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The group covariance effect and fitness trade-offs during evolutionary transitions in individuality.

Transforming our understanding of life is the realization that evolution occurs not only among individuals within populations but also through the integration of groups of preexisting individuals into a new higher-level individual, that is, through evolutionary transitions in individuality. During evolutionary transitions (such as during the origin of gene networks, bacteria-like cells, eukaryotic cells, multicellular organisms, and societies), fitness must be reorganized; specifically, it must be transferred from the lower- to the higher-level units and partitioned among the lower-level units that specialize in the fitness components of the new higher-level individual. This paper studies the role of fitness trade-offs in fitness reorganization, the evolution of cooperation, and the conversion of a group into a new individual during the origin of multicellular life. Specifically, this study shows that the fitness of the group is augmented over the average fitness of its members according to a covariance effect. This covariance effect appears to be one of the first emergent properties of the group and a general aspect of groups with multiplicative properties that are themselves averages of properties of lower-level units. The covariance effect allows groups to break through the constraints that govern their members, and this effect likely applies to group dynamics in other fields.

Animals↗

Y chromosome polymorphism is a strong determinant of male fitness in Drosophila melanogaster.

In many species, the Y (or W) chromosome carries relatively few functional genes. This observation motivates the null hypothesis that the Y will be a minor contributor to genetic variation for fitness. Previous data and theory supported the null hypothesis, but evidence presented here shows that the Y of Drosophila melanogaster is a major determinant of a male's total fitness, with standing genetic variation estimated to be 68% of that of an entire X/autosome genomic haplotype. Most Y-linked genes are expressed during spermatogenesis, and correspondingly, we found that the Y influences fitness primarily through its effect on a male's reproductive success (sperm competition and/or mating success) rather than his egg-to-adult viability. But the fitness of a Y highly depended on the genetic makeup of its bearer, reverting from high to low in different genetic backgrounds. This pattern leads to large epistatic (inconsistent among backgrounds) but no additive (consistent among backgrounds) Y-linked genetic variance for fitness. On a microevolutionary scale, the observed large epistatic variation on the Y substantially reduces heritable variation for fitness among males, and on a macroevolutionary scale, the Y produces strong selection for genomic rearrangements that move interacting genes onto the nonrecombining region of the Y.

Animals↗

Increment of average fitness for multiple alleles.

When the various genotypes (A(i)A(i), A(i)A(i)) of an autosomal locus do not reproduce and contribute equally to the genetic composition of the next generation, we say that the "fitness" values of the genotypes are different, or that there is natural selection on the genotypes. Under natural selection, the average fitness for a random mating population increases with time until a stationary state is reached. This communication gives a general expression for the amount of increase of the average fitness per generation of selection. It was found that the amount of increase depends largely on the linear or additive component of the variance of the genotype fitness values and, to a lesser extent, on the dominance pattern for the values of heterozygote fitness relative to those of homozygote fitness.

Genetic Complementation Test↗

Rapid fitness losses in mammalian RNA virus clones due to Muller's ratchet.

Muller's ratchet is an important concept in population genetics. It predicts that when mutation rates are high and a significant proportion of mutations are deleterious, a kind of irreversible ratchet mechanism will gradually decrease the mean fitness of small populations of asexual organisms. In contrast, sexual recombination may stop or reverse this mutational ratchet by recombinational repair of genetic damage. Experimental support for Muller's ratchet has previously been obtained in protozoa and in a tripartite RNA bacteriophage. We now show clear evidence that Muller's ratchet can operate on a nonsegmented nonrecombining pathogenic RNA virus of animals and humans. We did genetic bottleneck passages (plaque-to-plaque transfers) of vesicular somatitis virus (VSV) and then quantitated relative fitness of the bottleneck clones by allowing direct replication competition in mixed infections in cell culture. We document variable fitness drops (some severe) following only 20 plaque-to-plaque transfers of VSV. In some clones no fitness changes (or only insignificant changes) were observed. Surprisingly, the most regular and severe fitness losses occurred during virus passages on a new host cell type. These results again demonstrate the extreme genetic and biological variability of RNA virus populations. Muller's ratchet could have significant implications for variability of disease severity during virus outbreaks, since genetic bottlenecks must often occur during respiratory droplet transmissions and during spread of low-yield RNA viruses from one body site to another (as with human immunodeficiency virus). Likewise, the lower-probability generation of increased-fitness clones during repeated genetic bottleneck transfers of RNA viruses in nature might also affect disease pathogenesis in infected individuals and in host populations. Whenever genetic bottlenecks of RNA viruses occur, enhanced biological differences among viral subpopulations may result.

