Evolutionary Appearance Research Articles

Advanced crown‐group Rossellidae (Porifera: Hexactinellida) resembling extant taxa from the Hirnantian (Late Ordovician) Anji Biota

AbstractAmong living hexactinellids (glass sponges), the Rossellidae are one of the most distinctive species‐rich families because of their unique macroscopic characters and, due to the resulting fossil record, are among the most useful for tracing the origins of hexactinellid diversification. Recent discoveries have extended the origin of the total group back to the Ordovician, but these fossils have, so far, all been identified either as stem‐group representatives or as sponges with uncertain interpretation. New material described here from the Hirnantian (latest Ordovician) Anji Biota of Zhejiang, China includes demonstrable crown‐group representatives of the family, which also appear morphologically similar to living genera and may be closely related to them. The new taxa are described as Crateromorpha? (Neopsacas?) macrospicula sp. nov., Pseudanoxycalyx verrucosus gen. et sp. nov., Eorosselloides antiquus gen. et sp. nov. and Archaeaphorme conica gen. et sp. nov. The similarity to modern forms implies extraordinary evolutionary stasis of at least some members of the modern deep‐sea hexactinellid fauna since that time, and suggests that they had an even earlier origin and initial diversification. Possible examples of late Precambrian stem‐group hexactinellids remain ambiguous, potentially implying very rapid evolutionary appearance and divergence of the class during the Cambrian or earlier Ordovician. A possible driver of this evolutionary rate change is the remarkable longevity of modern hexactinellids as an adaptation to cold‐water environments; this potentially offers an explanation for discordances with molecular clock results, which have previously indicated a much later (late Palaeozoic) origin for total‐group Rossellidae.

Two different complement Factor B (Bf) alleles of the orangutan major histocompatibility complex (MHC) are also conserved in chimpanzee and humans showing importance in primate immunity.

MajorHuman Histocompatibility complex (MHC or HLA in humans) has been associated to autoimmune diseases. However, only statistical phenomenological and no pathogenetic description has been reached after decades. This shows that MHC single locus association studies are probably useless for HLA/diseases association. Extended HLA (class I and class II genes) haplotypes should also be studied conjointly with class III or complement alleles (complotypes). Complotypes in humans are defined as alleles belonging to C2, C4 and Bf (Factor B) genes/proteins (class III). Also, the placing of MHC class I and class II genes close together with complement genes from at least birds to humans shows existence of a strong selection to gather conjointly these loci that fight microbes, help self-maintenance and avoid autoimmunity. In this paper we aim to study Bf alleles in primates in order to rise again interest to study the role of Bf alleles together with other MHC genes in their physiopathology and evolution. Orangutan (Pongo pygmaeus, Popy) cell lines RNA from 6 different individuals were retrotranscribed, PCR amplified, cloned and DNA sequenced in order to study Bf alleles. A Bf allele identical to that found in chimpanzee (Patr-Bf*A01) and human (rs641153) was found in two of the six studied orangutans: Popy-Bf*A01 and Popy-Bf*A02. This polymorphism is placed in Factor B codon 32 that defines BF*S and Bf*F proteins in man and produce Leu instead of Arg (Bf*S) or Gln (Bf*F). In addition, each new orangutan allele present synonymous differences with each other at codon 25: Popy-Bf*A01 shows ACG while Popy-Bf*A02 bears ACA, both codifying for Thr. The selection for about 15 million years (time gap of evolutionary appearance between orangutan and hominids) shows the importance of this particular allele conservation in immune and self defense in primates. The complotypes (Bf,C2 and C4 loci) alleles together with other MHC class I and Cass II loci alleles are often transmitted in block to the germinal line: this indicates that all specific alleles from the MHC different loci may work together to accomplish MHC functions. All MHC loci alleles should be studied together to unveil their physiopathology and also maintenance of specific alleles (like the one described in this paper) for so long time in evolution should be further studied in Bf and the other neighbouring complement loci (C2 and C4).

An Orthologics Study of the Notch Signaling Pathway.

