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The evolution of adaptive immunity.

Approximately 500 mya two types of recombinatorial adaptive immune systems appeared in vertebrates. Jawed vertebrates generate a diverse repertoire of B and T cell antigen receptors through the rearrangement of immunoglobulin V, D, and J gene fragments, whereas jawless fish assemble their variable lymphocyte receptors through recombinatorial usage of leucine-rich repeat (LRR) modular units. Invariant germ line-encoded, LRR-containing proteins are pivotal mediators of microbial recognition throughout the plant and animal kingdoms. Whereas the genomes of plants and deuterostome and chordate invertebrates harbor large arsenals of recognition receptors primarily encoding LRR-containing proteins, relatively few innate pattern recognition receptors suffice for survival of pathogen-infected nematodes, insects, and vertebrates. The appearance of a lymphocyte-based recombinatorial system of anticipatory immunity in the vertebrates may have been driven by a need to facilitate developmental and morphological plasticity in addition to the advantage conferred by the ability to recognize a larger portion of the antigenic world.

Adaptation, Physiological↗

The pelvis as a passageway. I. Evolution and adaptations.

Man has an unusual pelvis, a large fetal head, and a complicated mechanism of labour. The evolution of the pelvic girdle, like that of the hind limb, is a story with some chapters still missing. In mammals the pelvis has been modified in various interesting ways to effect compromises between its locomotor function and the requirements of parturition. Among the primates, the increasing size of the fetal brain may have necessitated a change in the attitude of the head from extension to flexion as it goes through the pelvis. The distinctive shape of the hominid pelvis is probably an adaptation to bipedal gait. The birth canal has changed from a shallow bony ring to a deep curved tube, through which the mature fetal head can only pass by rotating as it descends.

Adaptation, Physiological↗

Salivary glands, cellular evolution, and adaptive radiation in mammals.

Evolutionary theory provides the foundation for interpretation of the natural world, but one remaining major challenge is to link genetic variation and particular gene products to natural selection and adaptation. Another challenge is to describe the role of cells, especially secretory cells, in the evolutionary process. Comparative studies of mammalian salivary glands could serve as an insightful model. Our theoretical approach combines data on genomic and chromosomal evolution with data on secretory cells and proteins so that adaptation can be understood in context of these interrelated components. The present paper reviews patterns and types of interspecific salivary gland cell variation that we have documented at the ultrastructural level. This data set shows ways in which secretory cells and the secretory process may have been evolutionarily modified in mammals. As a further example of our approach we also review how proline-rich-proteins (PRPs) may have had adaptive significance in the evolution of mammals, especially rodents. Our working hypotheses are: that Ca+2-binding acidic PRPs are incompatible with ever-growing dentition; and that presence or absence of tannin-binding basic PRPs was a major factor in ecological diversification in rodents. Some rodents might even exhibit behavioral compensation for absence of basic PRPs in their saliva and this illustrates the complex alternatives available to natural selection.

Animals↗

Biological adaptation and social behaviour.

In 1930, both Fisher and Wright identified Darwin's initial concept of adaptive evolution in the light of the genetical theory with intergenerational variation in allelic frequencies brought about by the action of natural selection through differential reproduction. They emphasized that selection only works at the level of the individual and that its only consequence is to increase fitness. One genetical evolution not easy to explain on these bases was that of social behaviour because any altruistic gene disadvantageous for its carriers in an antisocial environment would have been opposed by selection. In the 1950s, ethologists focusing on what appeared to be evolved collective behaviours, hypothesized that selection could operate at group level. Though the controversy between group selectionists and evolutionary geneticists ended by the rejection of the evolutionary role of group selection, it has remained a subject of investigation until now. Kin selection, proposed by Hamilton, offered a solution to the problem of the evolution of altruism and gave the impetus to the trend of adaptive explanations of basic behaviours, which was to become the core of human sociobiology. The intrusion of behaviour into the process of adaptive evolution was an invitation to investigate culture as an evolutive process. The first sociobiological interpretations of culture as a derivative of genetic processes were followed by other ideas in which culture, though channelled by evolved predispositions, was essentially free from biological determinism. It is concluded that as we have come to better understand human adaptation, its complexities have been further revealed, a development already implicit in Darwin's notion.

