve toward a stationary state of maximum structuring and minimum dissipation with respect to this applied gradient. Since biological evolution occurs opportunistically through chance and selection, rather than as a direct response to the free energy gradient, the conformance of this phase of evolution with thermodynamics requires that natural selection, and the particular adaptive strategies employed by species of organisms, be related to the principles of increasing structuring and decreasing dissipation. In this paper, some general features of this relationship are proposed.     

Local adaptation to biocontrol agents: A multi-objective data-driven optimization model for the evolution of resistance

Spatial and temporal variability in the application of biological control agents such as parasites or pathogenic bacteria can cause the evolution of resistance in pest organisms. Because biocontrol will be more effective if organisms are not resistant, it is desirable to examine the evolution of resistance under different application strategies.We present a computational method that integrates a genetic algorithm with experimental data for predicting when local populations are likely to evolve resistance to biocontrol pathogens. The model incorporates parameters that can be varied as part of pest control measures such as the distribution and severity of the biocontrol agent (e.g., pathogenic fungi). The model predicts the evolution of pathogen defense as well as indirect selection on several aspects of the organism's genetic system. Our results show that both variability of selection within populations as well as mean differences among populations are important in the evolution of defenses against biocontrol pathogens. The mean defense is changed through the pest organism's genotype and the variance is affected by components of the genetic system, namely, the resiliency, recombination rate and number of genes.The data-driven model incorporates experimental data on pathogen susceptibility and the cost of defense. The results suggest that spatial variability rather than uniform application of biological control will limit the evolution of resistance in pest organisms.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

Morphogenetic bases and parametric system of trichome evolution in plants of the genus <Emphasis Type="Italic">Draba</Emphasis> L.

Using quantitative morphological analysis of light microscopy data, the normal variation of trichome morphogenesis is studied in six whitlow grass species (Draba L.) and the morphological variation of adult trichomes in 11 species. The evolution consists in the transition from a radial morphogenesis pattern to bilateral and replacement of complex (branched) trichome rays with simple (unbranched) rays. A parametric system is constructed for classification of the ray morphology; this system includes two parameters—the ratio of the numbers of complex to simple rays, characterizing the probability of secondary branching of primary buds, and the number of primary buds, characterizing the probability of primary branching on the surface of the trichome cell. In this parametric space, all of the studied species fit well a third-order curve consisting of two ascending branches displaying a positive correlation between the primary and secondary branchings and a descending branch, located between them, where the primary and secondary branches are negatively correlated. The deduced evolutionary direction is almost independent of the size of the trichome cells and is explained exclusively by the mechanics of morphogenesis: acceleration in the development of the primary bud of the ray decreases the probability of its own branching and creates additional elastic extension of the cell surface, preventing other buds from branching. The morphogenesis itself appears to be a mechanically nonholonomic system, filtering in a selective manner the fluctuations of the same sign, which explains the directed pattern of its evolution. In the evolutionarily initial state, trichome ontogenesis is absent because its modules (primary buds) are formed by a mirror duplication. The ontogenesis commences when mirror symmetry in the formation of modules is lost and replaced with an axial pattern; thus, the change in the morphological type of buds is a direct consequence of the emergence of ontogenesis and its further evolution. Its main material is intraindividual variation, the only source of which is the mechanics of morphogenesis itself. It is found that morphological evolution can take place at an initially zero heritability and zero adaptive value of morphological differences.     

Compensatory vs. pseudocompensatory evolution in molecular and developmental interactions

The evolution of molecules, developmental circuits, and new species are all characterized by the accumulation of incompatibilities between ancestors and descendants. When specific interactions between components are necessary at any of these levels, this requires compensatory coevolution. Theoretical treatments of compensatory evolution that only consider the endpoints predict that it should be rare because intermediate states are deleterious. However, empirical data suggest that compensatory evolution is common at all levels of molecular interaction. A general solution to this paradox is provided by plausible neutral or nearly neutral intermediates that possess informational redundancy. These intermediates provide an evolutionary path between coadapted allelic combinations. Although they allow incompatible end points to evolve, at no point was a deleterious mutation ever in need of compensation. As a result, what appears to be compensatory evolution may often actually be “pseudocompensatory.” Both theoretical and empirical studies indicate that pseudocompensation can speed the evolution of intergenic incompatibility, especially when driven by adaptation. However, under strong stabilizing selection the rate of pseudocompensatory evolution is still significant. Important examples of this process at work discussed here include the evolution of rRNA secondary structures, intra- and inter-protein interactions, and developmental genetic pathways. Future empirical work in this area should focus on comparing the details of intra- and intergenic interactions in closely related organisms.     

