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The current state of insect molecular systematics: a thriving Tower of Babel.

Insect molecular systematics has undergone remarkable recent growth. Advances in methods of data generation and analysis have led to the accumulation of large amounts of DNA sequence data from most major insect groups. In addition to reviewing theoretical and methodological advances, we have compiled information on the taxa and regions sequenced from all available phylogenetic studies of insects. It is evident that investigators have not usually coordinated their efforts. The genes and regions that have been sequenced differ substantially among studies and the whole of our efforts is thus little greater than the sum of its parts. The cytochrome oxidase I, 16S, 18S, and elongation factor-1 alpha genes have been widely used and are informative across a broad range of divergences in insects. We advocate their use as standards for insect phylogenetics. Insect molecular systematics has complemented and enhanced the value of morphological and ecological data, making substantial contributions to evolutionary biology in the process. A more coordinated approach focused on gathering homologous sequence data will greatly facilitate such efforts.

Animals↗

Hormonal pleiotropy and the juvenile hormone regulation of Drosophila development and life history.

Understanding how traits are integrated at the organismal level remains a fundamental problem at the interface of developmental and evolutionary biology. Hormones, regulatory signaling molecules that coordinate multiple developmental and physiological processes, are major determinants underlying phenotypic integration. The probably best example for this is the lipid-like juvenile hormone (JH) in insects. Here we review the manifold effects of JH, the most versatile animal hormone, with an emphasis on the fruit fly Drosophila melanogaster, an organism amenable to both genetics and endocrinology. JH affects a remarkable number of processes and traits in Drosophila development and life history, including metamorphosis, behavior, reproduction, diapause, stress resistance and aging. While many molecular details underlying JH signaling remain unknown, we argue that studying "hormonal pleiotropy" offers intriguing insights into phenotypic integration and the mechanisms underlying life history evolution. In particular, we illustrate the role of JH as a key mediator of life history trade-offs.

Aging↗

Human-specific features of the cerebellum and ZP2-regulated synapse development.

Understanding the unique features of the human brain compared to non-human primates has long intrigued humankind. The cerebellum refines motor coordination and cognitive functions, contributing to the evolutionary development of human adaptability and dexterity. To identify shared and divergent features across primates, we conducted single-nucleus transcriptomic and chromatin accessibility profiling of the adult cerebellar cortex in humans, chimpanzees, macaques, and marmosets. We revealed human-specific transcriptomic and regulatory features, particularly those involved in synaptogenesis. Notably, we identified an enrichment of the sperm receptor zona pellucida glycoprotein 2 (ZP2) and its potential interactors, known for their roles in gamete interaction, in human granule cells. Experimental data show that ZP2 expression in human granule cells is induced by pontine mossy fibers, reducing synaptic proteins at pontocerebellar glomerular synapses, and decreasing cerebellar neuron electrophysiological activity. This unexpected co-option of ZP2 in human-specific synapse regulation provides insights into the evolutionary specialization of the human cerebellum.

Brain evolution↗

Zebrafish to humans: evolution of the alpha3-chain of type IV collagen and emergence of the autoimmune epitopes associated with Goodpasture syndrome.

Goodpasture syndrome is an autoimmune vascular disease associated with kidney and lung failure, with pathogenic circulating autoantibodies targeted to a set of discontinuous epitope sequences within the noncollagenous domain-1 (NC1) of the alpha3 chain of type IV collagen (alpha3(IV)NC1), the Goodpasture autoantigen. We demonstrate that basement membrane extracted NC1 domain preparations from Caenorhabditis elegans, Drosophila melanogaster, and Danio rerio do not bind Goodpasture autoantibodies, while Xenopus laevis, chicken, mouse and human alpha3(IV)NC1 domains bind autoantibodies. The alpha3(IV) chain is not present in C elegans and Drosophila melanogaster, but is first detected in the Danio rerio. Interestingly, native Danio rerio alpha3(IV)NC1 does not bind Goodpasture autoantibodies. Next, we cloned, sequenced, and generated recombinant Danio rerio alpha3(IV)NC1 domain. In contrast to recombinant human alpha3(IV)NC1 domain, there was complete absence of autoantibody binding to recombinant Danio rerio alpha3(IV)NC1. Three-dimensional molecular modeling from existing x-ray coordinates of human NC1 domain suggest that evolutionary alteration of electrostatic charge and polarity due to the emergence of critical serine, aspartic acid, and lysine residues, accompanied by the loss of asparagine and glutamine, contributes to the emergence of the 2 major Goodpasture epitopes on the human alpha3(IV)NC1 domain, as it evolved from the Danio rerio over 450 million years.