Animals↗

Morphological evolution through complex domains of fitness.

Computer simulated phenotypic walks through multi-dimensional fitness-landscapes indicate that (i) the number of phenotypes capable of reconciling conflicting morphological requirements increases in proportion to the number of manifold functional obligations an organism must perform to grow, survive, and reproduce, and (ii) walks over multi-task fitness-landscapes require fewer but larger phenotypic transformations than those through single-task landscapes. These results were determined by (i) simulating a "morphospace" containing 200,000 phenotypes reminiscent of early Paleozoic vascular sporophytes, (ii) evaluating the capacity of each morphology to perform each of three tasks (light interception, mechanical support, and reproduction) as well as the ability to reconcile the conflicting morphological requirements for the four combinatorial permutations of these tasks, (iii) simulating the walks obtaining all phenotypic maxima or optima within the seven "fitness-landscapes," and (iv) computing the mean morphological variation attending these walks. The results of these simulations, whose credibility is discussed in the context of early vascular land-plant evolution, suggest that both the number and the accessibility of phenotypic optima increase as the number of functional obligations contributing to total fitness increases (i.e., as the complexity of optimal phenotypes increases, the fitnesses of optima fall closer to the mean fitness of all the phenotypes under selection).

Biological Evolution↗

Adult fitness consequences of sexual selection in Drosophila melanogaster.

Few experiments have demonstrated a genetic correlation between the process of sexual selection and fitness benefits in offspring, either through female choice or male competition. Those that have looked at the relationship between female choice and offspring fitness have focused on juvenile fitness components, rather than fitness at later stages in the life cycle. In addition, many of these studies have not controlled for possible maternal effects. To test for a relationship between sexual selection and adult fitness, we carried out an artificial selection experiment in the fruit fly, Drosophila melanogaster. We created two treatments that varied in the level of opportunity for sexual selection. Increased opportunity for female choice and male competition was genetically correlated with an increase in adult survivorship, as well as an increase in male and female body size. Contrary to previous, single-generation studies, we did not find an increase in larval competitive ability. This study demonstrates that mate choice and/or male-male competition are correlated with an increase in at least one adult fitness component of offspring.

Aging↗

Short cognitive/neuropsychological test battery for first-tier fitness-to-drive assessment of older adults.

Since fitness-to-drive evaluation of elderly drivers has become an important issue, we developed a short first-tier screening battery to evaluate the necessity for further referral to specialised centres. Our sample consisted of 84 subjects between 65 and 96 years who came to the Belgian Road Safety institute for a fitness-to-drive evaluation. Using cross-validated discriminant analyses, the predictive power of a battery consisting of the Trail Making Test, Part A, a visual acuity test, a clock drawing test, the Mini-Mental State Examination (MMSE) and age was analysed. The judgement by an independent driver instructor (fit-to-drive vs. not unconditional fit-to-drive), based on a real world road test was used as the dependent variable. Classification functions based on the significant discriminant function yielded a specificity score of 85% (subjects fit-to-drive correctly classified) and a sensitivity score of 80% (subjects as not unconditional fit-to-drive correctly classified). These results highlight the potential value of a short screening instrument that can be used in primary health care settings. This instrument may be useful as a first step in a multi-tier assessment procedure.

Aged↗

Comparison of six respirator fit-test methods with an actual measurement of exposure in a simulated health care environment: Part III--Validation.

This article, the last in a series of three, describes the validation phase of a study conducted to test the correlation of respirator fit factors to the subject's actual exposure using biological sampling. The study consisted of three phases: protocol development, method comparison testing, and validation. Six quantitative fit-test methods were evaluated in the method comparison testing phase. The two fit methods with the highest correlation with the wearers' measured exposure were a corn oil method (R2 = 0.81) and an ambient aerosol method (R2 = 0.78). Because the ambient aerosol method is more commonly used in the workplace, it was selected for further analysis. In this validation phase, the fit factors measured during the ambient aerosol fit-test were used to calculate the exposures to Freon-113 by using the model determined in the method comparison testing phase of the study. The actual Freon-113 exposures were then measured and compared with the predicted exposures. The results verified that the ambient aerosol method fit factors are highly correlated to the total Freon-113 exposure dose and thus that the model had a predictive ability.