The Notch signaling pathway plays a major role in embryological development and in the ongoing life processes of many animals. Its role is to provide cell-to-cell communication in which a Sender cell, bearing membrane-embedded ligands, instructs a Receiver cell, bearing membrane-embedded receptors, to adopt one of two available fates. Elucidating the evolution of this pathway is the topic of this paper, which uses an orthologs approach, providing a comprehensive basis for the study. Using BLAST searches, orthologs were identified for all the 49 components of the Notch signaling pathway. The historical time course of integration of these proteins, as the animals evolved, was elucidated. Insofar as cell-to-cell communication is of relevance only in multicellular animals, it is not surprising that the Notch system became functional only with the evolutionary appearance of Metazoa, the first multicellular animals. Porifera contributed a quarter of the Notch pathway proteins, the Cnidaria brought the total to one-half, but the system reached completion only when humans appeared. A literature search elucidated the roles of the Notch system's components in modern descendants of the ortholog-contributing ancestors. A single protein, the protein tyrosine kinase (PTK) of the protozoan Ministeria vibrans, was identified as a possible pre-Metazoan ancestor of all three of the Notch pathway proteins, DLL, JAG, and NOTCH. A scenario for the evolution of the Notch signaling pathway is presented and described as the co-option of its components, clade by clade, in a repurposing of genes already present in ancestral unicellular organisms.

The evolution of flexibility and function in the Fc domains of IgM, IgY, and IgE.

Antibody Fc regions harbour the binding sites for receptors that mediate effector functions following antigen engagement by the Fab regions. An extended "hinge" region in IgG allows flexibility between Fab and Fc, but in both the most primitive antibody, IgM, and in the evolutionarily more recent IgE, the hinge is replaced by an additional domain pair in the homodimeric six-domain Fc region. This permits additional flexibility within the Fc region, which has been exploited by nature to modulate antibody effector functions. Thus, in pentameric or hexameric IgM, the Fc regions appear to adopt a planar conformation in solution until antigen binding causes a conformational change and exposes the complement binding sites. In contrast, IgE-Fc principally adopts an acutely bent conformation in solution, but the binding of different receptors is controlled by the degree of bending, and there is allosteric communication between receptor binding sites. We sought to trace the evolution of Fc conformational diversity from IgM to IgE via the intermediate avian IgY by studying the solution conformations of their Fc regions by small-angle X-ray scattering. We compared four extant proteins: human IgM-Fc homodimer, chicken IgY-Fc, platypus IgE-Fc, and human IgE-Fc. These are examples of proteins that first appeared in the jawed fish [425 million years ago (mya)], tetrapod (310 mya), monotreme (166 mya), and hominid (2.5 mya) clades, respectively. We analysed the scattering curves in terms of contributions from a pool of variously bent models chosen by a non-negative linear least-squares algorithm and found that the four proteins form a series in which the proportion of acutely bent material increases: IgM-Fc < IgY-Fc < plIgE-Fc < huIgE-Fc. This follows their order of appearance in evolution. For the huIgM-Fc homodimer, although none are acutely bent, and a significant fraction of the protein is sufficiently bent to expose the C1q-binding site, it predominantly adopts a fully extended conformation. In contrast, huIgE-Fc is found principally to be acutely bent, as expected from earlier studies. IgY-Fc, in this first structural analysis of the complete Fc region, exhibits an ensemble of conformations from acutely bent to fully extended, reflecting IgY's position as an evolutionary intermediate between IgM and IgE.

The evolution of tenascins

BackgroundThe evolution of extracellular matrix is tightly linked to the evolution of organogenesis in metazoans. Tenascins are extracellular matrix glycoproteins of chordates that participate in integrin-signaling and morphogenetic events. Single tenascins are encoded by invertebrate chordates, and multiple tenascin paralogs are found in vertebrates (designated tenascin-C, tenascin-R, tenascin-W and tenascin-X) yet, overall, the evolution of this family has remained unclear.ResultsThis study examines the genomes of hemichordates, cephalochordates, tunicates, agnathans, cartilaginous fishes, lobe-finned fishes, ray-finned fishes and representative tetrapods to identify predicted tenascin proteins. We comprehensively assess their evolutionary relationships by sequence conservation, molecular phylogeny and examination of conservation of synteny of the encoding genes. The resulting new evolutionary model posits the origin of tenascin in an ancestral chordate, with tenascin-C-like and tenascin-R-like paralogs emerging after a whole genome duplication event in an ancestral vertebrate. Tenascin-X appeared following a second round of whole genome duplication in an ancestral gnathostome, most likely from duplication of the gene encoding the tenascin-R homolog. The fourth gene, encoding tenascin-W (also known as tenascin-N), apparently arose from a local duplication of tenascin-R.ConclusionsThe diversity of tenascin paralogs observed in agnathans and gnathostomes has evolved through selective retention of novel genes that arose from a combination of whole genome and local duplication events. The evolutionary appearance of specific tenascin paralogs coincides with the appearance of vertebrate-specific cell and tissue types where the paralogs are abundantly expressed, such as the endocranium and facial skeleton (tenascin-C), an expanded central nervous system (tenascin-R), and bone (tenascin-W).