Animals↗

The challenge of paleoecological stasis: reassessing sources of evolutionary stability.

The paleontological record of the lower and middle Paleozoic Appalachian foreland basin demonstrates an unprecedented level of ecological and morphological stability on geological time scales. Some 70-80% of fossil morphospecies within assemblages persist in similar relative abundances in coordinated packages lasting as long as 7 million years despite evidence for environmental change and biotic disturbances. These intervals of stability are separated by much shorter periods of ecological and evolutionary change. This pattern appears widespread in the fossil record. Existing concepts of the evolutionary process are unable to explain this uniquely paleontological observation of faunawide coordinated stasis. A principle of evolutionary stability that arises from the ecosystem is explored here. We propose that hierarchical ecosystem theory, when extended to geological time scales, can explain long-term paleoecological stability as the result of ecosystem organization in response to high-frequency disturbance. The accompanying stability of fossil morphologies results from "ecological locking," in which selection is seen as a high-rate response of populations that is hierarchically constrained by lower-rate ecological processes. When disturbance exceeds the capacity of the system, ecological crashes remove these higher-level constraints, and evolution is free to proceed at high rates of directional selection during the organization of a new stable ecological hierarchy.

Journal Article↗

Dual nature of the genome: genes for the individual life and genes for the evolutionary progress of the population.

Biological evolution is here postulated to be driven coordinately by the products of specific evolution genes and by non-genetic elements such as the intrinsic properties of matter and random encounter with environmental factors. Evolution genes are supposed to have their own evolutionary history in which second-order selection was exerted at the population level. The products of evolution genes can act as generators of genetic variations and/or as modulators of the frequency of genetic variation. Three major natural strategies, each with a number of specific mechanisms contribute to the overall spontaneous production of genetic variants. Each of these three strategies contributes its own specific quality to genetic variation. The difficulties of experimentally investigating these strategies and a wider discussion of some of the postulates within the scientific community are outlined. Finally, the general relevance of the postulated duality of the genome for our world view is briefly mentioned.

Animals↗

Localization of genes encoding apolipoproteins CI, CII, and E to the p13----cen region of human chromosome 19.

The genes encoding apolipoproteins CI, CII, and E have been previously localized to chromosome 19. By use of rodent-human hybrid cell lines containing translocations of chromosome 19 we have now mapped these three genes to the region 19p13-19q13 and most probably 19p13-19cen. The clustering of APOC1, APOC2, and APOE must reflect their common evolutionary background and suggests that they may be coordinately regulated. Polymorphisms detected for any one gene will be useful for inheritance studies of all three.

Animals↗

A coordinated interplay: proteins with multiple functions in DNA replication, DNA repair, cell cycle/checkpoint control, and transcription.

In eukaryotic cells, DNA transactions such as replication, repair, and transcription require a large set of proteins. In all of these events, complexes of more than 30 polypetides appear to function in highly organized and structurally well-defined machines. We have learned in the past few years that the three essential macromolecular events, replication, repair, and transcription, have common functional entities and are coordinated by complex regulatory mechanisms. This can be documented for replication and repair, for replication and checkpoint control, and for replication and cell cycle control, as well as for replication and transcription. In this review we cover the three different protein classes: DNA polymerases, DNA polymerase accessory proteins, and selected transcription factors. The "common enzyme-different pathway strategy" is fascinating from several points of view: first, it might guarantee that these events are coordinated; second, it can be viewed from an evolutionary angle; and third, this strategy might provide cells with backup mechanisms for essential physiological tasks.