Air Pollutants, Occupational↗

Investigations of the impact of altered auditory feedback in-the-ear devices on the speech of people who stutter: initial fitting and 4-month follow-up.

BACKGROUND: Self-contained ear-level devices delivering altered auditory feedback (AAF) for the application with those who stutter have only been recently developed. AIMS: The paper examines the first therapeutic application of self-contained ear-level devices in three experiments. The effect of the device on the proportion of stuttered syllables and speech naturalness was investigated following initial fitting and at 4 months post-fitting. METHODS & PROCEDURES: Three experiments were undertaken: In Experiment 1, the effect of a self-contained in-the-ear device delivering AAF was investigated with those who stutter during reading and monologue. Two adolescents and five adults who stuttered read and produced monologue with and without a device fit monaurally. The device provided a frequency shift of +500 Hz in combination with a delayed auditory feedback of 60 ms. Custom-made ITC and CIC devices were fabricated for four adults and four youths in Experiment 2. The effect of group (i.e. youth versus adult), time (i.e. initial fitting versus 4-month follow-up), speech task (i.e. reading versus monologue), and device (i.e. present versus absent) on stuttering rate was examined. In Experiment 3, 15 naïve listeners rated the speech naturalness of speech produced by the participants in Experiment 2. Speech samples from six conditions were rated: reading and monologue without the device at the initial visit, reading and monologue with the device at the initial visit, and reading and monologue with the device at 4 months. OUTCOMES & RESULTS: In Experiment 1, the proportion of stuttered syllables was significantly (p=0.011) reduced by approximately 90% during reading and 67% during monologue with the device relative to no device. Only a significant main effect of device (p=0.0028) was found in Experiment 2. That is, stuttering rate was significantly reduced with the device in place regardless of speech task or group and remained so 4 months later. In Experiment 3, speech samples generated while wearing the device were judged to be more natural sounding than those without the device (p<0.0001) for reading and monologue with both adults and youths. There was no significant difference between the mean naturalness ratings of speech samples generated during the initial fitting with the device relative to that at 4 months with the device (p>0.05) in all cases except with the youths while engaged in monologue. For that condition, raters judged the speech produced at the initial fitting as more natural. CONCLUSIONS: These findings support the notion that a self-contained in-the-ear device delivering AAF assists those who stutter. With the device in place, stuttering is reduced and speech produced is judged to be more natural than with out the device.

Adolescent↗

Single-generation estimates of individual fitness as proxies for long-term genetic contribution.

Individual fitness is a central evolutionary concept, but the question of how it should be defined in empirical studies of natural selection remains contentious. Using founding cohorts from long-term population studies of two species of individually marked birds (collared flycatcher Ficedula albicollis and Ural owl Strix uralensis), we compared a rate-sensitive (lambdaind) and a rate-insensitive (lifetime reproductive success [LRS]) estimate of individual fitness with an estimate of long-term genetic fitness. The latter was calculated as the number of gene copies present in the population after more than two generations, as estimated by tracing genetic lineages and accounting for the fact that populations were not completely closed. When counting fledglings, rate-insensitive estimates of individual fitness correlated better than rate-sensitive estimates with estimated long-term genetic contribution. When counting recruits, both classes of estimates performed equally well. The results support the contention that simple, rate-insensitive measures of fitness, such as LRS, provide a valid and good estimate of fitness in evolutionary studies of natural populations.

Animals↗

Unifying the theories of inclusive fitness and reciprocal altruism.

Inclusive fitness and reciprocal altruism are widely thought to be distinct explanations for how altruism evolves. Here we show that they rely on the same underlying mechanism. We demonstrate this commonality by applying Hamilton's rule, normally associated with inclusive fitness, to two simple models of reciprocal altruism: one, an iterated prisoner's dilemma model with conditional behavior; the other, a mutualistic symbiosis model where two interacting species differ in conditional behaviors, fitness benefits, and costs. We employ Queller's generalization of Hamilton's rule because the traditional version of this rule does not apply when genotype and phenotype frequencies differ or when fitness effects are nonadditive, both of which are true in classic models of reciprocal altruism. Queller's equation is more general in that it applies to all situations covered by earlier versions of Hamilton's rule but also handles nonadditivity, conditional behavior, and lack of genetic similarity between altruists and recipients. Our results suggest changes to standard interpretations of Hamilton's rule that focus on kinship and indirect fitness. Despite being more than 20 years old, Queller's generalization of Hamilton's rule is not sufficiently appreciated, especially its implications for the unification of the theories of inclusive fitness and reciprocal altruism.

