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International audience; Cultivated pearls frequently exhibit morphological irregularities making obvious that mineral deposition was irregularly distributed onto nucleus surface. Taking advantage of experimental cultivations with short durations (from 10 days to few months), these irregular deposits predating occurrence of the nacre were investigated in Polynesian pearls by biochemical characterizations and a series of physical methods. Diversity in the resulting data suggests that various in‐depth alterations of the biomineralization mechanism may have occurred during the grafting process, leading to diversity in the biochemical pathways to nacreous deposition. This allows a precise discussion of current views about pearl formation. The “reversed shell theory” is formally disproved through point to point comparison with development of the shell growing edge. Similarity of pearl formation with “regeneration” or “shell repair” is also discussed, emphasizing the differences between these concepts.

International audience; The quality of natural and cultured pearls depends on the size, shape and properties of the surface. The inner structure of the pearl is not known, and not taken into account. Indeed, it is assumed that the cultured pearl is a nucleus covered by a thin organic layer, a thin prismatic layer and a thick nacreous layer. The examination of spherical and pear shape samples resulting from the grafting of the black lip pearl oyster (Pinctada margaritifera) shows that this assumption is, in most cases, erroneous. Optical, electron and atomic force microscopic observations, infrared and Raman analyses, as well as micro X-ray fluorescence and micro X-ray absorption near-edge spectroscopy data show that the inner structure of a pearl does not depend on its shape. Both circle or pear shape and spherical pearls show a large variability of the inner structures and composition. There are no two identical samples. Moreover, the arrangement is irregular within a single pearl, and some structures (aragonitic pseudo-prisms) do not exist in the shell.

In a series of Polynesian pearls collected after short cultivation periods, early post-grafting mineral deposits were characterized by high resolution synchrotron-based X-ray fluorescence with unprecedented accuracy. Morphological patterns and elemental composition are correlated through simultaneous imaging processes. Evidence that aragonite and calcite occur in neighboring units during the earliest biomineralization stages reveals that the grafting process can result in a greater degradation than usually admitted in the widely shared ‘reversed shell’ concept. Compared with ultrastructure of the pristine nacreous tablets, this method enables a precise evaluation of the possible biological changes in the biomineralization mechanism during grafting.

The pearl oyster Pinctada margaritifera has the specific ability to produce pearls with the widest range of colours among all pearl oyster species. This pearl colour diversity originates from the mantle biomineralising tissue (graft) of the donor oyster, which is originally responsible for the variety of colours of the inner shell surface. This study aimed to: 1) assess the geographic distribution and establish a first stocklist of the colourful oyster phenotypes used as donors in French Polynesia, and 2) investigate the phenotypic relation between inner shell colouration and the corresponding colour of harvested pearls. With the support of a pearl farmers' network, we investigated the different donor phenotype frequencies among five collection sites (Ahe, Apataki, Takaroa, TakumeandMangareva). This donor evaluation was made during grafting of pearl oysters (N=49,938) obtained from collector stations. Results showed that pearl production is mainly based on six common colourful donor phenotypes classified as monochromatic and polychromatic profiles, which shown different frequencies among the collection sites. Experimental grafts (N=4640)were then realised and subsequent culture conducted at a single site in order to avoid pearl colour variation due to environmental influences. Traceability between donors (N=232) and pearls (N=2776), revealed that: 1) yellow (gold) and aubergine (reddish) pearls could be mostly obtained by using the monochromatic yellowand red donor phenotypes, respectively, and 2) one third to one quarter of grey pearlswas inevitably harvested, whatever the polychromatic phenotype chosen as the donor, which leaves at least half the harvest composed of the attractive green and peacock colours. This preliminary stocklist of colour range together with analysis of the colour phenotype transmission between inner shell and pearl provide the basis for producing multiple pearl oyster “colour lines” through hatchery propagation and would be helpful for future selective breeding programs. Statement of relevance: Donor shell colour selection predict colours of pearls