Cell Cycle↗

Synthesis of adenylated molybdopterin: an essential step for molybdenum insertion.

The molybdenum cofactor (Moco) is part of the active site of all molybdenum (Mo)-dependent enzymes, except nitrogenase. Moco consists of molybdopterin (MPT), a phosphorylated pyranopterin with an enedithiolate coordinating Mo and it is synthesized by an evolutionary old multistep pathway. The plant protein Cnx1 from Arabidopsis thaliana catalyzes with its two domains (E and G) the terminal step of Moco biosynthesis, the insertion of Mo into MPT. Recently, the high-resolution MPT-bound structure of the Cnx1 G domain (Cnx1G) has been determined (Kuper, J., Llamas, A., Hecht, H. J., Mendel, R. R., and Schwarz, G. (2004) Nature 430, 803-806). Besides defining the MPT-binding site a novel and unexpected modification of MPT has been identified, adenylated MPT. Here we demonstrate that it is Cnx1G that catalyzes the adenylation of MPT. In vitro synthesized MPT was quantitatively transferred from Escherichia coli MPT synthase to Cnx1G. The subsequent adenylation reaction by Cnx1G was Mg(2+)- and ATP-dependent. Whereas Mn(2+) could partially replace Mg(2+), ATP was the only nucleotide accepted by Cnx1G. Consequently the formation of pyrophosphate was demonstrated, which was dependent on the ability of Cnx1G to bind MPT. Pyrophosphate, either formed in the reaction or added externally, inhibited the Cnx1G-catalyzed MPT adenylation reaction. Catalytically inactive Cnx1G mutant variants showed impaired MPT adenylation confirming that MPT-AMP is the reaction product of Cnx1G. Therefore Cnx1G is a MPT adenylyltransferase catalyzing the activation of MPT, a universal reaction in the Moco synthetic pathway because Cnx1G is able to reconstitute also bacterial and mammalian Moco biosynthesis.

Adenosine Triphosphate↗

Effect of zooid spacing on bryozoan feeding success: is competition or facilitation more important?

Most Recent bryozoan species are encrusting sheets, and many of these colonies have densely packed feeding zooids. In this study, I tested whether tight packing of feeding zooids affects food capture. Colonies of a bryozoan with an encrusting sheet form (Membranipora membranacea) were dissected to produce individuals whose feeding zooids were (1) closely packed, (2) more widely spaced, or (3) isolated. For each type, rates of particle ingestion were measured in still water and in a freestream velocity of 2.7 cm s(-1). Ingestion rate increased when zooids were closest together, probably because of reduced refiltration and increased feeding current strength farther from the lophophores. The mean incurrent velocity within 0.02 cm above the center of the lophophore was 0.28 cm s(-1) regardless of zooid spacing; however, when the incurrent velocity was measured more than 0.1 cm from the lophophores, zooids that were close together or spaced one zooid's width apart had significantly faster incurrent velocities than single zooids. Flow visualization suggests that isolated zooids and those spaced far apart refilter more water than zooids that are close together. These results along with the observed trend of increased zooid integration over evolutionary time suggest that the benefits of increasing coordination outweigh the consequences of intrazooid competition.

Analysis of Variance↗

Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in a evolution reactor, I.

