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At least 19 recordsLinked to original sources

Evolvability and genetic constraint in Dalechampia blossoms: genetic correlations and conditional evolvability.

The short-term evolvability of a character is closely related to its level of additive genetic variation. However, a large component of the variation in any one character may be pleiotropically linked to other characters under the influence of different selective factors. Therefore, the organization of the organism into quasi-independent modules may be an important prerequisite for evolvability. In this paper we propose to study character evolvability in terms of conditional genetic variation. By estimating the amount of genetic variation in a character, y, that is independent of other characters, x, we can assess the evolvability of y when there is stabilizing selection on x. We suggest that systematic use of conditioning may help build a picture of modular organization and quasi-independent evolvability. As an illustration, we use this approach to assess the evolvability of floral characters in Dalechampia scandens (Euphorbiaceae). Although our study population had relatively low levels of genetic variation at the outset, we find evidence that conditioning may lead to substantial further reduction in the genetic variation available for independent adaptation. This provides additional evidence that the D. scandens blossom is constrained in its short-term evolvability.

Biological Evolution↗

Is modularity necessary for evolvability? Remarks on the relationship between pleiotropy and evolvability.

Evolvability is the ability to respond to a selective challenge. This requires the capacity to produce the right kind of variation for selection to act upon. To understand evolvability we therefore need to understand the variational properties of biological organisms. Modularity is a variational property, which has been linked to evolvability. If different characters are able to vary independently, selection will be able to optimize each character separately without interference. But although modularity seems like a good design principle for an evolvable organism, it does not therefore follow that it is the only design that can achieve evolvability. In this essay I analyze the effects of modularity and, more generally, pleiotropy on evolvability. Although, pleiotropy causes interference between the adaptation of different characters, it also increases the variational potential of those characters. The most evolvable genetic architectures may often be those with an intermediate level of integration among characters, and in particular those where pleiotropic effects are variable and able to compensate for each other's constraints.

Adaptation, Psychological↗

Evolvability and genetic constraint in Dalechampia blossoms: components of variance and measures of evolvability.

Many evolutionary arguments are based on the assumption that quantitative characters are highly evolvable entities that can be rapidly moulded by changing selection pressures. The empirical evaluation of this assumption depends on having an operational measure of evolvability that reflects the ability of a trait to respond to a given external selection pressure. We suggest short-term evolvability be measured as expected proportional response in a trait to a unit strength of directional selection, where strength of selection is defined independently of character variation and in units of the strength of selection on fitness itself. We show that the additive genetic variance scaled by the square of the trait mean, IA, is such a measure. The heritability, h2, does not measure evolvability in this sense. Based on a diallel analysis, we use IA to assess the evolvability of floral characters in a population of the neotropical vine Dalechampia scandens (Euphorbiaceae). Although we are able to demonstrate that there is additive genetic variation in a number of floral traits, we also find that most of the traits are not expected to change by more than a fraction of a percent per generation. We provide evidence that the degree of among-population divergence of traits is related to their predicted evolvabilities, but not to their heritabilities.

Biological Evolution↗

Evolve II: a computer model of an evolving ecosystem.

The Evolve II program is a model of an ecosystem in which organisms are allowed to evolve. Organisms are subject to a changeable environment and competition from other organisms for a limited food supply. The gene structure may change through mutation. A feature of Evolve II is that the magnitude of phenotypic change resulting from mutation is itself a property of the gene. The system was studied under a number of environmental variation schemes. We report three significant findings. Two species (lineages with distinctly different survival strategies) evolved and coexisted in the same environmental conditions. Organisms developed a resistance to phenotypic change in response to mutation in slowly varying environments. However, traits which favor survival of the individual at the expense of reproduction could in some cases undergo phenotypic change in response to mutation despite the fact that this did not favor the survival of the offspring. This demonstrates that gene structures can evolve which are advantageous from the standpoint of the lineage, but not advantageous from the standpoint of individual offspring.

Aging↗

Recent advances in evolvable systems--ICES 96 (International Conference on Evolvable Systems).

This paper reviews the developments in evolvable hardware systems presented at the First International Conference on Evolvable Systems (ICES 96). The main body of the review gives an overview of the 34 papers presented orally, splitting them into three broad groups according to whether they involve (1) evolving a fit solution to a problem as a member of a population of competing candidates, (2) evolving solutions that can individually learn from and adapt to their environments, or (3) the embryonic growth of solutions. We also review the discussion sessions of the conference and give pointers to related upcoming events.