Adaptation, Physiological↗

Encephalization, adaptation and evolution of chiroptera: A statistical analysis with further evidence for bat monophyly.

As part of a large-scale study on brain morphometrics and adaptations in mammals, we addressed the problem of chiropteran evolution. A specific statistical framework was designed to test which of two competing hypotheses (bat monophyly vs. diphyly) is more strongly supported by quantitative brain data. Our analyses, based on 120 species, revealed that megabats and microbats were more closely related to each other than to primates, and illustrated the convergent adaptations of the brain of bats to similar trophic (i.e. feeding related) niches. Ecologically-corrected characters were then used to derive a new phylogeny which also supports the chiropteran clade. The monophyletic origin of bats is the preferred hypothesis to explain brain quantitative evolution in chiropterans and primates.

Adaptation, Physiological↗

Replicated evolution of integrated plastic responses during early adaptive divergence.

Colonization of a novel environment is expected to result in adaptive divergence from the ancestral population when selection favors a new phenotypic optimum. Local adaptation in the new environment occurs through the accumulation and integration of character states that positively affect fitness. The role played by plastic traits in adaptation to a novel environment has generally been ignored, except for variable environments. We propose that if conditions in a relatively stable but novel environment induce phenotypically plastic responses in many traits, and if genetic variation exists in the form of those responses, then selection may initially favor the accumulation and integration of functionally useful plastic responses. Early divergence between ancestral and colonist forms will then occur with respect to their plastic responses across the gradient bounded by ancestral and novel environmental conditions. To test this, we compared the magnitude, integration, and pattern of plastic character responses in external body form induced by shallow versus open water conditions between two sunfish ecomorphs that coexist in four postglacial lakes. The novel sunfish ecomorph is present in the deeper open water habitat, whereas the ancestral ecomorph inhabits the shallow waters along the lake margin. Plastic responses by open water ecomorphs were more correlated than those of their local shallow water ecomorph in two of the populations, whereas equal levels of correlated plastic character responses occurred between ecomorphs in the other two populations. Small but persistent differences occurred between ecomorph pairs in the pattern of their character responses, suggesting a recent divergence. Open water ecomorphs shared some similarities in the covariance among plastic responses to rearing environment. Replication in the form of correlated plastic responses among populations of open water ecomorphs suggests that plastic character states may evolve under selection. Variation between ecomorphs and among lake populations in the covariance of plastic responses suggests the presence of genetic variation in plastic character responses. In three populations, open water ecomorphs also exhibited larger plastic responses to the environmental gradient than the local shallow water ecomorph. This could account for the greater integration of plastic responses in open water ecomorphs in two of the populations. This suggests that the plastic responses of local sunfish ecomorphs can diverge through changes in the magnitude and coordination of plastic responses. Although these results require further investigation, they suggest that early adaptive evolution in a novel environment can include changes to plastic character states. The genetic assimilation of coordinated plastic responses could result in the further, and possibly rapid, divergence of such populations and could also account for the evolution of genes of major effect that contribute to suites of phenotypic differences between divergent populations.

Acclimatization↗

The two tempos of nuclear pore complex evolution: highly adapting proteins in an ancient frozen structure.

BACKGROUND: The origin of the nuclear compartment has been extensively debated, leading to several alternative views on the evolution of the eukaryotic nucleus. Until recently, too little phylogenetic information was available to address this issue by using multiple characters for many lineages. RESULTS: We analyzed 65 proteins integral to or associated with the nuclear pore complex (NPC), including all the identified nucleoporins, the components of their anchoring system and some of their main partners. We used reconstruction of ancestral sequences of these proteins to expand the detection of homologs, and showed that the majority of them, present all over the nuclear pore structure, share homologs in all extant eukaryotic lineages. The anchoring system, by contrast, is analogous between the different eukaryotic lineages and is thus a relatively recent innovation. We also showed the existence of high heterogeneity of evolutionary rates between these proteins, as well as between and within lineages. We show that the ubiquitous genes of the nuclear pore structure are not strongly conserved at the sequence level, and that only their domains are relatively well preserved. CONCLUSION: We propose that an NPC very similar to the extant one was already present in at least the last common ancestor of all extant eukaryotes and it would not have undergone major changes since its early origin. Importantly, we observe that sequences and structures obey two very different tempos of evolution. We suggest that, despite strong constraints that froze the structural evolution of the nuclear pore, the NPC is still highly adaptive, modern, and flexible at the sequence level.