A theory for describing evolution as adaptive walks by a finite population with M walkers (M > or = 1) on an anisotropic Mt. Fuji-type fitness landscape is presented, from a thermodynamical point of view. Introducing the 'free fitness' as the sum of a fitness term and an entropy term and 'evolutionary force' as the gradient of free fitness on a fitness coordinate, we demonstrate that the behavior of these theoretical walkers is almost consistent with the thermodynamical schemes. The major conclusions are as follows: (1) an adaptive walk (=evolution) is driven by an evolutionary force in the direction in which free fitness increases; (2) the expectation of the climbing rate obeys an equation analogous to the Einstein relation in Brownian motion; (3) the standard deviation of the climbing rate is a quantity analogous to the mean thermal energy of a particle, kT (x constant). In addition, on the interpretation that the walkers climb the landscape by absorbing 'fitness information' from the surroundings, we succeeded in quantifying the fitness information and formulating a macroscopic scheme from an informational point of view.

Adaptation, Biological↗

An n locus multiallele model for gene substitution.

We discuss the conceptual conflict between a slow series of gene substitutions as the mechanism of evolutionary change, and the apparent need for rapid and coordinated changes at many loci simultaneously in producing complex adaptations. To improve on the limitations of classical theory and accommodate the enormous amount of variability disclosed by electrophoretic studies, we develop a model that can deal with gene substitution at n loci, with numerous alleles at each locus. Fitness is treated somewhat differently from the usual way by allowing it to vary between zero and the number of offspring an individual of a particular species can produce. As maximum fitnesses, we chose five as typical of large mammals, 100 for insects like Drosophila, and 1000 for very prolific species. When our model is applied to the classical problem of determining the number of generations required to change the gene frequency from 0.0001 to 0.9999 (but for 100 loci rather than one), we find that it requires 22,899 generations when maximum fitness is five, 7,984 generations when maximum fitness is 100 and 5,333 generations when it is 1000. This is something of an improvement over the 300,000 generations calculated by Haldane (1957). By allowing the fitnesses in our model to be explicitly frequency dependent, these results are reduced considerably. In addition, allowing varying proportions of the population to inbreed reduces the number of generations required for the classical problem by as much as 50%. We also point out that, given the large amount of observed genetic variation, evolutionary change may not be so much a matter of classical gene substitution as it is of changing from one array of alleles to another. With our model, the array (0.5, 0.15, 0.2, 0.1, 0.05) can be changed to (0.03, 0.1, 0.2, 0.17, 0.5) at 1000 loci in 6,043, 2,108, or 1,408 generations, depending on whether the maximum fitness is five, 100, or 1000. Finally, we note that it is possible to substitute one array for another while continuously favoring heterozygotes.

Alleles↗

Human facial expressions as adaptations: Evolutionary questions in facial expression research.

The importance of the face in social interaction and social intelligence is widely recognized in anthropology. Yet the adaptive functions of human facial expression remain largely unknown. An evolutionary model of human facial expression as behavioral adaptation can be constructed, given the current knowledge of the phenotypic variation, ecological contexts, and fitness consequences of facial behavior. Studies of facial expression are available, but results are not typically framed in an evolutionary perspective. This review identifies the relevant physical phenomena of facial expression and integrates the study of this behavior with the anthropological study of communication and sociality in general. Anthropological issues with relevance to the evolutionary study of facial expression include: facial expressions as coordinated, stereotyped behavioral phenotypes, the unique contexts and functions of different facial expressions, the relationship of facial expression to speech, the value of facial expressions as signals, and the relationship of facial expression to social intelligence in humans and in nonhuman primates. Human smiling is used as an example of adaptation, and testable hypotheses concerning the human smile, as well as other expressions, are proposed.

Adaptation, Biological↗

[Central nervous system control of micturition in patients with bladder dysfunctions in comparison with healthy control probands. An electrophysiological study].