Algorithms↗

Catalytic consequences of experimental evolution: catalysis by a 'third-generation' evolvant of the second beta-galactosidase of Escherichia coli, ebgabcde, and by ebgabcd, a 'second-generation' evolvant containing two supposedly 'kinetically silent' mutations.

The kinetics of hydrolysis of a series of synthetic substrates by two experimentally evolved forms ('evolvants'), ebgabcd and ebgabcde, of the second beta-galactosidase of Escherichia coli have been measured. The ebgabcd enzyme differs from the wild-type (ebgo) enzyme by Asp92-->Asn (a) and Trp977-->Cys (b) changes in the large subunit, as well as two changes hitherto considered to have no kinetic effect, Ser979-->Gly in the large subunit (c) and Glu122-->Gly in the small subunit (d). The enzyme ebgabcde contains in addition a Glu93-->Lys change in the large subunit (e). Comparison of ebgabcd with ebgab [Elliott, K, Sinnott, Smith, Bommuswamy, Guo, Hall and Zhang (1992) Biochem. J. 282, 155-164] indicates that the c and d changes in fact accelerate the hydrolysis of the glycosyl-enzyme intermediate by a factor of 2.5, and also decrease the charge on the aglycone oxygen atom at the first transition state; the charge on the glycone, however, is unaltered [see K, Konstantinidis, Sinnott and Hall (1993) Biochem. J. 291, 15-17]. The e mutation causes a fall in the degalactosylation rate of about a factor of 3, and its occurrence only together with c and d mutations [Hall, Betts and Wootton (1989) Genetics 123, 635-648] suggests that degalactosylation of a hypothetical ebgabe enzyme would be so slow that the enzyme would have no biological advantage over the ancestral ebgab. The transfer products from galactosyl-ebgabcd and galactosyl-ebgabcde to high concentrations to glucose have been measured; the predominant product is allolactose, but significant quantities of lactose are also formed; however, at apparent kinetic saturation of the galactosyl-enzyme, hydrolysis rather than transfer is the preponderant pathway. A knowledge of the rates of enzyme-catalysed exchange of 18O from [1-18O]galactose to water permits the construction of the free-energy profiles for hydrolysis of lactose by begabcd and ebgabcde. As with the other evolvants, changes in the profile away from the rate-determining transition state are essentially random, and there is no correlation between the changes in the free energies of intermediates and of their flanking transition states. We consider the aggregate of our kinetic data on the ebg system to be telling experimental support for the theoretical objections of Pettersson [Pettersson (1992) Eur. J. Biochem. 206, 289-295 and previous papers] to the Albery-Knowles theory of the evolution of enzyme kinetic activity.

Catalysis↗

Early-branching or fast-evolving eukaryotes? An answer based on slowly evolving positions.

The current paradigm of eukaryotic evolution is based primarily on comparative analysis of ribosomal RNA sequences. It shows several early-emerging lineages, mostly amitochondriate, which might be living relics of a progressive assembly of the eukaryotic cell. However, the analysis of slow-evolving positions, carried out with the newly developed slow-fast method, reveals that these lineages are, in terms of nucleotide substitution, fast-evolving ones, misplaced at the base of the tree by a long branch attraction artefact. Since the fast-evolving groups are not always the same, depending on which macromolecule is used as a marker, this explains most of the observed incongruent phylogenies. The current paradigm of eukaryotic evolution thus has to be seriously re-examined as the eukaryotic phylogeny is presently best summarized by a multifurcation. This is consistent with the Big Bang hypothesis that all extant eukaryotic lineages are the result of multiple cladogeneses within a relatively brief period, although insufficiency of data is also a possible explanation for the lack of resolution. For further resolution, rare evolutionary events such as shared insertions and/or deletions or gene fusions might be helpful.

Actins↗

Beta-lactamase inhibitors: evolving compounds for evolving resistance targets.

The many and diverse beta-lactamases produced by bacteria, particularly by Gram-negative pathogens, are increasingly posing a serious threat to the clinical utility of beta-lactams. First-generation inhibitors (clavulanic acid, sulbactam, tazobactam) focus on Ambler class A enzymes. However, recent structural upgrades of class A beta-lactamases (e.g. TEM, SHV) have extended their spectrum (extended-spectrum beta-lactamases and carbapenemases [Sme, NMC-A, IMI-1]) and have brought about the possibility of beta-lactamase-inhibitor resistance. Furthermore, the mobilisation and spread of originally chromosomal class C enzymes (CMY, MIR), the growing clinical importance of class B enzymes (IMP, VIM), the emergence of inhibitor-resistant, broad spectrum class D (OXA) enzymes and the co-existence of different classes of beta-lactamases in the same pathogen have spurred research toward universal inhibitors. A complicating issue is target accessibility in Gram-negative bacteria, particularly in Enterobacter, Acinetobacter, Pseudomonas, Stenotrophomonas and other organisms, which is necessary in order for the inhibitor to synergise with vulnerable beta-lactam antibiotics. Several new, broad-spectrum inhibitors have emerged: cephem sulfones and oxapenems are upgrades of penam sulfones and oxapenams, respectively, with cephem sulfones possibly extending their inhibition to class B metallo-enzymes; and boronates and phosphonates are designed de novo, based on common structural and mechanistic features of serine beta-lactamases.

