Outer Mantle Research Articles

Molecular characterization, light-dependent expression, and cellular localization of a host vacuolar-type H+-ATPase (VHA) subunit A in the giant clam, Tridacna squamosa, indicate the involvement of the host VHA in the uptake of inorganic carbon and its supply to the symbiotic zooxanthellae

The giant clam, Tridacna squamosa, represents a clam-zooxanthellae association. In light, the host clam and the symbiotic zooxanthellae conduct light-enhanced calcification and photosynthesis, respectively. We had cloned the cDNA coding sequence of a Vacuolar-type Proton ATPase (VHA) subunit A, ATP6V1A, from T. squamosa, whereby the VHA is an electrogenic transporter that actively ‘pumps’ H+ out of the cell. The ATP6V1A of T. squamosa comprised 1866 bp, encoding a protein of 622 amino acids and 69.9 kDa, and had a host-origin. Its gene expression was strong in the ctenidium and the colorful outer mantle, but weak in the whitish inner mantle, corroborating a previous proposition that VHA might have a trivial role in light-enhanced calcification. Light exposure led to significant increases in the gene and protein expression levels of ATP6V1A/ATP6V1A in the ctenidium and the outer mantle. In the ctenidium, the ATP6V1A was localized in the apical epithelia of the filaments and tertiary water channels, indicating that the VHA could participate in the increased excretion of H+ produced during light-enhanced calcification. Additionally, the excreted H+ would augment HCO3– dehydration in the external medium and facilitate the uptake of CO2 by the ctenidium during insolation. In the outer mantle, the ATP6V1A was detected in intracellular vesicles in a type of cells, presumably iridocytes, surrounding the zooxanthellal tubules, and in the apical epithelium of zooxanthellal tubules. Hence, the host VHA could participate in the transfer of inorganic carbon from the hemolymph to the luminal fluid of the tubules by increasing the supply of H+ for the dehydration of HCO3– to CO2 during insolation to benefit the photosynthesizing zooxanthellae.

Walking the thin line… ten years later: the dilemma of above- versus below-ground features to support phylogenies in the Russulaceae (Basidiomycota)

For the very first time, morpho-anatomical features of both fruiting bodies as well as below-ground structures have been confronted with a newly produced multigene phylogeny of root symbiotic basidiomycetes using one of the most speciose genera of ectomycorrhizal fungi (Russula, Russulales) as an example. In this first of two papers, the authors focus more specifically on below-ground structures. Our five-gene phylogeny divides the genus in five main clades, here interpreted as representing seven subgenera, all significantly supported. Although more conserved than features of fruiting bodies, the anatomy of ectomycorrhiza does not allow for an unambiguous characterization of the main clades resolved by phylogenetic analysis, but the anatomy of ectomycorrhiza performs better to naturally classify the species of this genus. Features of fruiting bodies remain much more adequate for the delimitation of terminal clades and are irreplaceable for morphological species identification. Tropical taxa mostly nest in ancient lineages, but are also present in some terminal clades of otherwise temperate species groups. The shift from plectenchymatic to pseudoparenchymatic ECM outer mantle structures happened most likely already in the paleotropics, and is here hypothesized to have facilitated a major diversification of the genus with new hosts in the northern hemisphere. Available data as well as our own observations on below ground structures of several Lactifluus species suggests that this genus shares with Russula the absence of lactifers in ECM mantles and rhizomorphs, contrary to species of Lactarius where lactifers are always present. First observations on rhizomorphs of species in Multifurca confirm the presence of vessel-like and ladder-like hyphae, also found in the other agarioid genera of this family, while distinct lactifers are only present in the lactarioid, but not in russuloid members of this genus.