How Cancer Shapes Evolution and How Evolution Shapes Cancer

Evolutionary theories are critical for understanding cancer development at the level of species as well as at the level of cells and tissues, and for developing effective therapies. Animals have evolved potent tumor-suppressive mechanisms to prevent cancer development. These mechanisms were initially necessary for the evolution of multi-cellular organisms and became even more important as animals evolved large bodies and long lives. Indeed, the development and architecture of our tissues were evolutionarily constrained by the need to limit cancer. Cancer development within an individual is also an evolutionary process, which in many respects mirrors species evolution. Species evolve by mutation and selection acting on individuals in a population; tumors evolve by mutation and selection acting on cells in a tissue. The processes of mutation and selection are integral to the evolution of cancer at every step of multistage carcinogenesis, from tumor genesis to metastasis. Factors associated with cancer development, such as aging and carcinogens, have been shown to promote cancer evolution by impacting both mutation and selection processes. While there are therapies that can decimate a cancer cell population, unfortunately cancers can also evolve resistance to these therapies, leading to the resurgence of treatment-refractory disease. Understanding cancer from an evolutionary perspective can allow us to appreciate better why cancers predominantly occur in the elderly and why other conditions, from radiation exposure to smoking, are associated with increased cancers. Importantly, the application of evolutionary theory to cancer should engender new treatment strategies that could better control this dreaded disease.     

Adaptation and plasticity in life-history theory: How to derive predictions.

Why is life paced so differently across as well as within organisms? Can one expect across-species patterns to be repeated within a species too, among individuals? The answer to these questions requires understanding conditions under which reaction norms evolve. We provide an overview of what we believe to be understudied areas of life-history theory, to foster theoretical work and to help deriving predictions for the evolution of human reaction norms. We discuss both why one might expect reaction norms to be aligned with patterns across species, and why that expectation might sometimes fail. It is not impossible for environmental cues to shape life histories in the current generation, but compared with cue-independent genetic adaptation, the adaptive task is now more complex; cues may be unreliable or change in value with time; and parental strategies may differ between situations where offspring have the possibility to disperse to new habitats and situations where environmental conditions remain the same across generations. In that regard, we comment on the value of source-sink theory and on the importance of being specific about the way density regulation affects individual vital rates. We also remind the reader that adaptation does not necessarily optimize population growth rates when conflict between entities (e.g. between the two sexes) is a feature of the adaptive process. All these factors likely play an important role on the evolution of reaction norms, and we argue in favour of their more systematic inclusion in human life-history research.     

Tardigrades in Space Research - Past and Future

To survive exposure to space conditions, organisms should have certain characteristics including a high tolerance for freezing, radiation and desiccation. The organisms with the best chance for survival under such conditions are extremophiles, like some species of Bacteria and Archea, Rotifera, several species of Nematoda, some of the arthropods and Tardigrada (water bears). There is no denying that tardigrades are one of the toughest animals on our planet and are the most unique in the extremophiles group. Tardigrada are very small animals (50 to 2,100 μm in length), and they inhabit great number of Earth environments. Ever since it was proven that tardigrades have high resistance to the different kinds of stress factors associated with cosmic journeys, combined with their relatively complex structure and their relative ease of observation, they have become a perfect model organism for space research. This taxon is now the focus of astrobiologists from around the world. Therefore, this paper presents a short review of the space research performed on tardigrades as well as some considerations for further studies.     