MRNA context and translation factors determine decoding in alternative nuclear genetic codes.

The genetic code is a set of instructions that determine how the information in our genetic material is translated into amino acids. In general, it is universal for all organisms, from viruses and bacteria to humans. However, in the last few decades, exceptions to this rule have been identified both in pro- and eukaryotes. In this review, we discuss the 16 described alternative eukaryotic nuclear genetic codes and observe theories of their appearance in evolution. We consider possible molecular mechanisms that allow codon reassignment. Most reassignments in nuclear genetic codes are observed for stop codons. Moreover, in several organisms, stop codons can simultaneously encode amino acids and serve as termination signals. In this case, the meaning of the codon is determined by the additional factors besides the triplets. A comprehensive review of various non-standard coding events in the nuclear genomes provides a new insight into the translation mechanism in eukaryotes.

Developmental process of the modern house shrew's molars: implications for the evolution of the tribosphenic molar in Mesozoic mammals.

Phylogenetically, the tribosphenic molars-prototypes of multi-cusped cheek teeth in marsupial and placental mammals-are derived from the single-cusped conical teeth of reptiles through the addition of cusps. Ontogenetically, mammalian molars are formed through the interface between the dental epithelium and mesenchyme (future enamel-dentin junction), becoming geometrically complex by adding epithelial signaling centers, called enamel knots, which determine future cusp positions. To reevaluate cusp homologies in Mesozoic mammals from an ontogenetic perspective, this study tracked molar development in a living placental mammal species, the house shrew (Suncus murinus), whose molars are morphologically the least derived from tribosphenic prototypes. The development of shrew molars proceeded as if it replayed the evolutionary process of tribosphenic molars. The first formed enamel knots gave rise to the evolutionarily oldest cusps-upper paracone and lower protoconid. The order of formation of other enamel knots and their location in development seemed to trace the order of cusp appearance in evolution. The parallel relationship between ontogeny and phylogeny of mammalian molars, if any, suggests that a change in the timing between developmental events rather than a change in the morphogenetic mechanism itself, should have been a major causal factor for the evolutionary transformation of tooth morphology.

Revision of the Quaternary calcareous nannofossil biochronology of Arctic Ocean sediments

Despite extensive chronological studies, the relationship between the age and sub-seafloor depth of Arctic Ocean sediments remains ambiguous. This prevents confident identification of paleoceanographic changes in the Arctic during the Quaternary. Currently, age-depth models derived from uranium-series decay in Arctic sediments diverge by hundreds of thousands of years compared to those built on known evolutionary appearances and extinctions of calcareous nannoplankton, a group of globally valuable age-markers. Here we report on high-resolution biostratigraphic analysis of late Quaternary sediments in six cores from the central Arctic Ocean (CAO). We applied paired light microscope (LM) and scanning electron microscope (SEM) imaging to improve nannofossil diagnosis. We argue that low abundances and poor preservation have led to misidentification of the true stratigraphic depth of the critical Pleistocene nannofossil bio-events that have underpinned age models for many Arctic sedimentary records for decades. The revised calcareous nannofossil biochronology provides a radically different geochronological framework for CAO sediments – indicating that what had previously been identified as Marine Isotope Stage (MIS) 7 (191–243 ka) in many sedimentary records is older than MIS 12 (424–478 ka). Furthermore, it suggests that previously inferred sub-stages of MIS 5 could represent full interglacial periods rather than interstadials. The results help reconcile the different dating approaches and provide a transformative step towards resolving the disparity in Quaternary Arctic age-depth models, bringing us one step closer to accurate paleoceanographic reconstructions based on sediment cores.

