Late Pleistocene‐Holocene Slip Rates in the Northwestern Zagros Mountains (Kurdistan Region of Iraq) Derived From Luminescence Dating of River Terraces and Structural Modeling

Abstract A significant amount of the ongoing shortening between the Eurasian and Arabian plates is accommodated within the Zagros Fold‐Thrust Belt. However, the spatial and temporal distribution of active shortening within the belt, especially in its NW part, is not yet well constrained. We determined depositional ages of uplifted river terraces crossing the belt along the Greater Zab River using luminescence dating. Kinematic modeling of the fault‐related fold belt was then used to calculate long‐term slip rates during the Late Pleistocene to Holocene. Our results provide new insight into the rates of active faulting and folding in the area. The Zagros Mountain Front Fault accommodates about 1.46 ± 0.60 mm a −1 of slip, while a more external basement fault further to the SW accommodates less than 0.41 ± 0.16 mm a −1 . Horizontal slip rates related to detachment folding of two anticlines within the Zagros Foothills are 0.40 ± 0.10 and 1.24 ± 0.36 mm a −1 . Basement thrusting and thickening of the crust are restricted to the NE part of the Zagros belt. This is also reflected in the regional topography and in the distribution of uplifted terraces. In the southwestern part, the deformation is limited mainly to folding and thrusting of the sedimentary cover above a Triassic basal detachment. In the NE, deformation is associated with slip on basement thrusts. Our study sheds light on the distribution of shortening in the Zagros Mountains and helps to understand the regional tectonic system. Our results may be the foundation for a better seismic hazard assessment of the entire area.Plain Language Summary In active mountain belts, river terraces found above the present‐day river level can be indicative of differences in uplift rates due to the thickening, faulting, and folding processes in the Earth's crust. These processes, driven by the motion of tectonic plates, are responsible for the formation of mountain belts. Here, we took sediment samples from uplifted river terraces along the Greater Zab River that crosses the Zagros Mountains in the Kurdistan Region of Iraq. We determined their deposition age using luminescence dating. From their age and elevation, we calculated uplift rates. We built a geometrical model of the fault zones in the area and determined how fast the slip occurs on these faults based on the uplift rates. Our results indicate that there were less than two millimeter per year of slip on these faults on average during the last 60 thousand years. This motion is a result of the convergence between the Arabian and Eurasian plates. With studies like this we can measure how fast fault blocks move, even if they were not associated with large earthquakes in the recent past. This approach helps to better assess the potential earthquake hazard in the area under investigation.Key Points We estimated fault slip rates in the NW Zagros Mountains by luminescence dating of river terraces and structural modeling There is c. 1.46 mm a −1 slip on the Mountain Front Fault and c. 1.64 mm a −1 slip from detachment folding in the NE part of the Foothill Zone Crustal thickening and basement thrusting occur in the NE parts of the Foothill Zone and only cover deformation occur in the SW part

Disruption and pseudoautosomal localization of the major histocompatibility complex in monotremes

The characterization and chromosomal mapping of major histocompatibility complex (MHC)-containing BAC clones from platypus and the short-beaked echidna reveals new insights into the evolution of both the mammalian MHC and monotreme sex chromosomes

Loss of genes implicated in gastric function during platypus evolution

Several genes implicated in food digestion have been deleted or inactivated in platypus. This loss perhaps explains the anatomical and physiological differences in the gastrointestinal tract between monotremes and other vertebrates and provides insights into platypus genome evolution

Flavors of non-random meiotic segregation of autosomes and sex chromosomes

Segregation of chromosomes is a multistep process occurring both at mitosis and meiosis to ensure that daughter cells receive a complete set of genetic information. Critical components in the chromosome segregation include centromeres, kinetochores, components of sister chromatid and homologous chromosomes cohesion, microtubule organizing centres, and spindles. Based on the cytological work in the grasshopper Brachystola, it has been accepted for decades that segregation of homologs at meiosis is fundamentally random. This ensures that alleles on chromosomes have equal chance to be transmitted to progeny. At the same time mechanisms of meiotic drive and an increasing number of other examples of non-random segregation of autosomes and sex chromosomes provide insights into the underlying mechanisms of chromosome segregation but also question the textbook dogma of random chromosome segregation. Recent advances provide a better understanding of meiotic drive as a prominent force where cellular and chromosomal changes allow autosomes to bias their segregation. Less understood are mechanisms explaining observations that autosomal heteromorphism may cause biased segregation and regulate alternating segregation of multiple sex chromosome systems or translocation heterozygotes as an extreme case of non-random segregation. We speculate that molecular and cytological mechanisms of non-random segregation might be common in these cases and that there might be a continuous transition between random and non-random segregation which may play a role in the evolution of sexually antagonistic genes and sex chromosome evolution

The multiple sex chromosomes of platypus and echidna are not completely identical and several share homology with the avian Z.