Molecular characterization of a novel algal glutamine synthetase (GS) and an algal glutamate synthase (GOGAT) from the colorful outer mantle of the giant clam, Tridacna squamosa, and the putative GS-GOGAT cycle in its symbiotic zooxanthellae

Giant clams harbor symbiotic zooxanthellae (Symbiodinium), which are nitrogen-deficient, mainly in the fleshy and colorful outer mantle. This study aimed to sequence and characterize the algal Glutamine Synthetase (GS) and Glutamate Synthase (GLT), which constitute the glutamate synthase cycle (or GS-GOGAT cycle, whereby GOGAT is the protein acronym of GLT) of nitrogen assimilation, from the outer mantle of the fluted giant clam, Tridacna squamosa. We had identified a novel GS-like cDNA coding sequence of 2325 bp, and named it as T. squamosa Symbiodinium GS1 (TSSGS1). The deduced TSSGS1 sequence had 774 amino acids with a molecular mass of 85 kDa, and displayed the characteristics of GS1 and Nucleotide Diphosphate Kinase. The cDNA coding sequence of the algal GLT, named as T. squamosa Symbiodinium GLT (TSSGLT), comprised 6399 bp, encoding a protein of 2133 amino acids and 232.4 kDa. The zooxanthellal origin of TSSGS1 and TSSGOGAT was confirmed by sequence comparison and phylogenetic analyses. Indeed, TSSGS1 and TSSGOGAT were expressed predominately in the outer mantle, which contained the majority of the zooxanthellae. Immunofluorescence microscopy confirmed the expression of TSSGS1 and TSSGOGAT in the cytoplasm and the plastids, respectively, of the zooxanthellae in the outer mantle. It can be concluded that the symbiotic zooxanthellae of T. squamosa possesses a glutamate synthase (TSSGS1-TSSGOGAT) cycle that can assimilate endogenous ammonia produced by the host clam into glutamate, which can act as a substrate for amino acid syntheses. Thus, our results provide insights into why intact giant clam-zooxanthellae associations do not excrete ammonia under normal circumstances.

Light exposure enhances urea absorption in the fluted giant clam, Tridacna squamosa, and up-regulates the protein abundance of a light-dependent urea active transporter, DUR3-like, in its ctenidium.

Giant clams live in nutrient-poor reef waters of the Indo-Pacific and rely on symbiotic dinoflagellates (Symbiodinium spp., also known as zooxanthellae) for nutrients. As the symbionts are nitrogen deficient, the host clam has to absorb exogenous nitrogen and supply it to them. This study aimed to demonstrate light-enhanced urea absorption in the fluted giant clam, Tridacna squamosa, and to clone and characterize the urea active transporter DUR3-like from its ctenidium (gill). The results indicate that T. squamosa absorbs exogenous urea, and the rate of urea uptake in the light was significantly higher than that in darkness. The DUR3-like coding sequence obtained from its ctenidium comprised 2346 bp, encoding a protein of 782 amino acids and 87.0 kDa. DUR3-like was expressed strongly in the ctenidium, outer mantle and kidney. Twelve hours of exposure to light had no significant effect on the transcript level of ctenidial DUR3-like However, between 3 and 12 h of light exposure, DUR3-like protein abundance increased progressively in the ctenidium, and became significantly greater than that in the control at 12 h. DUR3-like had an apical localization in the epithelia of the ctenidial filaments and tertiary water channels. Taken together, these results indicate that DUR3-like might participate in light-enhanced urea absorption in the ctenidium of T. squamosa When made available to the symbiotic zooxanthellae that are known to possess urease, the absorbed urea can be metabolized to NH3 and CO2 to support amino acid synthesis and photosynthesis, respectively, during insolation.

Do squid breathe through their skin?

Squid are thought to obtain a large portion of their oxygen via simple diffusion across the skin in addition to uptake at the gills. Although this hypothesis has support from indirect evidence and is widely accepted, no empirical examinations have been conducted to assess the validity of this hypothesis. In this study, we examined cutaneous respiration in two squid species, Doryteuthis pealeii and Lolliguncula brevis, using a divided chamber to physically separate the mantle cavity and gills from the outer mantle surface. We measured oxygen consumption and ammonia excretion rates in the two compartments and found that, at rest, squid only obtain enough oxygen cutaneously to meet the demand of the skin tissue locally (12% of total) and excrete little ammonia across the skin. The majority of oxygen is obtained via the traditional branchial pathway. In light of these findings, we re-examine and discuss the indirect evidence that has supported the cutaneous respiration hypothesis.