The fossil record of appendicular muscle evolution in Synapsida on the line to mammals: Part II-Hindlimb.

This paper is the second in a two-part series that charts the evolution of appendicular musculature along the mammalian stem lineage, drawing upon the exceptional fossil record of extinct synapsids. Here, attention is focused on muscles of the hindlimb. Although the hindlimb skeleton did not undergo as marked a transformation on the line to mammals as did the forelimb skeleton, the anatomy of extant tetrapods indicates that major changes to musculature have nonetheless occurred. To better understand these changes, this study surveyed the osteological evidence for muscular attachments in extinct mammalian and nonmammalian synapsids, two extinct amniote outgroups, and a large selection of extant mammals, saurians, and salamanders. Observations were integrated into an explicit phylogenetic framework, comprising 80 character-state complexes covering all muscles crossing the hip, knee, and ankle joints. These were coded for 33 operational taxonomic units spanning >330 Ma of tetrapod evolution, and ancestral state reconstruction was used to evaluate the sequence of muscular evolution along the stem lineage from Amniota to Theria. The evolutionary history of mammalian hindlimb musculature was complex, nonlinear, and protracted, with several instances of convergence and pulses of anatomical transformation that continued well into the crown group. Numerous traits typically regarded as characteristically "mammalian" have much greater antiquity than previously recognized, and for some traits, most synapsids are probably more reflective of the ancestral amniote condition than are extant saurians. More broadly, this study highlights the utility of the fossil record in interpreting the evolutionary appearance of distinctive anatomies.

Evolution of the jasmonate ligands and their biosynthetic pathways.

Different plant species employ different jasmonates to activate a conserved signalling pathway in land plants, where (+)-7-iso-JA-Ile (JA-Ile) is the ligand for the COI1/JAZ receptor in angiosperms and dn-cis-OPDA, dn-iso-OPDA and Δ4 -dn-iso-OPDA act as ligands in Marchantia polymorpha. In addition, some jasmonates play a COI1-independent role. To understand the distribution of bioactive jasmonates in the green lineage and how their biosynthetic pathways evolved, we performed phylogenetic analyses and systematic jasmonates profiling in representative species from different lineages. We found that both OPDA and dn-OPDA are ubiquitous in all tested land plants and present also in charophyte algae, underscoring their importance as ancestral signalling molecules. By contrast, JA-Ile biosynthesis emerged within lycophytes coincident with the evolutionary appearance of JAR1 function. We identified that the OPR3-independent JA biosynthesis pathway is ancient and predates the evolutionary appearance of the OPR3-dependent pathway. Moreover, we identified a negative correlation between dn-iso-OPDA and JA-Ile in land plants, which supports that in bryophytes and lycophytes dn-iso-OPDA represents the analogous hormone to JA-Ile in other vascular plants.

Planktonic foraminifera document palaeoceanographic changes across the middle Cenomanian carbon-isotope excursion MCE 1: new evidence from the UK chalk

AbstractPlanktonic foraminifera were studied at Lydden Spout, near Folkestone (southeast England, UK), the reference section of the middle Cenomanian Event 1 (MCE 1) characterized by a prominent double-peak δ13C excursion of 1 ‰ identified in different ocean basins and considered a global event. Biostratigraphic and quantitative analysis of planktonic foraminifera are correlated to the δ13C perturbation, to the positive δ18O shifts identified within MCE 1 and to the occurrence of Boreal macrofossils (the bivalvesChlamys arlesiensisandOxytoma seminudum, and the belemnitePraectinocamax primus). Variations in abundance and species richness of planktonic foraminifera and the inferred palaeoecological preferences of taxa permit the identification of distinct palaeoenvironmental settings across MCE 1. The stratigraphic interval corresponding to MCE 1 is characterized by the absence of oligotrophic rotaliporids, and by the evolutionary appearance of meso-eutrophic dicarinellids and ofMuricohedbergella portsdownensis, a cold-water species that occurs at the same level as the Boreal macrofossils. These observations indicate a palaeoceanographic scenario characterized by reduced stratification of surface waters and absence/disruption of the thermocline in a dominantly eutrophic regime during MCE 1. Evidence provided by planktonic foraminifera, Boreal macrofossils and δ18O records documented for the late Cenomanian Plenus Cold Event (PCE) at Eastbourne (UK) reveal similarities that confirm the periodic inflow of cold Boreal seawater originating in the Norwegian Sea as previously postulated to explain the occurrence of Boreal fauna in the Anglo-Paris Basin. The southerly extension of this water mass may be related to the reorganization of circulation driven by the long eccentricity cycle.

