Petrogenetic evolution of Uralian-Alaskan-type mafic-ultramafic complexes in the southern and middle Urals, Russia

The PhD thesis at hand consists of three parts and describes the petrogenetic evolution of Uralian-Alaskan-type mafic ultramafic complexes in the Ural Mountains, Russia. Uralian-Alaskan-type mafic-ultramafic complexes are recognized as a distinct class of intrusions. Characteristic petrologic features are the concentric zonation of a central dunite body grading outward into wehrlite, clinopyroxenite and gabbro, the absence of orthopyroxene and frequently occurring platinum group element (PGE) mineralization. In addition, the presence of ferric iron-rich spinel discriminates Uralian-Alaskan-type complexes from most other mafic ultramafic rock assemblages. The studied Uralian-Alaskan-type complexes (Nizhnii Tagil, Kytlym and Svetley Bor) belong to the southern part of a 900 km long, N–S-trending chain of similar intrusions between the Main Uralian Fault to the west and the Serov-Mauk Fault to the east. The first chapter of this thesis studies the evolution of the ultramafic rocks tracing the compositional variations of rock forming and accessory minerals. The comparison of the chemical composition of olivine, clinopyroxene and chromian spinel from the Urals with data from other localities indicates that they are unique intrusions having a characteristic spinel and clinopyroxene chemistry. Laser ablation-ICPMS (LA-ICPMS ) analyses of trace element concentrations in clinopyroxene are used to calculate the composition of their parental melt which is characterized by enriched LREE (0.5-5.2 prim. mantle) and other highly incompatible elements (U, Th, Ba, Rb) relative to the HREE (0.25-2.0 prim. mantle). A subduction-related geotectonic setting is indicated by a positive anomaly for Sr and negative anomalies for Ti, Zr and Hf. The mineral compositions monitor the evolution of the parental magmas and decipher differences between the studied complexes. In addition, the observed variation in LREE/HREE (for example La/Lu = 2-24) can be best explained with the model of an episodically replenished and erupted open magma chamber system with the extensive fractionation of olivine, spinel and clinopyroxene. The data also show that ankaramites in a subduction-related geotectonic setting could represent parental magmas of Uralian-Alaskan-type complexes. The second chapter of the thesis discusses the chemical variation of major and trace elements in rock-forming minerals of the mafic rocks. Electron microprobe and LA-ICPMS analyses are used to quantitatively describe the petrogenetic relationship between the different gabbroic lithologies and their genetic link to the ultramafic rocks. The composition of clinopyroxene identifies the presence of melts with different trace element abundances on the scale of a thin section and suggests the presence of open system crustal magma chambers. Even on a regional scale the large variation of trace element concentrations and ratios in clinopyroxene (e.g. La/Lu = 3-55) is best explained by the interaction of at least two fundamentally different magma types at various stages of fractionation. This requires the existence of a complex magma chamber system fed with multiple pulses of magmas from at least two different coeval sources in a subduction-related environment. One source produces silica saturated Island arc tholeiitic melts. The second source produces silica undersaturated, ultra-calcic, alkaline melts. Taken these data collectively, the mixing of the two different parental magmas is the dominant petrogenetic process explaining the observed chemical variations. The results further imply that this is an intrinsic feature of Uralian-Alaskan-type complexes and probably of many similar mafic-ultramafic complexes world-wide. In the third chapter of this thesis the major element composition of homogeneous and exsolved spinel is used as a petrogenetic indicator. Homogeneous chromian spinel in dunites and wehrlites monitors the fractionation during the early stages of the magma chamber and the onset of clinopyroxene fractionation as well as the reaction of spinel with interstitial liquid. Exsolved spinel is present in mafic and ultramafic rocks from all three studied complexes. Its composition lies along a solvus curve which defines an equilibrium temperature of 600°C, given that spinel coexists with olivine. This temperature is considered to be close to the temperature of the host rocks into which the studied Uralian-Alaskan-type complexes intruded. The similarity of the exsolution temperatures in the different complexes over a distance of several hundred kilometres implies a regional tectonic event that terminated the exsolution process. This event is potentially associated with the final exhumation of the Uralian-Alaskan-type complexes along the Main Uralian Fault and the Serov-Mauk Fault in the Uralian fold belt.