The French Polynesian islands are internationally known for their black pearls, produced by culture of the black lipped pearl oyster Pinctada margaritifera. The ongoing development of hatcheries for P. margaritifera in French Polynesia poses new challenges for the industry, particularly regarding the maintenance of genetic diversity in the hatchery stocks. This emphasizes the necessity to characterize the genetic diversity and differentiation within natural and exploited populations, to carefully select putative parental populations. The present study aimed at validating the phylogenetic status and investigating genetic attributes of populations from the only two non-exploited archipelagos of French Polynesia, the Marquesas archipelago, and the Australes archipelago, never analysed before. We found that individuals from both archipelagos belonged to P. margaritifera species. However, while the Australes population was genetically similar to non-exploited populations of the Tuamotu, the Marquesas populations were highly differentiated from the rest of the populations. This differentiation cannot not be only attributed to geographic distance and aquaculture status, but likely to hydrodynamic barriers allowing vicariant events to take place. Our results add up to other studies describing the Marquesas archipelago as a hotspot for biodiversity and differentiation, with some of the highest levels of endemism and vicariance found among marine species worldwide and provide precious information on available genetic resources for the implementation of P. margaritifera selective breeding and its genetic conservation in French Polynesia.

In French Polynesia, the P. margaritifera pearl aquaculture industry is spread over a vast area, as large as Europe. All the oysters for this the highly economically important activity are supplied from just a few collection lagoons, but they are grown in numerous sites across three archipelagos (Gambier, Society and Tuamotu). Many oyster transfers thus indirectly bring about grafting combinations mixing different geographic origins and production sites. This study aims to examine the impact of such graft combinations on cultured pearl quality traits. For this, six homogeneous and standardised experimental graft combinations (N = 6197) were conducted at commercial scale in the two growing locations the most frequently used in French Polynesia: Arutua atoll (Tuamotu) and Mangareva island (Gambier), using oysters supplied from by the top three collection sites: Ahe, Takapoto and Mangareva lagoons. At harvest, four main pearl quality traits: nacre weight deposition speed, pearl colour components (darkness level and green overtone), grade and shape categories were recorded by a professional sorter from the Tahiti auction and compared. Results revealed effects of the combinations of oyster origin and grow-out location, with: 1) significant origin × site interaction for nacre weight deposition speed; 2) colour variation at intra- and inter-site scales, with Ahe origin producing the most dark pearls and Gambier highest rate of the attractive green coloured pearls; and 3) higher grade categories for the Gambier origin and rearing location. These oyster-site combination effects highlight the benefit for the Polynesian pearl industry of switching from a mono-site/ company production system to a new multi-site production strategy to maximize overall cultured pearl quality expression. Keywords: Pearl oyster, Pinctada margaritifera, Graft combinations, Pearl quality traits, Environment

Pinctada margaritifera is an economically important marine bivalve species for cultured pearl production in French Polynesian aquaculture. In order to evaluate the influence of donor oyster age on pearl quality traits, experiments were conducted over 6 years using both grafts and surgreffe operations. At harvest, 6 pearl quality traits were recorded and compared: surface defects, luster, grade, darkness level, and visual color. Analyzing the quality traits of pearls harvested in the initial graft process and those of pearls obtained from surgreffe experiments allowed a comparison of the influence of pearl sac cells originating from the initial mantle graft, which aged together with their recipient oysters. The results demonstrated a significant decrease between these successive grafts in luster, grade (A-B-C,) darkness level, and green color – traits that are of major importance in the pearl market. The duplicated graft experiment allowed the comparison of donor oyster families at 2 and 5 years old, where a mantle graft was inserted into recipient oysters aged 2.5 years old. The results showed the same tendencies to a lesser extent, with 1) an improved pearl grade, predominantly through a most important rate of 0 surface defect category, and 2) a green / grey ratio in favor of the younger donor. A comparison between the graft-surgreffe and the duplicated graft experiments also highlighted: 1) the indirect role played by the younger recipient oysters, which must be optimized for optimal pearl quality realization, and 2) the complex interplay between donor and recipient oysters.