Defective Thyroglobulin: Cell Biology of Disease.

The primary functional units of the thyroid gland are follicles of various sizes comprised of a monolayer of epithelial cells (thyrocytes) surrounding an apical extracellular cavity known as the follicle lumen. In the normal thyroid gland, the follicle lumen is filled with secreted protein (referred to as colloid), comprised nearly exclusively of thyroglobulin with a half-life ranging from days to weeks. At the cellular boundary of the follicle lumen, secreted thyroglobulin becomes iodinated, resulting from the coordinated activities of enzymes localized to the thyrocyte apical plasma membrane. Thyroglobulin appearance in evolution is essentially synchronous with the appearance of the follicular architecture of the vertebrate thyroid gland. Thyroglobulin is the most highly expressed thyroid gene and represents the most abundantly expressed thyroid protein. Wildtype thyroglobulin protein is a large and complex glycoprotein that folds in the endoplasmic reticulum, leading to homodimerization and export via the classical secretory pathway to the follicle lumen. However, of the hundreds of human thyroglobulin genetic variants, most exhibit increased susceptibility to misfolding with defective export from the endoplasmic reticulum, triggering hypothyroidism as well as thyroidal endoplasmic reticulum stress. The human disease of hypothyroidism with defective thyroglobulin (either homozygous, or compound heterozygous) can be experimentally modeled in thyrocyte cell culture, or in whole animals, such as mice that are readily amenable to genetic manipulation. From a combination of approaches, it can be demonstrated that in the setting of thyroglobulin misfolding, thyrocytes under chronic continuous ER stress exhibit increased susceptibility to cell death, with interesting cell biological and pathophysiological consequences.

Trace classical conditioning impairment after lesion of the lateral part of the goldfish telencephalic pallium suggests a long ancestry of the episodic memory function of the vertebrate hippocampus.

There is an ongoing debate on the evolutionary origin of the episodic memory function of the hippocampus. A widely accepted hypothesis claims that the hippocampus first evolved as a dedicated system for spatial navigation in ancestral vertebrates, being transformed later in phylogeny to support a broader role in episodic memory with the emergence of mammals. On the contrary, an alternative hypothesis holds that the hippocampus of ancestral vertebrates originally encoded both the spatial and temporal dimensions of relational memories since its evolutionary appearance, thus suggesting that the episodic-like memory function of the hippocampus could be the primitive condition in vertebrate forebrain evolution. The present experiment was aimed at scrutinizing these opposing hypotheses by investigating whether the hippocampal pallium of teleost fish, a vertebrate group that shares with mammals a common ancestor that lived about 400 Mya, is, like the hippocampus of mammals, essential to associate time-discontiguous events. Thus, goldfish with lesions in the ventral part of the dorsolateral pallium (Dlv), a telencephalic region considered homologous to the hippocampal pallium of land vertebrates, were trained in trace versus delay eyeblink-like classical conditioning, two learning procedures that differ only in the temporal relationships between the stimuli to be associated in memory. The results showed that hippocampal pallium lesion in goldfish severely impairs trace conditioning, but spares delay conditioning. This finding challenges the idea that navigation preceded relational memory in evolutionary appearance and suggests the possibility that a relational memory function that associates the experienced events in both the spatial and temporal dimensions could be a primitive feature of the hippocampus that pre-existed in the common ancestor of vertebrates.

Energetic costs and benefits of sleep

Energy derived from food is a precious resource to animals. Those finite calories are often well-earned through exhaustive foraging effort, which can dominate waking hours, to support physiological processes (e.g. body maintenance and growth) and ecological necessities (e.g. predator avoidance and courting) that are pertinent to the production of progeny. So, it is unsurprising to find that animals have evolved strategies to guard against the gratuitous waste of hard-won caloric energy. Yet, it remains surprising to find such diversity, and elegant creativity, in those solutions. Brief examples of energy-saving innovation could include the very shape of animals and how they move, from streamlined swimming sharks to skyward-soaring seabirds; or the evolutionary appearance of various states of dormancy, such as endothermic animals sacrificing high body temperature through modest (torpor) or severe (hibernation) curtailments to metabolic heat production. Another reversibly dormant state with energetic benefits is sleep.