Adaptation, Physiological↗

Hierarchical organization of biological systems and the structure of adaptation in evolution and tumorigenesis.

Biological systems are structurally organized according to patterns repeated at each hierarchical level. Complex units are composed of so-called interactors, systems that by cooperative interaction maintain the structure of the complex unit. Interactors are composed of large numbers of assemblies of complex units of a limited number of types of a lower hierarchical level. Thus, macromolecules, cells, organisms and ecological communities should be defined as complex units, and cellular organelles, organs and oligospecies populations as the interactors between those units. The similarity of organization at each level should make it possible to describe patterns of structure at one level and apply it to organization at another level. This was tested for the structural aspects of adaptation as viewed in a pathobiological context. Adaptation is then viewed as the result of stress seen at the level of the interactors or at the level of the lower complex units related to the type of stress. Subsequently, this structure of adaptation was applied to adaptation in biological evolution and tumorigenesis, which has led to the conclusion that stress is a driving force for both and that an increase in number of organisms or cells may precede heritable changes or mutations, respectively.

Adaptation, Physiological↗

An empirical test of the mutational landscape model of adaptation using a single-stranded DNA virus.

The primary impediment to formulating a general theory for adaptive evolution has been the unknown distribution of fitness effects for new beneficial mutations. By applying extreme value theory, Gillespie circumvented this issue in his mutational landscape model for the adaptation of DNA sequences, and Orr recently extended Gillespie's model, generating testable predictions regarding the course of adaptive evolution. Here we provide the first empirical examination of this model, using a single-stranded DNA bacteriophage related to phiX174, and find that our data are consistent with Orr's predictions, provided that the model is adjusted to incorporate mutation bias. Orr's work suggests that there may be generalities in adaptive molecular evolution that transcend the biological details of a system, but we show that for the model to be useful as a predictive or inferential tool, some adjustments for the biology of the system will be necessary.

Adaptation, Biological↗

Duplication and relocation of the functional DPY19L2 gene within low copy repeats.

BACKGROUND: Low copy repeats (LCRs) are thought to play an important role in recent gene evolution, especially when they facilitate gene duplications. Duplicate genes are fundamental to adaptive evolution, providing substrates for the development of new or shared gene functions. Moreover, silencing of duplicate genes can have an indirect effect on adaptive evolution by causing genomic relocation of functional genes. These changes are theorized to have been a major factor in speciation. RESULTS: Here we present a novel example showing functional gene relocation within a LCR. We characterize the genomic structure and gene content of eight related LCRs on human Chromosomes 7 and 12. Two members of a novel transmembrane gene family, DPY19L, were identified in these regions, along with six transcribed pseudogenes. One of these genes, DPY19L2, is found on Chromosome 12 and is not syntenic with its mouse orthologue. Instead, the human locus syntenic to mouse Dpy19l2 contains a pseudogene, DPY19L2P1. This indicates that the ancestral copy of this gene has been silenced, while the descendant copy has remained active. Thus, the functional copy of this gene has been relocated to a new genomic locus. We then describe the expansion and evolution of the DPY19L gene family from a single gene found in invertebrate animals. Ancient duplications have led to multiple homologues in different lineages, with three in fish, frogs and birds and four in mammals. CONCLUSION: Our results show that the DPY19L family has expanded throughout the vertebrate lineage and has undergone recent primate-specific evolution within LCRs.