The black-lipped pearl oyster Pinctada margaritifera is of high economic importance in French Polynesia. Arecent decline in pearl value led to the initiation of several genetic breeding programs aiming to improve production traits, including oyster shell growth, which dictates the time of grafting, size of the implanted nuclei, and biomineralization capacities. We assessed shell diameters on hatchery-produced spat to analyze juvenile growth performance of four half-sib families derived from polyandry (one dam crossed with two sires) and polygyny (two dams crossed with one sire) using gametes obtained by thermal shock or gonad scarification. Spat growth was monitored over 3 mo, with shell diameter measured weekly. Results revealed that the spawning method had no significant effect on juvenile growth; however, the half-sib families produced with the polygyny mating design showed significant differences in average shell diameter between dams throughout the experiment, whereas none were observed between sires with the polyandry mating design. Precocious larval size selection within each family was performed by separation into batches of small, medium, and large size, and their maintenance through juvenile stages, providing the possibility for early growth selection. These findings are important for genetic breeding programs (1) as breeding of sires and dams exhibiting the most colorful inner shell phenotypes would be possible with the gonad scarification spawning procedure without affecting overall growth performance and (2) because P. margaritifera is a protandrous hermaphrodite species, genetic selection strategies in the pearl industry must take into account the differential influence of polygyny and polyandry mating designs.

International audience; n French Polynesia, the aquaculture of Pinctada margaritifera is carried out in numerous grow-out sites, located over three archipelagos (Gambier, Society and Tuamotu). To evaluate the impact of macro-geographical effects of these growing sites on pearl quality traits, five hatcheries produced families were used as homogeneous donor oysters in an experimental graft. The molluscs were then reared in two commercial locations: Tahaa Island (Society) and Rangiroa atoll (Tuamotu). At harvest, eight pearl quality traits were recorded and compared: surface defects, lustre, grade, circles, shape categories, darkness level, body and secondary colour and visual colour categories. Overall inter-site comparison revealed that: (1) all traits were affected by grow-out location except for lustre and round shape, and (2) a higher mean rate of valuable pearls was produced in Rangiroa. Indeed, for pearl grade, Rangiroa showed twice as many A-B and less reject samples than Tahaa. This was related to the number of surface defects (grade component): in Rangiroa, twice as many pearls had no defects and less pearls had up to 10 defects. Concerning pearl shape, more circled and baroque pearls were found in Tahaa (+10%). For colour variation, 10% more pearls have an attractive green overtone in Rangiroa than in Tahaa, where more grey body colour were harvested. Lustre does not seem to be affected by these two culture site (except at a family scale). This is the first time P. margaritifera donor family have been shown to vary in the quality of pearls they produce depending on their grow-out location.

International audience; Spat collecting of the black lip oyster (Pinctada margaritifera) is the foundation of cultured black pearl production, the second source of income for French Polynesia. To understand spat collecting temporal and spatial variations, larval supply and its origin need to be characterized. To achieve this, it is necessary to account for the stock of oysters, its distribution and population characteristics (size distribution, sex ratio). While the farmed stock in concessions can be easily characterized, the wild stock is elusive. Here, we investigate the distribution and population structure of the wild stock of Ahe and Takaroa atolls using fine-scale bathymetry and in situ census data. Stocks were surprisingly low (similar to 666,000 and similar to 1,030,000 oysters for Ahe and Takaroa respectively) considering these two atolls have both been very successful spat collecting atolls in the past. Furthermore, in Ahe atoll, wild populations are aging with a dominant but small female population. Comparison with the cultured stock population (-14 millions oysters) and its dominant young male population suggests that to maximize larval supply and spat collecting on the long term, it would be useful to increase the number of females in selected sanctuaries. We discuss the implication of our findings for the long-term management of stocks and for spat collection in pearl farming atolls, and for on-going numerical modelling studies on larval dispersal.