Vestibular Influence on Vertebrate Skeletal Symmetry and Body Shape.

Vestibular endorgans in the vertebrate inner ear form the principal sensors for head orientation and motion in space. Following the evolutionary appearance of these organs in pre-vertebrate ancestors, specific sensory epithelial patches, such as the utricle, which is sensitive to linear acceleration and orientation of the head with respect to earth’s gravity, have become particularly important for constant postural stabilization. This influence operates through descending neuronal populations with evolutionarily conserved hindbrain origins that directly and indirectly control spinal motoneurons of axial and limb muscles. During embryogenesis and early post-embryonic periods, bilateral otolith signals contribute to the formation of symmetric skeletal elements through a balanced activation of axial muscles. This role has been validated by removal of otolith signals on one side during a specific developmental period in Xenopus laevis tadpoles. This intervention causes severe scoliotic deformations that remain permanent and extend into adulthood. Accordingly, the functional influence of weight-bearing otoconia, likely on utricular hair cells and resultant afferent discharge, represents a mechanism to ensure a symmetric muscle tonus essential for establishing a normal body shape. Such an impact is presumably occurring within a critical period that is curtailed by the functional completion of central vestibulo-motor circuits and by the modifiability of skeletal elements before ossification of the bones. Thus, bilateral otolith organs and their associated sensitivity to head orientation and linear accelerations are not only indispensable for real time postural stabilization during motion in space but also serve as a guidance for the ontogenetic establishment of a symmetric body.

The evolution of anteriorly directed molar occlusion in mammals

AbstractIn non-mammalian synapsids and early mammals, evolutionary transformations in the feeding and hearing apparatuses are posited to have been prerequisites for the radiation of extant mammals. Unlike most vertebrates, including many early synapsids, mammals have precise dental occlusion, a lower jaw composed of one bone, and middle ear ossicles derived from ancestral jaw bones. We illuminate a related functional transition: therian mammals (eutherians and metatherians) evolved anteriorly directed chewing strokes, which are absent in other synapsid lineages. Anteriorly directed jaw movement during occlusion necessitates anteriorly directed muscle force vectors, and we posit that a shift in muscle orientation is reflected in the fossil record by the evolutionary appearance of a posteriorly positioned angular process in cladotherians (therians and their close kin). Anteriorly directed occlusion might have been absent in earlier synapsids because of the presence of attached middle ear elements in the posterior region of the jaw that prohibited the posterior insertion of jaw musculature. These changes to the masticatory apparatus in cladotherians are likely to have permitted the evolution of novel masticatory movements, including grinding in both the anterior and medial directions (e.g. rodents and ungulates, respectively). Thus, this evolutionary transition might have been a crucial prerequisite for the dietary diversification of therians.

Early Cretaceous radiolarians and chert geochemistry from western Yarlung Tsangpo suture zone in Jiangyema section, Purang county, SW Tibet

Moderately to well-preserved radiolarian faunas belonging to 51 genera and 81 species have been identified in a continuous bedded chert succession in the Jiangyema section in the west segment of the Yarlung Tsangpo suture zone (YTSZ), Purang county, SW Tibet. Based on the occurrence of some characteristic species, especially the evolutionary appearance of age-diagnostic species two radiolarian zones and two subzones are recognized and respectively named as the Cecrops septemporatus zone, Aurisaturnalis carinatus zone, Aurisaturnalis carinatus carinatus subzone and Aurisaturnalis carinatus perforatus subzone. These can be correlated with coeval Hauterivian to upper Barremian radiolarian biozones in both the western Tethys and Japan. They provide a better biostratigraphical framework and reliable age constraints for parts of the Neotethyan Ocean floor that were subducted beneath supra-subduction zone (SSZ) ophiolites in the western part of the YTSZ.Geochemical data indicate that SiO2 contents of bedded cherts range from 79.84 to 94.65% with an average of 89.39%. Comparison of geochemical characteristics on discriminant graphs suggests that the bedded cherts were deposited in a deep oceanic basin near a continental margin. Our results and integration of available radiolarian studies from bedded cherts along the YTSZ suggest that the Neo-Tethys was a deep ocean between the Indian and Eurasian continents in which pelagic sedimentation was on-going until at least the late Barremian.