Altruism↗

The alcohol dehydrogenase polymorphism in Drosophila melanogaster: fitness measurements and predictions under conditions with no alcohol stress.

The relative fitnesses of the different Adh genotypes under normal laboratory conditions and in the absence of alcohol stress were estimated in Drosophila melanogaster according to Prout's method. The larval component (viability) did not reveal fitness differences between the genotypes but for the adult component significant differences were observed. The female adult component (fecundity) showed an overdominant pattern: both homozygous genotypes showed a relative fitness significantly lower than the heterozygous genotype. For the male adult component (virility) also differences were observed. The homozygous SS genotype showed a lower relative fitness than the other two genotypes. Predictions for gene frequency changes based on the estimated fitness values do show a reasonably good correspondence with frequency changes actually observed in a number of experimental cage populations and indicate a globally stable equilibrium around a frequency of the F allele of 0.40-0.70. The relevance of these fitness estimates, obtained under conditions with no alcohol stress, for the explanation of the Adh polymorphisms observed in nature is discussed.

Alcohol Dehydrogenase↗

Viabilities of third chromosomes of Drosophila pseudoobscura differing in relative competitive fitness.

Arrowhead (AR) third chromosome arrangements of Drosophila pseudoobscura, whose competitive fitnesses had been determined in population cages, were tested for their genetic loads in homozygous, heterozygous (homokaryotypic), and heterokaryotypic (AR/CH) combinations. The results showed that their competitive population cage performances were correlated to their viabilities as homozygotes but were not correlated to their viabilities as heterozygotes or as heterokaryotypes. However, the results do not fit in too simply with the mutational model of population structure, since the improvement of homozygous viability with increased competitive fitness was not accompanied by a significant degree of dominance as measured by the regression of viabilities of heterozygotes on homozygotes. Only the AR chromosomes derived from the population with poorest competitive fitness showed marked partial dominance (h=.35). The viabilities of heterokaryotypes were markedly uniform for all chromosomes tested and produced significantly greater numbers of flies per culture than the homokaryotypes. In general, the results show that the ranking of relative competitive fitnesses of these chromosomes is not a simple extrapolation of their viabilities, although marked changes in the populations tested have occurred. It is proposed that the differences in competitive fitness, homozygous viability, and degree of dominance observed among these chromosomes, arise from differences in genetic variability which enable different linkage relationships to be established for genes affecting these attributes.

Animals↗

A long-term perspective on person-environment fit and ADL dependence among older Swedish adults.

PURPOSE: This study described person-environment (P-E) fit and activities of daily living (ADLs) among older adults, and it explored the relationship between P-E fit and ADL dependence, testing Lawton's docility hypothesis at two points in time. DESIGN AND METHODS: From a random sample of individuals aged 75-84 living in a Swedish municipality, 72 persons were available for baseline and follow-up interviews over a 6-year period. Interviews and observations were used for data collection at home visits. RESULTS: P-E fit problems as well as ADL dependence increased from baseline to follow-up. Functional limitations and dependence on mobility devices increased; few changes were found concerning environmental barriers. At follow-up, ADL dependence and P-E fit correlated significantly in the frailest subsample. IMPLICATIONS: In advanced age, P-E fit problems in housing are related to ADL dependence. The increase of P-E fit problems mostly results from functional decline, although to some extent it is from environmental changes as well. The methodology used demonstrates the potential to increase our understanding of how the main components of Lawton's model change and interact over time.

Activities of Daily Living↗

The assessment of fitness to travel.

The majority of health problems encountered in association with travel stem from pre-existing, perhaps latent, illness in the individual which may be exacerbated by the rigours and hazards of travel. It is essential that the advising physician understands the hazards that are likely to be encountered during travel in order that they may develop informed decisions regarding fitness for travel and give appropriate advice. In an occupational health setting, the employer has a responsibility to safeguard the health of their employees whilst travelling on behalf of the organisation and will also have to fund any treatment abroad or the cost of repatriation. The dictating factor in determining fitness to travel will often be fitness to travel by air, consequent to the reduced partial pressure of oxygen and pressure changes in-flight. The majority of significant health problems encountered during travel are attributable to coronary heart disease and detailed guidance exists to determine fitness for travel. For many health problems little if any evidence based guidance exists and decisions will therefore have to be based on an understanding of the hazards likely to be encountered during travel. Access to appropriate standards of medical care abroad and the difficulties and expense of repatriation, should this be necessary, are also important factors to consider in addition to the basic determination of fitness for travel itself. This paper outlines the main factors to be considered when assessing fitness to travel and also examines available guidance for some of the more commonly encountered conditions.