Animals↗

The Annual African American Conference on Diabetes: evolving program evaluation with evolving program implementation.

BACKGROUND: According to 2003 Behavioral Risk Factor Surveillance System data, South Carolina has the fourth highest rate of overall diabetes among the 50 states (9.3%) but the second highest rate among African Americans (15.5%). Nationwide, African Americans are disproportionately affected by diabetes. In addition, 40% of the African American population in South Carolina lives in a rural area, and approximately 26% live at or below the poverty level. Lack of access to health care and diabetes education are additional barriers for people with diabetes and their families. CONTEXT: Since 1997, the South Carolina Diabetes Prevention and Control Program and the Diabetes Today Advisory Council have sponsored the African American Conference on Diabetes, which targets African Americans with diabetes, their families, and their caregivers. This article describes the evolution of the conference and its evaluation. METHODS: In 2002, we conducted focus groups with 20 African American conference attendees with diabetes to 1) assess the program's effects, 2) determine how to reach more individuals, and 3) improve programming. In 2004, we incorporated the preconference and postconference Diabetes Understanding Scale survey to assess the cognitive impact of the conference on participants. CONSEQUENCES: Focus group results revealed that participants wanted to attend the conference because of the opportunity to increase their knowledge and change their behaviors through 1) education, 2) social support, 3) resources, and 4) logistics. Self-rated understanding increased significantly after the conference for each cognitive understanding item on the Diabetes Understanding Scale. INTERPRETATION: Focus group results suggested that participants who continue to attend the conference year after year may improve diabetes self-management skills. A quantitative evaluation showed that this 1-day diabetes education conference significantly increased short-term, self-rated cognitive understanding of diabetes behaviors.

Black or African American↗

Cooperation and conflict in the evolution of individuality. IV. Conflict mediation and evolvability in Volvox carteri.

The continued well being of evolutionary individuals (units of selection and evolution) depends upon their evolvability, that is their capacity to generate and evolve adaptations at their level of organization, as well as their longer term capacity for diversifying into more complex evolutionary forms. During a transition from a lower- to higher-level individual, such as the transition between unicellular and multicellular organisms, the evolvability of the lower-level (cells) must be restricted, while the evolvability of the new higher-level unit (multicellular organism) must be enhanced. For these reasons, understanding the factors leading to an evolutionary transition should help us to understand the factors underlying the emergence of evolvability of a new evolutionary unit. Cooperation among lower-level units is fundamental to the origin of new functions in the higher-level unit. Cooperation can produce a new more complex evolutionary unit, with the requisite properties of heritable fitness variations, because cooperation trades fitness from a lower-level (the costs of cooperation) to the higher-level (the benefits for the group). For this reason, the evolution of cooperative interactions helps us to understand the origin of new and higher-levels of fitness and organization. As cooperation creates a new level of fitness, it also creates the opportunity for conflict between levels of selection, as deleterious mutants with differing effects at the two levels arise and spread. This conflict can interfere with the evolvability of the higher-level unit, since the lower and higher-levels of selection will often "disagree" on what adaptations are most beneficial to their respective interests. Mediation of this conflict is essential to the emergence of the new evolutionary unit and to its continued evolvability. As an example, we consider the transition from unicellular to multicellular organisms and study the evolution of an early-sequestered germ-line in terms of its role in mediating conflict between the two levels of selection, the cell and the cell group. We apply our theoretical framework to the evolution of germ/soma differentiation in the green algal group Volvocales. In the most complex member of the group, Volvox carteri, the potential conflicts among lower-level cells as to the "right" to reproduce the higher-level individual (i.e. the colony) have been mediated by restricting immortality and totipotency to the germ-line. However, this mediation, and the evolution of an early segregated germ-line, was achieved by suppressing mitotic and differentiation capabilities in all post-embryonic cells. By handicapping the soma in this way, individuality is ensured, but the solution has affected the long-term evolvability of this lineage. We think that although conflict mediation is pivotal to the emergence of individuality at the higher-level, the way in which the mediation is achieved can greatly affect the longer-term evolvability of the lineage.