PURPOSE: The pontine micturition center plays a central role in regulating the micturition reflex, but the precise neural mechanisms are unclear. The cerebral cortex is involved in coordinating micturition but there is little knowledge on specific evolutionary higher brain regions. The present study aimed to investigate whether cortical activation during micturition can be demonstrated by EEG power spectra patterns and to explore whether specific cortical regions involved in the interaction of inhibition and release during the micturition reflex can be discerned. We also aimed to test whether intravesical electrostimulation (IVES) therapy in patients with micturition disorders has an effect on patterns of cortical activity. METHODS: The healthy control group was divided into those who were able to void when requested (6 women, 12 men) and those who were not (8 women, 10 men). These subgroups were compared separately with the 14 patients before and after IVES for voiding dysfunction. Following IVES all patients were able to void spontaneously. Mean age of the patients and healthy volunteers was 52 and 30 years, respectively. At the beginning of the study all subjects had a bladder volume of approximately 250 mL as measured by sonography. The EEG was obtained at rest and during the attempt to void. In the patients' group EEG was obtained before IVES treatment and at the day of the last stimulation. The measurement period lasted about 6 minutes. At the beginning of the recording the proband was asked to close his/her eyes. During the resting period after 1 minute the patient was asked to open his/her eyes. After 10 seconds he/she was asked to close his/her eyes again. Then, with eyes still closed, the patient was asked to void. During the entire EEG recording the patient was seated in a comfortable, electrically isolated chair in a darkened room and separated from the examiner by a partition. The subject was asked to relax and not move his/her eyes. The EEG was recorded from the 19 standard points (10-20 System) versus an averaged mastoid electrode with a gold-plated cup electrode (Glass). An EOG was recorded simultaneously to register eye artefacts. The amplification chain was calibrated with a 10-Hz 100-microVss sinus signal generated with a biosignal amplifier. The transitional resistances of all EEG channels were less than 5 kOhm and established as soon as possible. EEG and EOG signals were amplified and recorded with a B.E.S.T. Brain Mapping System. The recording frequency was 256 Hz and the resolution of the analog digital conversion was 12 bit. A high pass and a low pass filter were set to 0.53 Hz and 70 Hz, respectively. All recordings were inspected visually before computer analysis. Artefacts were marked and excluded from the further analysis. None of the EEG recordings showed clinical abnormalities. As expected, the EEGs during voiding attempts showed some muscle potentials and slow motion artefacts. For each subject two artefact-free resting segments of about 20 seconds, one from the resting phase and one from the voiding attempt, were defined by hand for automated analysis. Relative power spectra (microV2) were calculated for the defined segments. From the spectra the relative alpha band power (7.5-13.0 Hz) was calculated for each subject for rest and voiding. Group (patients vs. voiding probands vs. probands unable to void) and sex were independent variables. The alpha power of the 17 electrode positions of the 10-20 system (without Fp1 and Fp2) during rest and attempted voiding were repeated measurement variables. The frontopolar electrode was not used because of its susceptibility to artefacts. The number of dependent variables was due to the explorative nature of the study. With interactions of variables with more than two factor levels a Greenhouse-Geisser correction was performed. Interactions were subjected to contrast analysis and Newman-Keuls-Post tests. RESULTS: Significant effects were seen for BEDINGUNG (

Cerebral Cortex↗

Cyclin F disruption compromises placental development and affects normal cell cycle execution.

Human cyclin F was originally isolated as a cDNA capable of suppressing the temperature sensitivity of a Saccharomyces cerevisiae cdc4-1 mutant. Its tightly regulated expression and high conservation in the evolutionary progression from amphibians to mammals suggest that it coordinates the timing of a critical cell cycle event. The fact that it contains an F box and can form an SCF (Skp1-Cul1/Cdc53-F-box) complex in vivo further suggests that it may also function in proteolysis. To investigate the role of cyclin F in vivo, we generated mice deficient for cyclin F and conditionally deficient mice as well as mouse embryonic fibroblasts (MEFs) conditionally deficient for cyclin F. Heterozygous animals are normal and fertile, but CycF(-/-) animals, with a myriad of developmental anomalies due in large part to failures in yolk sac and chorioallantoic placentation, die around embryonic day 10.5. Tissue-specific deletion of cyclin F revealed that it was not required for the development and function of a number of different embryonic and adult tissues. In contrast, MEFs lacking cyclin F, while viable, do exhibit cell cycle defects, including reduced population-doubling time and a delay in cell cycle reentry from quiescence, indicating that cyclin F plays a role in cell cycle regulation.