Evolution of herbicide resistance in weeds: vertically transmitted fungal endophytes as genetic entities

The appearance of heritable resistance to herbicides in weeds is an evolutionary process driven by human selection. Assuming that spontaneous and random mutations originate herbicide resistance genes, which are selected by selection pressure imposed by herbicides, is the simplest model to understand how this phenomenon appears and increases in weed populations. However, the rate of herbicide resistance evolution is not only determined by the amount of genetic variation within the populations and the selection pressure exerted by herbicides, but also by factors related to genetics, biology and ecology of weeds. The inheritance of the resistance genes, the mating patterns of the populations, the relative fitness of susceptible and resistant phenotypes and gene flow processes also control the mentioned rate. Many cool season grasses are often infected by fungal symbiotic endophytes (Neotyphodium spp.). These organisms modify the physiology, ecology and reproductive biology of their hosts, conferring greater tolerance to biotic and abiotic stresses, greater competitive ability and the capacity of reducing ecosystem biodiversity. In this work, we present new empirical data and propose new theoretical support on how these microbial symbionts can modulate the evolution of herbicide resistance in weeds. Fungal endophytes are vertically transmitted, and may act as genetic entities altering the evolution of herbicide resistance by reducing herbicide efficacy (delaying effect on evolution). In addition, indirect evidence suggests that fungal endophytes might reduce the fitness penalty associated with the newly arisen resistant phenotypes. The importance and dynamic of these opposite effects is discussed.     

Nonlinear population dynamics: complication in the age structure influences transition-to-chaos scenarios

The work continues a series of studies on the evolution of a natural population of explicitly seasonal organisms. Model analyses have revealed relationships between the duration of ontogenesis and the pattern of temporal dynamics in size of an isolated population (i.e., the structure and dimensionality of the chaotic attractors). For nonlinear models of age-structured population dynamics (under long-lasting ontogenesis), increase in the reproductive potential is shown to result in the chaotic attractors whose structure and dimensionality changes in response to variations in the model parameters. When the ontogenesis becomes longer and more complicated, it does not, "on the average", augment the level of chaos in the attractors observed. There are wide enough regions in the space of the birth and death parameter values that provide for windows in the chaotic dynamics where the total or partial regularization occurs.     

Increases in environmental entropy demand evolution

An application of the entropic theory of perception to evolutionary systems indicates that environmental entropy increases will exert pressures on an organism to adapt. We speculate that the instability caused by such environmental changes will also cause an increase in the mutation rate of organisms leading to an eventual increase in their complexity. Such complexity generation allows organisms to adapt to the more entropic environment. Although we conclude that increases in environmental entropy cause an organism to evolve into a more complex organism, increases in entropy may not be necessary for complexity generationper se.     

Cost of resistance to parasites in digital organisms

Virtually all organisms are attacked by parasites and are therefore expected to evolve resistance against these natural enemies. Parasite resistance is costly in a wide range of organisms, although the generality of such costs has been questioned, especially when resistance is not based on reallocation of resources. Digital organisms are increasingly used to explore aspects of life in general. In the Tierra system, there is a trade-off between resistance against parasites and competitive ability. Because digital organisms are too simple to store resources, the finding that resistance to parasites is costly in digital organisms suggests that costs of parasite resistance can also occur when resistance is not resource based.     

Multilevel selection favors fragmentation modes that maintain cooperative interactions in multispecies communities

Reproduction is one of the requirements for evolution and a defining feature of life. Yet, across the tree of life, organisms reproduce in many different ways. Groups of cells (e.g., multicellular organisms, colonial microbes, or multispecies biofilms) divide by releasing propagules that can be single-celled or multicellular. What conditions determine the number and size of reproductive propagules? In multicellular organisms, existing theory suggests that single-cell propagules prevent the accumulation of deleterious mutations (e.g., cheaters). However, groups of cells, such as biofilms, sometimes contain multiple metabolically interdependent species. This creates a reproductive dilemma: small daughter groups, which prevent the accumulation of cheaters, are also unlikely to contain the species diversity that is required for ecological success. Here, we developed an individual-based, multilevel selection model to investigate how such multi-species groups can resolve this dilemma. By tracking the dynamics of groups of cells that reproduce by fragmenting into smaller groups, we identified fragmentation modes that can maintain cooperative interactions. We systematically varied the fragmentation mode and calculated the maximum mutation rate that communities can withstand before being driven to extinction by the accumulation of cheaters. We find that for groups consisting of a single species, the optimal fragmentation mode consists of releasing single-cell propagules. For multi-species groups we find various optimal strategies. With migration between groups, single-cell propagules are favored. Without migration, larger propagules sizes are optimal; in this case, group-size dependent fissioning rates can prevent the accumulation of cheaters. Our work shows that multi-species groups can evolve reproductive strategies that allow them to maintain cooperative interactions.     