Adaptation, Physiological↗

Evolvability of an RNA virus is determined by its mutational neighbourhood.

The ubiquity of mechanisms that generate genetic variation has spurred arguments that evolvability, the ability to generate adaptive variation, has itself evolved in response to natural selection. The high mutation rate of RNA viruses is postulated to be an adaptation for evolvability, but the paradox is that whereas some RNA viruses evolve at high rates, others are highly stable. Here we show that evolvability in the RNA bacteriophage phi6 is also determined by the accessibility of advantageous genotypes within the mutational neighbourhood (the set of mutants one or a few mutational steps away). We found that two phi6 populations that were derived from a single ancestral phage repeatedly evolved at different rates and toward different fitness maxima. Fitness measurements of individual phages showed that the fitness distribution of mutants differed between the two populations. Whereas population A, which evolved toward a higher maximum, had a distribution that contained many advantageous mutants, population B, which evolved toward a lower maximum, had a distribution that contained only deleterious mutants. We interpret these distributions to measure the fitness effects of genotypes that are mutationally available to the two populations. Thus, the evolvability of phi6 is constrained by the distribution of its mutational neighbours, despite the fact that this phage has the characteristic high mutation rate of RNA viruses.

Bacteriophage phi 6↗

Phylogenetic utility of rapidly evolving DNA at high taxonomical levels: contrasting matK, trnT-F, and rbcL in basal angiosperms.

The prevailing view in molecular systematics is that relationships among distantly related taxa should be inferred using DNA segments with low rates of evolution. However, recent analyses of sequences from the rapidly evolving matK and trnT-trnF regions yielded well resolved and highly supported trees for early diverging angiosperms. We compare here the phylogenetic structure in matK, trnT-F, and rbcL datasets for the same 42, primarily basal angiosperm taxa. Phylogenetic trees based on matK or trnT-F are far more robust than those based on rbcL. Combined analysis of the rapidly evolving regions provides support for higher-level relationships stronger than that derived from analyses of multi-gene datasets of up to several fold the number of characters analyzed here. In addition to displaying a higher percentage of parsimony-informative characters, the average phylogenetic signal per informative character is significantly higher in the datasets from rapidly evolving DNA than in the more slowly evolving rbcL, as detected using resampling of identical numbers of parsimony-informative characters from the data matrices and subjecting different statistics for overall tree robustness and phylogenetic signal to significance tests. Automated via a set of scripts, the method used here should be easily extendable to comparisons of a broader range of genomic regions for varying taxon samplings. The relative performance of markers correlates not only with a lower mean homoplasy in matK and trnT-trnF compared to rbcL, but in particular correlates negatively with the percentage of sites exhibiting maximum or close to maximum homoplasy. A likelihood ratio test confirms that the rapidly evolving gene matK evolves significantly closer to neutrality, which may be one of the underlying factors for lower levels of overall homoplasy. Our results are in line with evidence from simulation studies suggesting that the deleterious effect of multiple hits in using rapidly evolving DNA at rather deep phylogenetic levels may have been overestimated, and thus promote extending the use of rapidly evolving DNA to deeper phylogenetic levels.

Codon↗

Prokaryote and eukaryote evolvability.

The concept of evolvability covers a broad spectrum of, often contradictory, ideas. At one end of the spectrum it is equivalent to the statement that evolution is possible, at the other end are untestable post hoc explanations, such as the suggestion that current evolutionary theory cannot explain the evolution of evolvability. We examine similarities and differences in eukaryote and prokaryote evolvability, and look for explanations that are compatible with a wide range of observations. Differences in genome organisation between eukaryotes and prokaryotes meets this criterion. The single origin of replication in prokaryote chromosomes (versus multiple origins in eukaryotes) accounts for many differences because the time to replicate a prokaryote genome limits its size (and the accumulation of junk DNA). Both prokaryotes and eukaryotes appear to switch from genetic stability to genetic change in response to stress. We examine a range of stress responses, and discuss how these impact on evolvability, particularly in unicellular organisms versus complex multicellular ones. Evolvability is also limited by environmental interactions (including competition) and we describe a model that places limits on potential evolvability. Examples are given of its application to predator competition and limits to lateral gene transfer. We suggest that unicellular organisms evolve largely through a process of metabolic change, resulting in biochemical diversity. Multicellular organisms evolve largely through morphological changes, not through extensive changes to cellular biochemistry.

Adaptation, Physiological↗