Thermochronological evolution of the Eastern Pontides and the Eastern Anatolian Plateau and NW Lesser Caucasus (Turkey, Georgia, Armenia)

The analysis of apatite fission tracks is applied to the study of the syn- and post-collisional thermochronological evolution of a vast area that includes the Eastern Pontides, their continuation in the Lesser Caucasus of Georgia (Adjara-Trialeti zone) and northern Armenia, and the eastern Anatolian Plateau. The resulting database is then integrated with the data presented by Okay et al. (2010) for the Bitlis Pütürge Massif, i.e. the western portion of the Bitlis-Zagros collision zone between Arabia and Eurasia. The mid-Miocene exhumation episode along the Black Sea coast and Lesser Caucasus of Armenia documented in this dissertation mirrors the age of collision between the Eurasian and Arabian plates along the Bitlis suture zone. We argue that tectonic stresses generated along the Bitlis collision zone were transmitted northward across eastern Anatolia and focused (i) at the rheological boundary between the Anatolian continental lithosphere and the (quasi)oceanic lithosphere of the Black Sea, and (ii) along major pre-existing discontinuities like the Sevan-Akera suture zone.The integration of both present-day crustal dynamics (GPS-derived kinematics and distribution of seismicity) and thermochronological data presented in this paper provides a comparison between short- and long-term deformation patterns for the entire eastern Anatolia-Transcaucasian region. Two successive stages of Neogene deformation of the northern foreland of the Arabia-Eurasia collision zone can be inferred. (i) Early and Middle Miocene: continental deformation was concentrated along the Arabia-Eurasia (Bitlis) collision zone but tectonic stress was also transferred northward across eastern Anatolia, focusing along the eastern Black Sea continent-ocean rheological transition and along major pre-existing structural discontinuities. (ii) Since Late-Middle Miocene time the westward translation of Anatolia and the activation of the North and Eastern Anatolian Fault systems have reduced efficient northward stress transfer.

Neoproterozoic to middle Palaeozoic evolution of the Central Asian Orogenic Belt in south-central Mongolia: chronological and geochemical perspectives

Mongolia occupies a central position within the eastern branch of the large accretionary Central Asian Orogenic Belt (CAOB) or Altaids. The present work aims to outline the geodynamic environment and possible evolution of this part of the eastern CAOB, predominantly from the Cambrian to the middle Palaeozoic. The investigation primarily focussed on zircon geochronology as well as whole-rock geochemical and Sm–Nd isotopic analyses for a variety of metaigneous rocks from the southern Hangay and Gobi-Altai regions in south-central Mongolia. The southern slope of the Hangay Mountains in central Mongolia exposes a large NWSE-trending middle Neoproterozoic ophiolitic complex (c. 650 Ma), which is tectonically integrated within an accretionary complex developed between the Precambrian Baydrag and Hangay crustal blocks. Formation of the entire accretionary system along the north-eastern margin of the Baydrag block mainly occurred during the early Cambrian, but convergence within this orogenic zone continued until the early Ordovician, because of on-going southward subduction-accretion of the Baydrag block. An important discovery is the identification of a late Mesoproterozoic to early Neoproterozoic belt within the northern Gobi-Altai that was reworked during the late Cambrian and throughout the late Ordovician/Devonian. Early Silurian low-grade mafic and felsic metavolcanic rocks from the northern Gobi-Altai display subduction-related geochemical features and highly heterogeneous Nd isotopic compositions, which suggest an origin at a mature active continental margin. Early Devonian protoliths of granodioritic and mafic gneisses from the southern Gobi-Altai display geochemical and Nd isotopic compositions compatible with derivation and evolution from predominantly juvenile crustal and mantel sources and these rocks may have been emplaced within the outboard portion of the late Ordovician/early Silurian active continental margin. Moreover, middle Devonian low-grade metavolcanic rocks from the southwestern Gobi-Altai yielded geochemical and Nd isotopic data consistent with emplacement in a transitional arc-backarc setting. The combined U–Pb zircon ages and geochemical data obtained from the Gobi-Altai region suggest that magmatism across an active continental margin migrated oceanwards through time by way of subduction zone retreat throughout the Devonian. Progressive extension of the continental margin was associated with the opening of a backarc basin and culminated in the late Devonian with the formation of a Japan-type arc front facing a southward open oceanic realm (present-day coordinates).