Amino Acid Sequence↗

Divergent adaptation of Escherichia coli to cyclic ultraviolet light exposures.

The genetic changes taking place during adaptive evolution are particularly interesting in evolutionary biology. As a consequence of adaptive evolution, natural populations of an organism under selective conditions change genetically and phenotypically after a number of generations in order to survive in that particular environment. When a DNA-damaging and mutagenic agent like UV light is experimentally used as a selective factor, natural resistance of bacteria to this agent is normally increased through processes of mutation and selection. Since UV-induced mutagenesis is not restricted to particular chromosomal regions, different UV resistance mechanisms will equally probably evolve as a consequence of cyclic UV irradiation. However, it is also possible that as a consequence of the selective process, one UV resistance mechanism is preferentially selected, causing adaptive convergence of different bacterial cultures. This may occur if the most abundant or lethal kind of DNA lesion is preferentially managed by a particular DNA repair pathway and even by a specific repair enzyme or if resistance mechanisms that decrease bacterial fitness tend to be eliminated from the populations. To examine which of these two alternatives actually takes place, five cultures of Escherichia coli were treated in parallel for 80 successive UV irradiation cycles. At the end, these five cultures gave rise to different grades of UV resistance and after a preliminary characterization we found that adaptation to cyclic UV irradiation was a consequence of selection of advantageous mutations arising in different genes related to repair and replication of DNA.

Alkaline Phosphatase↗

Gradual evolution in bacteria: evidence from Bacillus systematics.

The bacterial genome projects have suggested a central role for horizontal transfer in bacterial adaptation, but it is difficult to rule out an adaptive role for ordinary genetic change in existing genes. The bacterial systematics literature can readily address the importance of gene acquisition in adaptive evolution, since phenotypic characterization typically assesses presence versus absence of metabolic capabilities, and metabolic gains and losses are most likely due to horizontal transfer and/or gene loss. Bacterial systematists have not geared their studies toward quantitative differences in metabolic capabilities, which are more likely to involve adjustments of existing genes. Here, quantitative variation in metabolism within and between three closely related Bacillus taxa has been assayed. While these taxa show no qualitative (i.e. presence versus absence) differences in resource utilization, they are quantitatively different in utilization of 8 % of 95 resources tested. Moreover, 93 % of the resources tested showed significant quantitative variation among strains within a single taxon. These results suggest that ordinary genetic changes in existing genes may play an important role in adaptation. If these results are typical, future genomically based assays of quantitative variation in phenotype (e.g. microarray analysis of mRNA concentrations) may identify hundreds of genes whose expression has been modified. A protocol is presented for identifying those modifications of gene expression and those gene acquisitions that are most likely to have played a role in adaptive evolution.

Adaptation, Physiological↗

Gene conversion may aid adaptive peak shifts.

Gene conversion is often viewed as a homogenizing force that opposes adaptive evolution. The objective of this study is to suggest a potential role for gene conversion in adaptive evolution of proteins through aiding the transfer of a population from one adaptive peak to another. Our hypothesis starts with the observation that a tandem gene duplication may result in an extra gene copy that is released from selective constraints. In such cases, individually deleterious mutations may accumulate on the extra copy of the gene, and through gene conversion these mutations may subsequently be presented to the functioning gene for selection en masse. Thus, groups of mutations that jointly confer a selective advantage may regularly be made available for selection. We present a mathematical model of this process and identify the range of rates of gene conversion, gene duplication and mutation under which it may operate. The results indicate that the process may be biologically feasible if the rate of appearance of the potentially beneficial mutations is not too small in relation to the rates of null mutation and of gene conversion. This process appears to be a possible mechanism for effecting adaptive peak shifts in large populations. We show that all the evolutionary steps in the proposed model may have occurred in the evolution of primate gamma -globin genes. We suggest that hide-and-release mechanisms for genetic variation may constitute a more general principal of evolvability.

Adaptation, Physiological↗

Diversifying selection of the tumor-growth promoter angiogenin in primate evolution.