Animals↗

Energetic considerations of ciliary beating and the advantage of metachronal coordination.

The internal mechanism of cilia is among the most ancient biological motors on an evolutionary scale. It produces beat patterns that consist of two phases: during the effective stroke, the cilium moves approximately as a straight rod, and during the recovery stroke, it rolls close to the surface in a tangential motion. It is commonly agreed that these two phases are designed for efficient functioning: the effective stroke encounters strong viscous resistance and generates thrust, whereas the recovery stroke returns the cilium to starting position while avoiding viscous resistance. Metachronal coordination between cilia, which occurs when many of them beat close to each other, is believed to be an autonomous result of the hydrodynamical interactions in the system. Qualitatively, metachronism is perceived as a way for reducing the energy expenditure required for beating. This paper presents a quantitative study of the energy expenditure of beating cilia, and of the energetic significance of metachronism. We develop a method for computing the work done by model cilia that beat in a viscous fluid. We demonstrate that for a single cilium, beating in water, the mechanical work done during the effective stroke is approximately five times the amount of work done during the recovery stroke. Investigation of multicilia configurations shows that having neighboring cilia beat metachronally is energetically advantageous and perhaps even crucial for multiciliary functioning. Finally, the model is used to approximate the number of dynein arm attachments that are likely to occur during the effective and recovery strokes of a beat cycle, predicting that almost all of the available dynein arms should participate in generating the motion.

Cilia↗

Comparative metal binding and genomic analysis of the avian (chicken) and mammalian metallothionein.

Chicken metallothionein (ckMT) is the paradigm for the study of metallothioneins (MTs) in the Aves class of vertebrates. Available literature data depict ckMT as a one-copy gene, encoding an MT protein highly similar to mammalian MT1. In contrast, the MT system in mammals consists of a four-member family exhibiting functional differentiation. This scenario prompted us to analyse the apparently distinct evolutionary patterns followed by MTs in birds and mammals, at both the functional and structural levels. Thus, in this work, the ckMT metal binding abilities towards Zn(II), Cd(II) and Cu(I) have been thoroughly revisited and then compared with those of the mammalian MT1 and MT4 isoforms, identified as zinc- and copper-thioneins, respectively. Interestingly, a new mechanism of MT dimerization is reported, on the basis of the coordinating capacity of the ckMT C-terminal histidine. Furthermore, an evolutionary study has been performed by means of in silico analyses of avian MT genes and proteins. The joint consideration of the functional and genomic data obtained questions the two features until now defining the avian MT system. Overall, in vivo and in vitro metal-binding results reveal that the Zn(II), Cd(II) and Cu(I) binding abilities of ckMT lay between those of mammalian MT1 and MT4, being closer to those of MT1 for the divalent metal ions but more similar to those of MT4 for Cu(I). This is consistent with a strong functional constraint operating on low-copy number genes that must cope with differentiating functional limitation. Finally, a second MT gene has been identified in silico in the chicken genome, ckMT2, exhibiting all the features to be considered an active coding region. The results presented here allow a new insight into the metal binding abilities of warm blooded vertebrate MTs and their evolutionary relationships.

Animals↗

The PIN auxin efflux facilitators: evolutionary and functional perspectives.

It is widely believed that the PIN proteins are crucial for proper cellular coordination. Since the analysis of the Arabidopsis pin-formed mutant in 1991, and the subsequent cloning of AtPIN1, a further seven members of the family have been discovered. Here, we present an overview of this family of auxin efflux facilitators in monocot and dicot plants, summarizing their evolutionary history, expression profiles and, where appropriate, relating them to protein function.

Arabidopsis↗