International audience; The paper presents a state of the art on knowledge of the black pearl oyster aquaculture at the scale of french Polynesia. lt also introduces the main results obtained throughout the multi-partners POLYPERL project. The goal of this project was to develop an integrated and participative action-research of thepearl oyster culture system, focusing on the envirorunental, technological, economical and societal dimensions impacting the industry. The research proposed ranges from the undentanding of biological phenomena of the production system to socio-economic aspects and governance of the industry,taking into account the management of anthropogenic, climate and health risks. we lntroduce here the different papers from the POLYPERL project that are compiled in this volume of ECSS dealing with recent scientific work that can contribute to sustainability of the pearl industry ln French Polynesia.

Biofouling increases the operational and economic costs associated with pearl production. As current procedures for reducing oyster biofouling can be detrimental to survival and growth and may pollute the surrounding environment developing alternative, biologically-mediated, methods could potentially increase both production and ecological sustainability. With this in mind, the present study investigated natural cleaning of black-lipped pearl oysters, Pinctada margaritifera , by butterflyfishes ( Chaetodon ). The feeding behaviour of six butterflyfish species was examined at Rangiroa Atoll, French Polynesia: Chaetodon auriga , Chaetodon citrinellus , Chaetodon ephippium , Chaetodon lunulatus , Chaetodon trifascialis and Chaetodon ulietensis . All species cleaned the surface of pearl oysters by removing epibionts (from 16% to 40% of total biomass), although dietary variation may explain different cleaning efficiencies. Generalist omnivores ( C. auriga , C. citrinellus , C. ephippium and C. ulietensis ) were the most efficient cleaners (% cleaning range: 26–40% of total biomass). Within this group, C. ephippium removed the most biomass (average of 41%) targeting algae and anemones. However, C. auriga targeted the most diverse range of epibionts, removing significant amounts of algae, sponges, tunicates, and anemones. These results suggest that foraging by butterflyfishes can substantially reduce biofouling on economically-important tropical bivalves.

International audience; Breeding strategies and parental effects play a major role in genetic selection programmes for quality trait improvement. The black-lip pearl oyster, Pinctada margaritifera, is a protandrous hermaphrodite species in which the number of mature females is low, and these only appear after 4 years of culture. This difference underlines the importance of studying the contributions of dams and sires in the production and selection of donor pearl oyster families to choose the most appropriate breeding strategies for cultured pearl quality trait improvement. Laboratory crosses were made using wild dams and sires to produce half-sib families from polyandry and polygyny mating designs. The resulting half-sib families were used as donor oysters in a large grafting experiment of 900 grafts with random recipient oysters, using nuclei of the same size and brand, the same operating technician and the same grow-out site. After 18 months of culture, 497 cultured pearls were harvested. Comparison between half-sib pairs revealed that for nacre thickness, weight and shape, a polygyny effect exists, whereas no significant effect was observed for the polyandry mating. By contrast, no evidence of dam- or sire-related benefits was found for the presence/absence of circles, darkness level and colour categories or grade and its components: lustre and surface defects. These results should have important implications for the selective breeding of donor pearl oysters for hatchery management.

Pearl oyster grafting is a complex surgical operation that should lead to pearl formation after approximately eighteen months. Although this technique has been used for many years in French Polynesia, the grafting process is still not standardised. While studies have been carried out in order to improve graft performance and yield, these remain highly variable due to post-grafting mortality, nucleus rejection and unreliable pearl quality, all of which constrain pearl farm profitability. The present study uses histological analysis to monitor oysters that either rejected or retained their nuclei. Both groups of oysters are compared in terms of evolution of the graft, which could influence retention, and the development of a pearl sac in cases where grafting was successful. Data show that rejection phenomena are linked to a number of causes, notably an inflammatory reaction in the "receiving" oyster, the presence of numerous tissue lesions and the quality of the grafted tissue. These results suggest that study is needed on the different concomitant elements of the grafting process: the graft "donor" oysters, the nucleus and the "receiving" oyster and their interactions.