STAGES OF DEVELOPMENT OF THE PLANET EARTH IN THE FRAMEWORK OF ANALYTICAL GEOLOGY

Based on the known geophysical data on the structure of the inner and outer core, mantle, oceanic and continental crust, within the framework of the concept of self-development and self-organization of protoplanetary matter underthe influence of the cosmic pressure, against the background of a pulsating reduction in the earth's radius, a possible mechanism for the formation of the planet's geospheres and its surface shell. Attention is drawn to the morphology of geo-textures and morphostructures of the ocean floor - as an information system that gives an idea of the course of deep processes. The prism blocks of increased stiffness (BCG) of the primary oceanic crust are located in deep-water basins, marginal and inland seas of the continents and their lowered areas. The assumption is that the BPZ ensembles, moving in expansion-contraction regimes under the influence of radial and rotational dynamics forces, formed sigmoid structures of the planet's surface and formed Procrustean beds for the development of icebergs of continents. The symmetry and antipodality of the surface structures of the Earth is explained by the superposition of two energy sources - external and internal: the internal energy substance, according to the rule of the borer, is more dynamic in the southern hemisphere, and the waves of the external energy of the universe are rolling on the Pacific Ocean.Key words: globule of the right-hand plane, high-energy substance, cosmic pressure, the wave field of the universe, sources-ooids, rheons, pulsation, expansion-contraction regimes.

Rapid and comprehensive discovery of unreported shellfish allergens using large-scale transcriptomic and proteomic resources

[Extract] Discovery of allergens in various food and inhalant sources is central to our understanding of the molecular mechanisms of allergic reactions. Allergen characterization is the most important underlying factor for better patient management and the design and development of novel immunotherapeutics.1 Of the "Big eight" allergen food groups, shellfish presents a unique challenge in terms of allergen discovery due to the large number and diversity of consumed species, leading to heterogeneity of allergen structure and cross-reactivity among various sources. At present, 31 crustacean allergens have been officially registered in the World Health Organization and International Union of Immunological Societies Allergen Nomenclature Database as compared with 4 mollusk allergens due to pitfalls in current allergy discovery approaches. Cosensitization of patients with crustacean and mollusk allergy is often described; however, the current diagnostic approaches to manage these patients are not based on sufficient molecular knowledge of these shellfish allergens.

Carbonic anhydrase 2-like in the giant clam, Tridacna squamosa: characterization, localization, response to light, and possible role in the transport of inorganic carbon from the host to its symbionts.

The fluted giant clam, Tridacna squamosa, lives in symbiosis with zooxanthellae which reside extracellularly inside a tubular system. Zooxanthellae fix inorganic carbon (Ci) during insolation and donate photosynthate to the host. Carbonic anhydrases catalyze the interconversion of CO 2 and HCO3−, of which carbonic anhydrase 2 (CA2) is the most ubiquitous and involved in many biological processes. This study aimed to clone a CA2 homolog (CA2‐like) from the fleshy and colorful outer mantle as well as the thin and whitish inner mantle of T. squamosa, to determine its cellular and subcellular localization, and to examine the effects of light exposure on its gene and protein expression levels. The cDNA coding sequence of CA2‐like from T. squamosa comprised 789 bp, encoding 263 amino acids with an estimated molecular mass of 29.6 kDa. A phenogramic analysis of the deduced CA2‐like sequence denoted an animal origin. CA2‐like was not detectable in the shell‐facing epithelium of the inner mantle adjacent to the extrapallial fluid. Hence, CA2‐like is unlikely to participate directly in light‐enhanced calcification. By contrast, the outer mantle, which contains the highest density of tertiary tubules and zooxanthellae, displayed high level of CA2‐like expression, and CA2‐like was localized to the tubule epithelial cells. More importantly, exposure to light induced significant increases in the protein abundance of CA2‐like in the outer mantle. Hence, CA2‐like could probably take part in the increased supply of inorganic carbon (Ci) from the host clam to the symbiotic zooxanthellae when the latter conduct photosynthesis to fix Ci during light exposure.