Early steps in ventral nerve cord development in chelicerates and myriapods and formation of brain compartments in spiders

The central point of this work is the investigation of neurogenesis in chelicerates and myriapods. By comparing decisive mechanisms in neurogenesis in the four arthropod groups (Chelicerata, Crustacea, Insecta, Myriapoda) I was able to show which of these mechanisms are conserved and which developmental modules have diverged. Thereby two processes of embryonic development of the central nervous system were brought into focus. On the one hand I studied early neurogenesis in the ventral nerve cord of the spiders Cupiennius salei and Achaearanea tepidariorum and the millipede Glomeris marginata and on the other hand the development of the brain in Cupiennius salei.rnWhile the nervous system of insects and crustaceans is formed by the progeny of single neural stem cells (neuroblasts), in chelicerates and myriapods whole groups of cells adopt the neural cell fate and give rise to the ventral nerve cord after their invagination. The detailed comparison of the positions and the number of the neural precursor groups within the neuromeres in chelicerates and myriapods showed that the pattern is almost identical which suggests that the neural precursors groups in these arthropod groups are homologous. This pattern is also very similar to the neuroblast pattern in insects. This raises the question if the mechanisms that confer regional identity to the neural precursors is conserved in arthropods although the mode of neural precursor formation is different. The analysis of the functions and expression patterns of genes which are known to be involved in this mechanism in Drosophila melanogaster showed that neural patterning is highly conserved in arthropods. But I also discovered differences in early neurogenesis which reflect modifications and adaptations in the development of the nervous systems in the different arthropod groups.rnThe embryonic development of the brain in chelicerates which was investigated for the first time in this work shows similarities but also some modifications to insects. In vertebrates and arthropods the adult brain is composed of distinct centres with different functions. Investigating how these centres, which are organised in smaller compartments, develop during embryogenesis was part of this work. By tracing the morphogenetic movements and analysing marker gene expressions I could show the formation of the visual brain centres from the single-layered precheliceral neuroectoderm. The optic ganglia, the mushroom bodies and the arcuate body (central body) are formed by large invaginations in the peripheral precheliceral neuroectoderm. This epithelium itself contains neural precursor groups which are assigned to the respective centres and thereby build the three-dimensional optical centres. The single neural precursor groups are distinguishable during this process leading to the assumption that they carry positional information which might subdivide the individual brain centres into smaller functional compartments.rn

Hydrodynamical simulations of early-type galaxies: effects of galaxy shape and stellar dynamics on hot coronae