The Rarity of Survival to Old Age Does Not Drive the Evolution of Senescence

The evolution of senescence is often explained by arguing that, in nature, few individuals survive to be old and hence it is evolutionarily unimportant what happens to organisms when they are old. A corollary to this idea is that extrinsically imposed mortality, because it reduces the chance of surviving to be old, favors the evolution of senescence. We show that these ideas, although widespread, are incorrect. Selection leading to senescence does not depend directly on survival to old age, but on the shape of the stable age distribution, and we discuss the implications of this important distinction. We show that the selection gradient on mortality declines with age even in the hypothetical case of zero mortality, when survivorship does not decline. Changing the survivorship function by imposing age independent mortality has no affect on the selection gradients. A similar result exists for optimization models: age independent mortality does not change the optimal result. We propose an alternative, brief explanation for the decline of selection gradients, and hence the evolution of senescence.     

Beyond society: the evolution of organismality

The evolution of organismality is a social process. All organisms originated from groups of simpler units that now show high cooperation among the parts and are nearly free of conflicts. We suggest that this near-unanimous cooperation be taken as the defining trait of organisms. Consistency then requires that we accept some unconventional organisms, including some social insect colonies, some microbial groups and viruses, a few sexual partnerships and a number of mutualistic associations. Whether we call these organisms or not, a major task is to explain such cooperative entities, and our survey suggests that many of the traits commonly used to define organisms are not essential. These non-essential traits include physical contiguity, indivisibility, clonality or high relatedness, development from a single cell, short-term and long-term genetic cotransmission, germ–soma separation and membership in the same species.     

Evolution of freezing susceptibility and freezing tolerance in terrestrial arthropods

Arthropods have evolved various adaptations to survive adverse seasons and it has long been discussed why some arthropods are freezing-susceptible and some are freezing-tolerant. However, which mode of frost resistance came first during the course of evolution? A commonly held opinion is that no choice of strategy has been offered in evolution, because each species of arthropod may have its own evolutionary and natural history, leading to cold-hardiness. Freezing tolerance is more frequent in holometabolous insect orders and partially used by certain vertebrates, like some terrestrially hibernating amphibians and reptiles. Supported by phylogenetic, ontogenetic and ecological arguments, we suggest here that freezing tolerance is more recent than freezing susceptibility in the course of arthropods evolution. In addition, we observe that three basic modes of freezing resistance in insect species exist in the field: (i) permanent or year-round freezing-susceptible species, (ii) alternative or seasonal freezing-susceptible/freezing-tolerant species, (iii) permanent or year-round freezing tolerant species.     

An application of information theory to biological evolution

An evolving population of two alleles which generates another new allele is investigated within the framework of information theory. A communication system model of self-reproducing organisms which reproduce the genetic information by the self-reproduction is proposed and the concept of distortion when the genetic information is reproduced is introduced. What genetic information should be transmitted to the next generation? This question may be answered by so called rate distortion theory. The theory is applied to our model and the exact solution is obtained. Numerical results show that in low distortion region, new allele cannot survive, hence no evolution occurs; in intermediate distortion region, all alleles coexist; and in high distortion region, minor allele cannot survive. Moreover, diversity of the progeny takes its maximum value in the intermediate distortion region. This suggests that there may exist an optimal distortion for a population to evolve.     

Evolution by epigenesis: farewell to Darwinism, neo- and otherwise.

In the last 25 years, criticism of most theories advanced by Darwin and the neo-Darwinians has increased considerably, and so did their defense. Darwinism has become an ideology, while the most significant theories of Darwin were proven unsupportable. The critics advanced other theories instead of 'natural selection' and the survival of the fittest'. 'Saltatory ontogeny' and 'epigenesis' are such new theories proposed to explain how variations in ontogeny and novelties in evolution are created. They are reviewed again in the present essay that also tries to explain how Darwinians, artificially kept dominant in academia and in granting agencies, are preventing their acceptance. Epigenesis, the mechanism of ontogenies, creates in every generation alternative variations in a saltatory way that enable the organisms to survive in the changing environments as either altricial or precocial forms. The constant production of two such forms and their survival in different environments makes it possible, over a sequence of generations, to introduce changes and establish novelties--the true phenomena of evolution. The saltatory units of evolution remain far-from-stable structures capable of self-organization and self-maintenance (autopoiesis).