The thermal evolution of Mercury's Fe–Si core

We have studied the thermal and magnetic field evolution of planet Mercury with a core of Fe–Si alloy to assess whether an Fe–Si core matches its present-day partially molten state, Mercury's magnetic field strength, and the observed ancient crustal magnetization. The main advantages of an Fe–Si core, opposed to a previously assumed Fe–S core, are that a Si-bearing core is consistent with the highly reduced nature of Mercury and that no compositional convection is generated upon core solidification, in agreement with magnetic field indications of a stable layer at the top of Mercury's core. This study also present the first implementation of a conductive temperature profile in the core where heat fluxes are sub-adiabatic in a global thermal evolution model.We show that heat migrates from the deep core to the outer part of the core as soon as heat fluxes at the outer core become sub-adiabatic. As a result, the deep core cools throughout Mercury's evolution independent of the temperature evolution at the core-mantle boundary, causing an early start of inner core solidification and magnetic field generation. The conductive layer at the outer core suppresses the rate of core growth after temperature differences between the deep and shallow core are relaxed, such that a magnetic field can be generated until the present. Also, the outer core and mantle operate at higher temperatures than previously thought, which prolongs mantle melting and mantle convection. The results indicate that S is not a necessary ingredient of Mercury's core, bringing bulk compositional models of Mercury more in line with reduced meteorite analogues.

The Whitish Inner Mantle of the Giant Clam, Tridacna squamosa, Expresses an Apical Plasma Membrane Ca2+-ATPase (PMCA) Which Displays Light-Dependent Gene and Protein Expressions.

Giant clams live in symbiosis with extracellular zooxanthellae and display high rates of growth and shell formation (calcification) in light. Light-enhanced calcification requires an increase in the supply of Ca2+ to, and simultaneously an augmented removal of H+ from, the extrapallial fluid where shell formation occurs. We have obtained the complete coding cDNA sequence of Plasma Membrane Ca2+-ATPase (PMCA) from the thin and whitish inner mantle, which is in touch with the extrapallial fluid, of the giant clam Tridacna squamosa. The deduced PMCA sequence consisted of an apical targeting element. Immunofluorescence microscopy confirmed that PMCA had an apical localization in the shell-facing epithelium of the inner mantle, whereby it can actively secrete Ca2+ in exchange for H+. More importantly, the apical PMCA-immunofluorescence of the shell-facing epithelium of the inner mantle increased significantly after 12 h of exposure to light. The transcript and protein levels of PMCA/PMCA also increased significantly in the inner mantle after 6 or 12 h of light exposure. These results offer insights into a light-dependable mechanism of shell formation in T. squamosa and a novel explanation of light-enhanced calcification in general. As the inner mantle normally lacks light sensitive pigments, our results support a previous proposition that symbiotic zooxanthellae, particularly those in the colorful and extensible outer mantle, may act as light-sensing elements for the host clam.

An analysis of the geodesy and relativity experiments of BepiColombo

BepiColombo is a ESA-JAXA mission aimed to a comprehensive exploration of Mercury, the innermost planet of the solar system. The Mercury Orbiter Radio science Experiment (MORE) will exploit a state of the art microwave tracking system, including an advanced Ka-band transponder, to determine the gravity field and the rotational state of the planet, and to perform extensive tests of relativistic gravity. In this work we analyze all the aspects of the radio science investigation, which include: (i) the solar conjunction experiment in cruise; (ii) the gravimetry and rotation experiments; (iii) the fundamental physics test. We report on the results of numerical simulations based on the latest mission scenario, with launch in October 2018 and arrival at Mercury in December 2025. We show that the gravity and rotation measurements expected from BepiColombo will allow to better characterize the size of an inner solid core inside the outer liquid core, and the properties of the outer mantle and the crust. We discuss how the current estimate of several parametrized post-Newtonian (PPN) parameters can be improved by MORE through the determination of the heliocentric motion of Mercury and by measuring the propagation time of radio waves. We also assess in a quantitative way the benefits of an extended mission.