Pragian conodont zonal classification in Nevada, western North America

A tripartite global zonal scale for the Pragian Stage (Devonian) was recommended by the Subcomission on Devonian Stratigraphy in 1989. Since that time, additions to the data on the two primary lineages used for the subdivision of the Pragian, Eognathodus and early Polygnathus, have shown: 1) that the Lochkovian-Pragian boundary criterion is useable as defi ned and is applicable on a global scale, but that the boundary-stratotype section in the Barrandian region of the Czech Republic has serious limitations as a reference section; 2) the criteria for the internal subdivision of the Pragian are not globally applicable; and 3) that the taxon that we have used in Nevada to mark the base of the Emsian is Polygnathus lenzi, whose range may be different from that of Australian P. dehiscens, the supposed criterion for the base of the Emsian. This paper reviews the Pragian of Nevada and compares it with Alaska, Canada, eastern Australia, and central Europe and suggests a regional scale for use in the western North American Cordillera. The subdivisions proposed here (irregularis-profunda Zone; profunda–brevicauda Zone; brevicauda-mariannae Zone; mariannae-lenzi Zone) are not the same of the SDS and cannot be applied on a global scale. The first two are based on evolutionary appearances; the last two on the lowest occurrences of distinctive and widespread taxa in North America. One new genus, Masaraella and four new species, Masaraella epsilon, Masaraella riosi, Gondwania profunda, and Pedavis longicauda, are described.

Tenascin-W: Discovery, Evolution, and Future Prospects.

Of the four tenascins found in bony fish and tetrapods, tenascin-W is the least understood. It was first discovered in the zebrafish and later in mouse, where it was mistakenly named tenascin-N. Tenascin-W is expressed primarily in developing and mature bone, in a subset of stem cell niches, and in the stroma of many solid tumors. Phylogenetic studies show that it is the most recent tenascin to evolve, appearing first in bony fishes. Its expression in bone and the timing of its evolutionary appearance should direct future studies to its role in bone formation, in stem cell niches, and in the treatment and detection of cancer.

An Ancient COI1-Independent Function for Reactive Electrophilic Oxylipins in Thermotolerance.

The jasmonate signaling pathway regulates development, growth, and defense responses in plants. Studies in the model eudicot, Arabidopsis thaliana, have identified the bioactive hormone (jasmonoyl-isoleucine [JA-Ile]) and its Coronatine Insensitive 1 (COI1)/Jasmonate-ZIM Domain (JAZ) co-receptor. In bryophytes, a conserved signaling pathway regulates similar responses but uses a different ligand, the JA-Ile precursor dinor-12-oxo-10,15(Z)-phytodienoic acid (dn-OPDA), to activate a conserved co-receptor. Jasmonate responses independent of JA-Ile and COI1, thought to be mediated by the cyclopentenone OPDA, have also been suggested, but experimental limitations in Arabidopsis have hindered attempts to uncouple OPDA and JA-Ile biosynthesis. Thus, a clear understanding of this pathway remains elusive. Here, we address the role of cyclopentenones in COI1-independent responses using the bryophyte Marchantia polymorpha, which is unable to synthesize JA-Ile but does accumulate OPDA and dn-OPDA. We demonstrate that OPDA and dn-OPDA activate a COI1-independent pathway that regulates plant thermotolerance genes, and consequently, treatment with these oxylipins protects plants against heat stress. Furthermore, we identify that these molecules signal through their electrophilic properties. By performing comparative analyses between M.polymorpha and two evolutionary distant species, A.thaliana and the charophyte alga Klebsormidium nitens, we demonstrate that this pathway is conserved in streptophyte plants and pre-dates the evolutionary appearance of the COI1-dependent jasmonate pathway, which later co-opted the pre-existing dn-OPDA as its ligand. Taken together, our data indicate that cyclopentenone-regulated COI1-independent signaling is an ancient conserved pathway, whose ancestral role was to protect plants against heat stress. This pathway was likely crucial for plants' successful land colonization and will be critical for adaption to current climate warming.