Diversifying selection drives the rapid differentiation of gene sequences and is one of the main forces behind adaptive evolution. Most genes known to be shaped by diversifying selection are those involved in host-pathogen or male-female interactions characterized as molecular "arms races." Here we report the unexpected detection of diversifying selection in the evolution of a tumor-growth promoter, angiogenin (ANG). A comparison among 11 primate species demonstrates that ANG has a significantly higher rate of nucleotide substitution at nonsynonymous sites than at synonymous sites, a hallmark of positive selection acting at the molecular level. Furthermore, we observed significant charge diversity at the molecular surface, suggesting the presence of selective pressures in the microenvironment of ANG, including its binding molecules. A population survey of ANG in chimpanzees, however, reveals no polymorphism, which may have resulted from a recent selective sweep of a charge-altering substitution in chimpanzee evolution. Functional assays of recombinant ANGs from the human and owl monkey indicate that primate ANGs retain angiogenic activity despite rapid evolution. Our study, together with findings of similar selection in the primate breast cancer suppressor gene, BRCA1, reveals an intriguing phenomenon of unusual selective pressures on, and adaptive evolution of, cancer-related genes in primate evolution.

Amino Acid Sequence↗

Rapid speciation, morphological evolution, and adaptation to extreme environments in South African sand lizards (Meroles) as revealed by mitochondrial gene sequences.

Data derived from the morphology of the seven species of South African sand lizards, Meroles (Reptilia, Lacertidae), and their outgroups produce a robust estimate of phylogeny when a maximum parsimony approach is applied. The estimate is fully resolved with little character conflict and internal branches are relatively long. This analysis indicates that Meroles is a true clade that includes the aberrant lacertid long separated as Aporosaura anchietae. The tree is pectinate, its successive external branches representing species with increasing adaptation to desert conditions, especially aeolian sand habitats. This pattern, and the robustness of the tree, support a model of invasion of severe habitats in which successive rounds of speciation, displacement, and adaptation result in spread into extreme ecological situations. To test the robust morphological phylogeny and, indirectly, the model as well, DNA from mitochondrial 12S and 16S ribosomal genes was sequenced and analyzed by both maximum parsimony and maximum likelihood approaches. Trees produced were largely congruent with that derived from morphology, although different from ones resulting from protein electrophoresis. However, in contrast to the internal branches of the morphological tree, those of the DNA maximum likelihood tree are quite short. The DNA data provide some corroboration for the relationships within Meroles based on morphology and consequently for the model as well. The disparity in internal branch lengths between the maximum parsimony morphological and maximum likelihood DNA trees may well indicate that the multiple adaptations to desert conditions arising on the main lineage of Meroles evolved quite rapidly. In this study DNA thus not only corroborates the phylogeny but also provides evidence about another aspect of evolutionary history.

Adaptation, Physiological↗

Population differentiation in an annual legume: local adaptation.

Studies of many plants species have demonstrated adaptive genetic differentiation to local environmental conditions. Typically these studies are conducted to evaluate adaptation to contrasting environments. As a consequence, although local adaptation has been frequently demonstrated, we have little information as to the spatial scale of adaptive evolution. We evaluated adaptive differentiation between populations of the annual legume Chamaecrista fasciculata using a replicated common-garden design. Study sites were established in three field locations that are home to native populations of C. fasciculata. Each location was planted for two years with seed from the population native to the study site (home population) and populations located six distances (0.1-2000 km) from each site (transplanted populations). Seeds were planted into the study sites with minimum disturbance to determine the scale of local adaptation, as measured by a home-site fitness advantage, for five fitness components: germination, survival, vegetative biomass, fruit production, and the number of fruit produced per seed planted (an estimate of cumulative fitness). For all characters there was little evidence for local adaptation, except at the furthest spatial scales. Patterns of adaptive differentiation were fairly consistent in two of the three sites, but varied between years. Little genetic variation was expressed at the third site. These results, combined with previous estimates of limited gene flow, suggest that metapopulation processes and temporal environmental variation act together to reduce local adaptation, except over long distances.

Adaptation, Physiological↗