A review of applications of fractional calculus in Earth system dynamics

Fractional calculus has been used to model various hydrologic processes for 15years. Yet, there are still major gaps between real-world hydrologic dynamics and fractional-order partial differential equations (fPDEs). In addition, the applicability of fPDEs in the broad field of Earth dynamics remains obscure. This study first reviews previous applications and then identifies new research directions for fPDEs simulating non-Fickian transport in both surface and subsurface hydrology. We then explore the applicability of fractional calculus in various anomalous dynamics with a wide range of spatiotemporal scales observed in the solid Earth, including internal dynamics (such as inner core rotation, outer core flow, mantle convection, and crustal deformation), large-scale surface dynamics (in fluvial, Aeolian, and glacial systems), and small vertical-scale surface kinetics (in crystal growth, rock/mineral weathering, and pedogenesis), where driven forces, previous modeling approaches, and the details of anomalous dynamics are analyzed. Results show that the solid Earth can provide an ideal and diverse base for the application of fractional calculus and fPDEs. Complex dynamics within and across spatiotemporal scales, multi-scale intrinsic heterogeneity, and intertwined controlling factors for dynamic processes in the solid Earth can motivate the application of fPDEs. Challenges for the future application of fPDEs in Earth systems are also discussed, including poor parameter predictability, the lack of mathematical specification of bounded fractional diffusion, lack of intermediate-scale geologic information in parsimonious and upscaling models, and a lack of models for multi-phase and coupled processes. Substantial extension of fPDE models is needed for the development of next-generation, solid Earth dynamic models, where potential solutions are discussed based on our experience gained in the development and application of fractional calculus and fPDEs over the last decade. Therefore, the current bottleneck in the application of fractional calculus in hydrologic sciences should not be the end of a promising stochastic approach, but could be the early stage of a decade-long effort filled with multiple new research and application directions in geology. This conclusion may shed light on the bottleneck challenging stochastic hydrogeology, where the advanced stochastic models (with more than 3500 journal publications in the last three decades) have not significantly impacted the practice of groundwater flow and transport modeling.

Cuticle of Polyplacophora: structure, secretion, and homology with the periostracum of conchiferans

The plesiomorphic state of the molluscan scleritome remains ambiguous. Chitons are significant because they show a mix of characters considered diagnostic in both aplacophorans and conchiferans, with both shell plates and small calcified girdle elements on an elongate body, and there is no consensus on the homology of structures involved in chiton plate calcification with those in conchiferan shells. Using light, scanning, and transmission microscopy, the present study examined the structure and formation of the cuticle and underlying epithelium in five chiton species collected over several years in southeast Spain and in California, USA. The cuticular matrix in chitons is similar to the translucent layer of the bivalve periostracum, although there are differences in the orientation of the densely spaced laminae with respect to the mantle surface. The cuticle arises in the accessory fold of the ventral girdle mantle. There is full continuity between the outermost layer of the cuticle and the thin outer layer of the tegmentum, suggesting that the latter is derived from the cuticular wedge permanently lying on the plate margins. A homology is proposed between the cuticle/associated girdle mantle epithelium of Polyplacophora and the periostracum/inner side of the outer mantle fold of the Bivalvia. Nevertheless, while the bivalve periostracum slides over the mantle epithelium during growth, the polyplacophoran cuticle remains stationary. The configuration of the cuticle, which overlaps the plate margins, allows it to efficiently close the plate biomineralization compartment, in a way comparable to that of the periostracum of conchiferans.

Biomineralization-related specialization of hemocytes and mantle tissues of the Pacific oyster Crassostrea gigas.