Early-Type galaxies (ETGs) are embedded in hot (10^6-10^7 K), X-ray emitting gaseous haloes, produced mainly by stellar winds and heated by Type Ia supernovae explosions, by the thermalization of stellar motions and occasionally by the central super-massive black hole (SMBH). In particular, the thermalization of the stellar motions is due to the interaction between the stellar and the SNIa ejecta and the hot interstellar medium (ISM) already residing in the ETG. A number of different astrophysical phenomena determine the X-ray properties of the hot ISM, such as stellar population formation and evolution, galaxy structure and internal kinematics, Active Galactic Nuclei (AGN) presence, and environmental effects. With the aid of high-resolution hydrodynamical simulations performed on state-of-the-art galaxy models, in this Thesis we focus on the effects of galaxy shape, stellar kinematics and star formation on the evolution of the X-ray coronae of ETGs. Numerical simulations show that the relative importance of flattening and rotation are functions of the galaxy mass: at low galaxy masses, adding flattening and rotation induces a galactic wind, thus lowering the X-ray luminosity; at high galaxy masses the angular momentum conservation keeps the central regions of rotating galaxies at low density, whereas in non-rotating models a denser and brighter atmosphere is formed. The same dependence from the galaxy mass is present in the effects of star formation (SF): in light galaxies SF contributes to increase the spread in Lx, while at high galaxy masses the halo X-ray properties are marginally sensitive to SF effects. In every case, the star formation rate at the present epoch quite agrees with observations, and the massive, cold gaseous discs are partially or completely consumed by SF on a time-scale of few Gyr, excluding the presence of young stellar discs at the present epoch.

Genetic dynamics across early life stages in the tropical tree Prunus africana (Rosaceae)

In many plant species, the genetic template of early life-stages is formed by animal-mediated pollination and seed dispersal and has profound impact on further recruitment and population dynamics. Understanding the impact of pollination and seed dispersal on genetic patterns is a central issue in plant population biology. In my thesis, I investigated (i) contemporary dispersal and gene flow distances as well as (ii) genetic diversity and spatial genetic structure (SGS) across subsequent recruitment stages in a population of the animal-pollinated and dispersed tree Prunus africana in Kakamega Forest, West Kenya. Using microsatellite markers and parentage analyses, I inferred distances of pollen dispersal (father-to-mother), seed dispersal/maternal gene flow (mother-to-offspring) as well as paternal gene flow (father-to-offspring) for four early life stages of the species (seeds and fruits, current year seedlings, seedlings ≤ 3yr, seedlings > 3yr). Distances of pollen and seed dispersal as well as paternal gene flow were significantly shorter than expected from the spatial arrangement of trees and sampling plots. They were not affected by the density of conspecific trees in the surrounding. At the propagule stage, mean pollen dispersal distances were considerably (23-fold) longer than seed dispersal distances, and paternal gene flow distances exceeded maternal gene flow by a factor of 25. Seed dispersal distances were remarkably restricted, potentially leading to a strong initial SGS. The initial genetic template created by pollination and seed dispersal was extensively altered during later recruitment stages. Potential Janzen-Connell effects led to markedly increasing distances between offspring and both parental trees in older life stages. This showed that distance and density-dependent mortality factors are not exclusively related to the mother tree, but also to the father. Across subsequent recruitment stages, the pollen to seed dispersal ratio and the paternal to maternal gene flow ratio dropped to 2.1 and 3.4, respectively, in seedlings > 3yr. The relative changes in effective pollen dispersal, seed dispersal, and paternal gene flow distances across recruitment stages elucidate the mechanisms affecting the contribution of the two processes pollen and seed dispersal to overall gene flow. Using the same six microsatellite loci, I analyzed genetic diversity and SGS across five life stages, from seed rain to adults. Levels of genetic diversity within the studied P. africana population were comparable to other Prunus species and did not vary across life stages. In congruence with the short seed dispersal distances, I found significant SGS in all life stages. SGS decreased from seed and early seedling stages to older juvenile stages, and it was higher in adults than in late juveniles of the next generation. A comparison of the data with direct assessments of contemporary gene flow patterns indicate that distance- or density-dependent mortality, potentially due to Janzen-Connell effects, led to the initial decrease in SGS. Intergeneration variation in SGS could have been driven by variation in demographic processes, the effect of overlapping generations, and local selection processes. Overall, my study showed that complex sequential processes during recruitment contribute to the spatial genetic structure of tree populations. It highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal-mediated pollen and seed dispersal on spatial population dynamics and genetic patterns of trees.