Aerospace Medicine↗

Effects of carriage and expression of the Tn10 tetracycline-resistance operon on the fitness of Escherichia coli K12.

We have been examining the consequences of alternative modes of regulation of plasmid-borne, Tn10-encoded tetracycline resistance for the fitness of Escherichia coli. In a tetracycline-free environment, we measured the effects on fitness that were caused by (1) maximally induced expression of the resistance operon, (2) low-level constitutive expression of the resistance protein, (3) residual expression of the repressed resistance operon, (4) carriage of the resistance operon, (5) the remainder of the plasmid genome, and (6) hyperexpression of the repressor protein. We observed large reductions in fitness that were associated with induction and with constitutive expression of the tetracycline-resistance protein, but there was no discernible effect of hyperexpression of the repressor protein. We also observed a small reduction in fitness associated with the remainder of the plasmid genome. However, any reductions in fitness that were caused by residual expression and by carriage of the repressed operon were not more than 0.3%. We conclude that tight gene regulation has eliminated antagonistic pleiotropic effects of the resistance gene on fitness, so that possession of an inducible Tn10-encoded tetracycline-resistance operon imposes essentially no burden in the absence of antibiotic.

DNA Transposable Elements↗

A comparison of fitting growth models with a genetic algorithm and nonlinear regression.

A genetic algorithm (GA), an optimization procedure based on the theory of evolution, was compared with nonlinear regression for the ability of the 2 algorithms to fit the coefficients of poultry growth models. It was hypothesized that the nonlinear approach of using GA to define the parameters of growth equations would better fit the growth equations than the use of nonlinear regression. Two sets of growth data from the literature, consisting of male broiler BW grown for 168 and 170 d, were used in the study. The growth data were fit to 2 forms of the logistic model, the Gompertz, the Gompertz-Laird, and the saturated kinetic models using the SAS nonlinear algorithm (NLIN) procedure and a GA. There were no statistical differences for the comparison of the residuals (the difference between observed and predicted BWs) of growth models fit by a GA or nonlinear regression. The plotted residuals for the nonlinear regression and GA-determined growth values confirmed observations of others that the residuals have oscillations resembling sine waves that are not represented by the growth models. It was found that GA could successfully determine the coefficients of growth equations. A disadvantage of slowness in converging to the solution was found for the GA. The advantage of GA over traditional nonlinear regression is that only ranges need be specified for the parameters of the growth equations, whereas estimates of the coefficients need to be determined, and in some programs the derivatives of the growth equations need to be identified. Depending on the goal of the research, solving multivariable complex functions with an algorithm that considers several solutions at the same time in an evolutionary mode can be considered an advantage especially where there is a chance for the solution to converge on a local optimum when a global optimum is desired. It was concluded that the fitting of the growth equations was not so much a problem with the fitting methodology as it is with the form of the equation.

Algorithms↗

Increased fitness of drug resistant HIV-1 protease as a result of acquisition of compensatory mutations during suboptimal therapy.

OBJECTIVE: It is thought as a consequence of continuous replication, HIV-1 has acquired an optimal fitness state and that suboptimal antiretroviral therapy selects for drug resistant variants which show impaired fitness in the absence of the drug. In this paper we studied the evolution and fitness of viral populations appearing in a patient who received protease monotherapy. METHODS: Two factors contributing to fitness, drug resistance and protease catalytic activity, were studied at the enzymatic and virological level. RESULTS: The first drug resistant viral variants that were selected in vivo harboured one to three protease substitutions. These mutants showed reduced protease activity and consequently a reduction in viral replication capacity. During continued in vivo replication of these viruses in the presence of the drug, novel variants harbouring additional substitutions in the viral protease appeared. These variants did not display any further increase in drug resistance but demonstrated clearly increased protease activity. Consequently the replication capacity of these viruses was raised to a level at which they replicated better than the original wild-type virus. CONCLUSION: This study indicates that the viral population in the patient does not have to represent the fittest possible variants, and thus antiretroviral therapy may drive the viral population first through a lower fitness level and then to a higher fitness level.

Amino Acid Sequence↗