The molluscan exoskeleton (shell) plays multiple important roles including structural support, protection from predators and stressors, and physiological homeostasis. Shell formation is a tightly regulated biological process that allows molluscs to build their shells even in environments unfavorable for mineral precipitation. Outer mantle edge epithelial cells (OME) and hemocytes were implicated in this process; however, the exact functions of these cell types in biomineralization are not clear. Pacific oysters (Crassostrea gigas) were used to study differences in the expression profiles of selected biomineralization-related genes in hemocytes and mantle cells, and the functional characteristics of hemocytes such as adhesion, motility and phagocytosis. The specialized role of OME in shell formation was supported by high expression levels of the extracellular matrix (ECM) related and cell-cell interaction genes. Density gradient separation of hemocytes revealed distinct phenotypes based on the cell morphology, gene expression patterns, motility and adhesion characteristics. These hemocyte fractions can be categorized into two functional groups, i.e. biomineralization and immune response cells. Gene expression profiles of the putative biomineralizing hemocytes indicate that in addition to their proposed role in mineral transport, hemocytes also contribute to the formation of the ECM, thus challenging the current paradigm of the mantle as the sole source of the ECM for shell formation. Our findings corroborate the specialized roles of hemocytes and the OME in biomineralization and emphasize complexity of the biological controls over shell formation in bivalves.

Intra-population variability of ocean acidification impacts on the physiology of Baltic blue mussels (Mytilus edulis): integrating tissue and organism response.

Increased maintenance costs at cellular, and consequently organism level, are thought to be involved in shaping the sensitivity of marine calcifiers to ocean acidification (OA). Yet, knowledge of the capacity of marine calcifiers to undergo metabolic adaptation is sparse. In Kiel Fjord, blue mussels thrive despite periodically high seawater PCO2, making this population interesting for studying metabolic adaptation under OA. Consequently, we conducted a multi-generation experiment and compared physiological responses of F1 mussels from 'tolerant' and 'sensitive' families exposed to OA for 1year. Family classifications were based on larval survival; tolerant families settled at all PCO2 levels (700, 1120, 2400µatm) while sensitive families did not settle at the highest PCO2 (≥99.8% mortality). We found similar filtration rates between family types at the control and intermediate PCO2 level. However, at 2400µatm, filtration and metabolic scope of gill tissue decreased in tolerant families, indicating functional limitations at the tissue level. Routine metabolic rates (RMR) and summed tissue respiration (gill and outer mantle tissue) of tolerant families were increased at intermediate PCO2, indicating elevated cellular homeostatic costs in various tissues. By contrast, OA did not affect tissue and routine metabolism of sensitive families. However, tolerant mussels were characterised by lower RMR at control PCO2 than sensitive families, which had variable RMR. This might provide the energetic scope to cover increased energetic demands under OA, highlighting the importance of analysing intra-population variability. The mechanisms shaping such difference in RMR and scope, and thus species' adaptation potential, remain to be identified.

Scleroderma areolatum ectomycorrhiza on Fagus sylvatica L.

Despite its broad host range and distribution and its potential applications in commercial plantation forests, comprehensive descriptions of Scleroderma ectomycorrhizae are available only for Scleroderma citrinum, Scleroderma bovista and Scleroderma sinnamariense. This study provides a morphological and anatomical description of tree nursery derived ectomycorrhizae of Scleroderma areolatum on Fagus sylvatica, grown for several years in a climatized room. Ectomycorrhizae of S. areolatum were silvery white with abundant rhizomorphs; all mantle layers were plectenchymatous, rhizomorphs of type E, with prominent emanating hyphae with thick cell wall. The distal ends of emanating hyphae of rhizomorphs were inflated and often merged with other emanating hyphae. All parts of the mycorrhiza were clampless. In hyphae of the outer mantle layer, rhizomorphs and emanating hyphae, oily droplets were observed that did not stain in sulfo-vanillin and disappeared in lactic acid after a few hours. Although the phylogenetic analysis positioned the newly described ectomycorrhiza together with Scleroderma verrucosum and Scleroderma cepa in a single clade with a taxon name SH005470.07FU, the ectomycorrhizae of these three species can be morphologically well separated based on rhizomorph type.