The dynamics of early-type galaxies as a tool to understand their hot coronae and their IMF

Dynamical models of galaxies are a powerful tool to study and understand several astrophysical problems related to galaxy formation and evolution. This thesis is focussed on a particular type of dynamical models, that are widely used in literature, and are based on the solution of the Jeans equations. By means of a numerical Jeans solver code, developed on purpose and able to build state-of-the-art advanced axisymmetric galaxy models, two of the main currently investigated issues in the field of research of early-type galaxies (ETGs) are addressed. The first topic concerns the hot and X-ray emitting gaseous coronae that surround ETGs. The main goal is to explain why flat and rotating galaxies generally exhibit haloes with lower gas temperatures and luminosities with respect to rounder and velocity dispersion supported systems. The second astrophysical problem addressed concerns instead the stellar initial mass function (IMF) of ETGs. Nowadays, this is a very controversial issue due to a growing number of works on ETGs, based on different and independent techniques, that show evidences of a systematic variation of the IMF normalization as a function of galaxy velocity dispersion or mass. These studies are changing the previous opinion that the IMF of ETGs was the same as that of spiral galaxies, and hence universal throughout the whole large family of galaxies.

Low-temperature thermochronological evolution of the Menderes and Alanya massifs (Turkey)

The application of two low-temperature thermochronometers [fission-track analysis and (U-Th)/He analyses, both on apatite] to various tectonostratigraphic units of the Menderes and Alanya Massifs of Turkey has provided significant new constraints to the understanding of their structural evolution. The Menderes Massif of western Anatolia is one of the largest metamorphic core complexes on Earth. The integration of the geochronometric dataset presented in this dissertation with preexisting ones from the literature delineates three groups of samples within the Menderes Massif. In the northern and southern region the massif experienced a Late Oligocene-Early Miocene tectonic denudation and surface uplift; whereas data from the central region are younger, with most ages ranging between the Middle-Late Miocene. The results of this study are consistent with the interpretation for a symmetric exhumation of the Menderes Massif. The Alanya Massif of SW Anatolia presents a typical nappe pile consisting of thrust sheets with contrasting metamorphic histories. Petrological and geochronological data clearly indicate that the tectonometamorphic evolution Alanya started from Late Cretaceous with the northward subduction of an ‘Alanya ocean’ under the Tauride plate. As an effect of the closure of the İzmir–Ankara–Erzincan ocean, northward backthrusting during the Paleocene-Early Eocene created the present stacking order. Apatite fission-track ages from this study range from 31.8 to 26.8 Ma (Late Rupelian-Early Chattian) and point to a previously unrecognized mid-Oligocene cooling/exhumation episode. (U-Th)/He analysis on zircon crystals obtained from the island of Cyprus evidentiate that the Late Cretaceous trondhjemites of the Troodos Massif not recorded a significant cooling event. Instead results for the Late Triassic turbiditic sandstones of the Vlambouros Formation show that the Mamonia mélange was never buried enough to reach the closure temperature of the ZHe radiometric system (ca. 200°C), thus retaining the Paleozoic signature of a previous sedimentary cycle.

Extracting evolutionary information from the spectral decomposition of early-type galaxies.