BACKGROUND: Sex-determining systems have evolved independently in vertebrates. Placental mammals and marsupials have an XY system, birds have a ZW system. Reptiles and amphibians have different systems, including temperature-dependent sex determination, and XY and ZW systems that differ in origin from birds and placental mammals. Monotremes diverged early in mammalian evolution, just after the mammalian clade diverged from the sauropsid clade. Our previous studies showed that male platypus has five X and five Y chromosomes, no SRY, and DMRT1 on an X chromosome. In order to investigate monotreme sex chromosome evolution, we performed a comparative study of platypus and echidna by chromosome painting and comparative gene mapping. RESULTS: Chromosome painting reveals a meiotic chain of nine sex chromosomes in the male echidna and establishes their order in the chain. Two of those differ from those in the platypus, three of the platypus sex chromosomes differ from those of the echidna and the order of several chromosomes is rearranged. Comparative gene mapping shows that, in addition to bird autosome regions, regions of bird Z chromosomes are homologous to regions in four platypus X chromosomes, that is, X1, X2, X3, X5, and in chromosome Y1. CONCLUSION: Monotreme sex chromosomes are easiest to explain on the hypothesis that autosomes were added sequentially to the translocation chain, with the final additions after platypus and echidna divergence. Genome sequencing and contig anchoring show no homology yet between platypus and therian Xs; thus, monotremes have a unique XY sex chromosome system that shares some homology with the avian Z.RIGHTS : This article is licensed under the BioMed Central licence at http://www.biomedcentral.com/about/license which is similar to the 'Creative Commons Attribution Licence'. In brief you may : copy, distribute, and display the work; make derivative works; or make commercial use of the work - under the following conditions: the original author must be given credit; for any reuse or distribution, it must be made clear to others what the license terms of this work are

The emergence of the brain non-CpG methylation system in vertebrates

Mammalian brains feature exceptionally high levels of non-CpG DNA methylation alongside the canonical form of CpG methylation. Non-CpG methylation plays a critical regulatory role in cognitive function, which is mediated by the binding of MeCP2, the transcriptional regulator that when mutated causes Rett syndrome. However, it is unclear whether the non-CpG neural methylation system is restricted to mammalian species with complex cognitive abilities or has deeper evolutionary origins. To test this, we investigated brain DNA methylation across 12 distantly related animal lineages, revealing that non-CpG methylation is restricted to vertebrates. We discovered that in vertebrates, non-CpG methylation is enriched within a highly conserved set of developmental genes transcriptionally repressed in adult brains, indicating that it demarcates a deeply conserved regulatory program. We also found that the writer of non-CpG methylation, DNMT3A, and the reader, MeCP2, originated at the onset of vertebrates as a result of the ancestral vertebrate whole-genome duplication. Together, we demonstrate how this novel layer of epigenetic information assembled at the root of vertebrates and gained new regulatory roles independent of the ancestral form of the canonical CpG methylation. This suggests that the emergence of non-CpG methylation may have fostered the evolution of sophisticated cognitive abilities found in the vertebrate lineage.This work was supported by the Australian Research Council (ARC) Centre of Excellence programme in Plant Energy Biology (grant no. CE140100008). R.L. was supported by a Sylvia and Charles Viertel Senior Medical Research Fellowship, ARC Future Fellowship (no. FT120100862) and Howard Hughes Medical Institute International Research Scholarship. A.d.M. was funded by an EMBO long-term fellowship (no. ALTF 144-2014). J.L.G.-S. was supported by the Spanish government (grant no. BFU2016- 74961-P) and the institutional grant Unidad de Excelencia María de Maeztu (no. MDM-2016-0687). B.V. was supported by the Biomedical Research Council of the Agency for Science, Technology and Research of Singapore. F.G. was supported by an ARC Future Fellowship (no. FT160100267). C.W.R. was supported by an NSF grant (no. IOS-1354898). J.R.E. is an investigator of the Howard Hughes Medical Institute. Genomic data was generated at the Australian Cancer Research Foundation Centre for Advanced Cancer Genomics

Cone visual pigments of monotremes: Filling the phylogenetic gap

We have determined the sequence and genomic organization of the genes encoding the cone visual pigment of the platypus (Ornithorhynchus anatinus) and the echidna (Tachyglossus aculeatus), and inferred their spectral properties and evolutionary pathways. We prepared platypus and echidna retinal RNA and used primers of the middle-wave-sensitive (MWS), long-wave-sensitive (LWS), and short-wave sensitive (SWS1) pigments corresponding to coding sequences that are highly conserved among mammals; to PCR amplify the corresponding pigment sequences. Amplification from the retinal RNA revealed the expression of LWS pigment mRNA that is homologous in sequence and spectral properties to the primate LWS visual pigments. However, we were unable to amplify the mammalian SWS1 pigment from these two species, indicating this gene was lost prior to the echidna-platypus divergence ~21 MYA. Subsequently, when the platypus genome sequence became available, we found an LWS pigment gene in a conserved genomic arrangement that resembles the primate pigment, but, surprisingly we found an adjacent (~20 kb) SWS2 pigment gene within this conserved genomic arrangement. We obtained the same result after sequencing the echidna genes. The encoded SWS2 pigment is predicted to have a wavelength of maximal absorption of about 440 nm, and is paralogous to SWS pigments typically found in reptiles, birds, and fish but not in mammals. This study suggests the locus control region (LCR) has played an important role in the conservation of photo receptor gene arrays and the control of their spatial and temporal expression in the retina in all mammals. In conclusion, a duplication event of an ancestral cone visual pigment gene, followed by sequence divergence and selection gave rise to the LWS and SWS2 visual pigments. So far, the echidna and platypus are the only mammals that share the gene structure of the LWS-SWS2 pigment gene complex with reptiles, birds and fishes.Matthew J. Wakefield, Mark Anderson, Ellen Chang, Ke-Jun Wei, Rajinder Kaul, Jennifer A. Marshall Graves, Frank Grutzner and Samir S. Dee