Chemical layering in the upper mantle of Mars: Evidence from olivine‐hosted melt inclusions in Tissint

AbstractMelting of Martian mantle, formation, and evolution of primary magma from the depleted mantle were previously modeled from experimental petrology and geochemical studies of Martian meteorites. Based on in situ major and trace element study of a range of olivine‐hosted melt inclusions in various stages of crystallization of Tissint, a depleted olivine–phyric shergottite, we further constrain different stages of depletion and enrichment in the depleted mantle source of the shergottite suite. Two types of melt inclusions were petrographically recognized. Type I melt inclusions occur in the megacrystic olivine core (Fo76‐70), while type II melt inclusions are hosted by the outer mantle of the olivine (Fo66‐55). REE‐plot indicates type I melt inclusions, which are unique because they represent the most depleted trace element data from the parent magmas of all the depleted shergottites, are an order of magnitude depleted compared to the type II melt inclusions. The absolute REE content of type II displays parallel trend but somewhat lower value than the Tissint whole‐rock. Model calculations indicate two‐stage mantle melting events followed by enrichment through mixing with a hypothetical residual melt from solidifying magma ocean. This resulted in ~10 times enrichment of incompatible trace elements from parent magma stage to the remaining melt after 45% crystallization, simulating the whole‐rock of Tissint. We rule out any assimilation due to crustal recycling into the upper mantle, as proposed by a recent study. Rather, we propose the presence of Al, Ca, Na, P, and REE‐rich layer at the shallower upper mantle above the depleted mantle source region during the geologic evolution of Mars.

A light-dependent ammonia-assimilating mechanism in the ctenidia of a giant clam

Giant clams harbor symbiotic zooxanthellae and manifest light-enhanced calcification. In light, they also increase the absorption and assimilation of exogenous ammonia, but the roles of the host and symbionts are unclear and the reason for light dependency remains enigmatic. Here, we report the complete coding cDNA sequence of a glutamine synthetase (GS), from the ctenidia (gill) of Tridacna squamosa. The deduced GS amino acid sequence was closely related to other animal GSs, indicating a host origin. The GS/GS transcript level and protein abundance increased significantly in the ctenidia after 12 h of light exposure. These results denote the ctenidia as the site of light-enhanced ammonia absorption with the absorbed ammonia being assimilated to glutamine by the host GS. Immunofluorescence microscopy confirmed that GS was expressed predominantly in the cytoplasm of the epithelial cells separating the tertiary water channels from the hemolymph inside the ctenidia. Hence, unlike symbiotic cnidarians, the host clam probably supplies exogenous nitrogen as glutamine from the ctenidia, through the hemolymph and tubular fluid, to the nitrogen-deficient zooxanthellae which reside extracellularly in the extensible outer mantle. This is the first report on light-dependent gene and protein expression of a host enzyme involved in nitrogen metabolism in an alga–invertebrate association.

GRAIL, LLR, and LOLA constraints on the interior structure of the Moon

AbstractThe interior structure of the Moon is constrained by its mass, moment of inertia, and k2 and h2 tidal Love numbers. We infer the likely radius, density, and (elastic limit) rigidity of all interior layers by solving the inverse problem using these observational constraints assuming spherical symmetry. Our results do not favor the presence of a low rigidity transition layer between a liquid outer core and mantle. If a transition layer exists, its rigidity is constrained to GPa, with a preference for the high rigidity values. Therefore, if a transition layer exists, it is more likely to have a rigidity similar to that of the mantle (∼70 GPa). The total (solid and liquid) core mass fraction relative to the lunar mass is constrained to and for interior structures with and without a transition layer, respectively, narrowing the range of possible giant impact formation scenarios.

Finite‐frequency wave propagation through outer rise fault zones and seismic measurements of upper mantle hydration

AbstractEffects of serpentine‐filled fault zones on seismic wave propagation in the upper mantle at the outer rise of subduction zones are evaluated using acoustic wave propagation models. Modeled wave speeds depend on azimuth, with slowest speeds in the fault‐normal direction. Propagation is fastest along faults, but, for fault widths on the order of the seismic wavelength, apparent wave speeds in this direction depend on frequency. For the 5–12 Hz Pn arrivals used in tomographic studies, joint‐parallel wavefronts are slowed by joints. This delay can account for the slowing seen in tomographic images of the outer rise upper mantle. At the Middle America Trench, confining serpentine to fault zones, as opposed to a uniform distribution, reduces estimates of bulk upper mantle hydration from ~3.5 wt % to as low as 0.33 wt % H2O.