Holding the major share of stellar mass in galaxies and being also old and passively evolving, early-type galaxies (ETGs) are the primary probes in investigating these various evolution scenarios, as well as being useful means to provide insights on cosmological parameters. In this thesis work I focused specifically on ETGs and on their capability in constraining galaxy formation and evolution; in particular, the principal aims were to derive some of the ETGs evolutionary parameters, such as age, metallicity and star formation history (SFH) and to study their age-redshift and mass-age relations. In order to infer galaxy physical parameters, I used the public code STARLIGHT: this program provides a best fit to the observed spectrum from a combination of many theoretical models defined in user-made libraries. the comparison between the output and input light-weighted ages shows a good agreement starting from SNRs of ∼ 10, with a bias of ∼ 2.2% and a dispersion 3%. Furthermore, also metallicities and SFHs are well reproduced. In the second part of the thesis I performed an analysis on real data, starting from Sloan Digital Sky Survey (SDSS) spectra. I found that galaxies get older with cosmic time and with increasing mass (for a fixed redshift bin); absolute light-weighted ages, instead, result independent from the fitting parameters or the synthetic models used. Metallicities, instead, are very similar from each other and clearly consistent with the ones derived from the Lick indices. The predicted SFH indicates the presence of a double burst of star formation. Velocity dispersions and extinctiona are also well constrained, following the expected behaviours. As a further step, I also fitted single SDSS spectra (with SNR∼ 20), to verify that stacked spectra gave the same results without introducing any bias: this is an important check, if one wants to apply the method at higher z, where stacked spectra are necessary to increase the SNR. Our upcoming aim is to adopt this approach also on galaxy spectra obtained from higher redshift Surveys, such as BOSS (z ∼ 0.5), zCOSMOS (z 1), K20 (z ∼ 1), GMASS (z ∼ 1.5) and, eventually, Euclid (z 2). Indeed, I am currently carrying on a preliminary study to estabilish the applicability of the method to lower resolution, as well as higher redshift (z 2) spectra, just like the Euclid ones.

On the Luminous and Dark Matter Distribution in Early-Type Galaxies

The way mass is distributed in galaxies plays a major role in shaping their evolution across cosmic time. The galaxy's total mass is usually determined by tracing the motion of stars in its potential, which can be probed observationally by measuring stellar spectra at different distances from the galactic centre, whose kinematics is used to constrain dynamical models. A class of such models, commonly used to accurately determine the distribution of luminous and dark matter in galaxies, is that of equilibrium models. In this Thesis, a novel approach to the design of equilibrium dynamical models, in which the distribution function is an analytic function of the action integrals, is presented. Axisymmetric and rotating models are used to explain observations of a sample of nearby early-type galaxies in the Calar Alto Legacy Integral Field Area survey. Photometric and spectroscopic data for round and flattened galaxies are well fitted by the models, which are then used to get the galaxies' total mass distribution and orbital anisotropy. The time evolution of massive early-type galaxies is also investigated with numerical models. Their structural properties (mass, size, velocity dispersion) are observed to evolve, on average, with redshift. In particular, they appear to be significantly more compact at higher redshift, at fixed stellar mass, so it is interesting to investigate what drives such evolution. This Thesis focuses on the role played by dark-matter haloes: their mass-size and mass-velocity dispersion correlations evolve similarly to the analogous correlations of ellipticals; at fixed halo mass, the haloes are more compact at higher redshift, similarly to massive galaxies; a simple model, in which all the galaxy's size and velocity-dispersion evolution is due to the cosmological evolution of the underlying halo population, reproduces the observed size and velocity-dispersion of massive compact early-type galaxies up to redshift of about 2.

Active processes, morphology, and dynamics of icy debris fans: Landform evolution along rapidly degrading escarpments in alpine regions undergoing recent deglaciation

In the past few decades the impacts of climate warming have been significant in alpine glaciated regions. Many valley glaciers formerly linked as distributary glaciers to high-level icecaps have decoupled at their icefalls, exposing major escarpments and generating a suite of dynamic landforrns dominated by mass wasting. Ice-dominated landforms, here termed icy debris fans, develop rapidly by ice avalanching, rockfall, and icy debris flow. Field-based reconnaissance studies at two alpine settings, the Wrangell Mountains of Alaska and the Southern Alps of New Zealand, provide a preliminary morphogenetic model of spatial and temporal evolution of icy debris fans in a range of alpine settings. The influence of these processes on landform evolution is largely unrecognized in the literature dealing with post-glacial landform adjustment known as the paraglacial. A better understanding of these dynamic processes will be increasingly important because of the extreme geohazards characterizing these areas. Our field studies show that after glacier decoupling, icy debris fans begin to form along the base of bedrock escarpments at the mouths of catchments and prograde over valley glaciers. The presence of a distinct catchment, apex, and fan morphology distinguishes these landforms from other landforms common in periglacial hillslope settings receiving abundant clastic debris and ice. Ice avalanching is the most abundant process involved in icy debris fan formation. Fans developed below weakly incised catchments are dominated by ice avalanching and are composed primarily of ice with minor lithic detritus. Typically, avalanches fall into the fan catchments where sediments transform into grainflows that flow onto the fans. Once on the fans, avalanche deposits ablate rapidly, flattening and concentrating lithic fragments at the surface. Icy debris fans may become thick enough to become glaciers with splay crevasse systems. Fans developed below larger, more complex catchments are composed of higher proportions of lithic detritus resulting from temporary storage of ice and lithic detritus deposits within the catchment. Episodic outbursts of meltwater from the icecap may mix with the stored sediments and mobilize icy debris flows (mixture of ice and lithic clasts) onto the fans. Our observations indicate that the entire evolutionary cycle of icy debris fans probably occurs during an early paraglacial interval (i.e., decades to 100 years). Observations comparing avalanche frequency, volume, and fan morphologic evolution at the Alaska site between 2006 and 2010 illustrate complex response between icy debris fans even within the same cirque - where one fan may be growing while others are downwasting because of differences in ice supply controlled by their respective catchments and icecap contributions. As ice supply from the icecap diminishes through time, icy debris fans rapidly downwaste and eventually evolve into talus cones that receive occasional but ephemeral ice avalanches.

Holocentric chromosomes: convergent evolution, meiotic adaptations, and genomic analysis

In most eukaryotes, the kinetochore protein complex assembles at a single locus termed the centromere to attach chromosomes to spindle microtubules. Holocentric chromosomes have the unusual property of attaching to spindle microtubules along their entire length. Our mechanistic understanding of holocentric chromosome function is derived largely from studies in the nematode Caenorhabditis elegans, but holocentric chromosomes are found over a broad range of animal and plant species. In this review, we describe how holocentricity may be identified through cytological and molecular methods. By surveying the diversity of organisms with holocentric chromosomes, we estimate that the trait has arisen at least 13 independent times (four times in plants and at least nine times in animals). Holocentric chromosomes have inherent problems in meiosis because bivalents can attach to spindles in a random fashion. Interestingly, there are several solutions that have evolved to allow accurate meiotic segregation of holocentric chromosomes. Lastly, we describe how extensive genome sequencing and experiments in nonmodel organisms may allow holocentric chromosomes to shed light on general principles of chromosome segregation.

Bacillus anthracis: Molecular taxonomy, population genetics, phylogeny and patho-evolution

Bacillus anthracis, the etiological agent of anthrax, manifests a particular bimodal lifestyle. This bacterial species alternates between short replication phases of 20-40 generations that strictly require infection of the host, normally causing death, interrupted by relatively long, mostly dormant phases as spores in the environment. Hence, the B. anthracis genome is highly homogeneous. This feature and the fact that strains from nearly all parts of the world have been analysed for canonical single nucleotide polymorphisms (canSNPs) and variable number tandem repeats (VNTRs) has allowed the development of molecular epidemiological and molecular clock models to estimate the age of major diversifications in the evolution of B. anthracis and to trace the global spread of this pathogen, which was mostly promoted by movement of domestic cattle with settlers and by international trade of contaminated animal products. From a taxonomic and phylogenetic point of view, B. anthracis is a member of the Bacillus cereus group. The differentiation of B. anthracis from B. cereus sensu strict, solely based on chromosomal markers, is difficult. However, differences in pathogenicity clearly differentiate B. anthracis from B. cereus and are marked by the strict presence of virulence genes located on the two virulence plasmids pXO1 and pXO2, which both are required by the bacterium to cause anthrax. Conversely, anthrax-like symptoms can also be caused by organisms with chromosomal features that are more closely related to B. cereus, but which carry these virulence genes on two plasmids that largely resemble the B. anthracis virulence plasmids. (C) 2011 Elsevier B.V. All rights reserved.