International audience; The present study investigated the effect of electrolysis on the biomineralization capacities of juveniles of the mollusk Pinctada argaritifera for the first time. Size-selected individuals from two groups, “Medium” and “Large”, from a multi-parental family produced in a hatchery system were subjected to electrolysis under a low voltage current over a nine-week experimental period. The growth of the juveniles was individually monitored and assessed weekly by wet weight and shell height measurements. At the end of the experiment, mantle tissue was sampled for biomineralization-related gene expression analysis. Electrolysis significantly increased pearl oyster growth in terms of shell height and wet weight for Large juveniles from the 5th and the 2nd week, respectively, until the end of the experiment. However, differences were only significant for Medium individuals from the 7th week for shell height and from the 9th week for wet weight. Furthermore, transcriptional analysis of six known biomineralization genes coding for shell matrix proteins of calcitic prisms and/or nacreous shell structures revealed that five were significantly overexpressed in the mantle mineralizing tissue under electrolysis: three in common between the two size class groups and two that were expressed exclusively in one or the other group. Finally, we found no statistical difference of the shell thickness ratio between individuals undergoing electrolysis and control conditions. Taken together, our results indicate, for the first time in a calcifying marine organism, that electrolysis influences molecular mechanisms involved in biomineralization and may stimulate some parameters of pearl oyster growth rate.

Cultured pearls are human creations formed by inserting a nucleus and a small piece of mantle tissue into a living shelled mollusc, usually a pearl oyster. Although many pearl observations intuitively suggest a possible rotation of the nucleated pearl inside the oyster, no experimental demonstration of such a movement has ever been done. This can be explained by the difficulty of observation of such a phenomenon in the tissues of a living animal. To investigate this question of pearl rotation, a magnetometer system was specifically engineered to register magnetic field variations with magnetic sensors from movements of a magnetic nucleus inserted in the pearl oyster. We demonstrated that a continuous movement of the nucleus inside the oyster starts after a minimum of 40 days post-grafting and continues until the pearl harvest. We measured a mean angular speed of 1.27° min −1 calculated for four different oysters. Rotation variability was observed among oysters and may be correlated to pearl shape and defects. Nature's ability to generate so amazingly complex structures like a pearl has delivered one of its secrets.

The top aquaculture species in French Polynesia is Pinctada margaritifera , a mollusc grown for the production of a unique gem: the black pearl. One of the challenges facing the pearl farming industry is to “produce less but better pearls” through genetic improvement. An experimental hatchery system was used to generate full-sib families to be tested for their potential as donor “oysters”. A large-scale grafting experiment was done and seven cultured pearl quality traits: grade, surface defects, lustre, darkness level, visual colour categories, circles and shape categories were recorded. Our results revealed, for the first time, significant phenotypic relationships between these quality traits. The grade A cultured pearl class had the largest proportion of pearls with a green overtone (65%), the lowest number of circled pearls (15%) and the maximum of round-shaped pearls (45%). In contrast, the “reject” cultured pearl class had the largest proportion of pearls with grey body color (65%), the greatest number of circled pearls (35%) and the maximum with a baroque shape (nearly 60%). When grade components were studied separately, cultured pearls in the zero surface defect class exhibited the same tendencies as grade A pearls, contrasting with the class where there were more than ten defects on the surface of each pearl. When cultured pearls were classified according to the presence or absence of lustre, pearls with lustre mostly had a green overtone colour, whilst pearls without lustre did not. These findings have major implications for cultured pearl quality improvement, as modern genetic breeding methods can increase the proportion of high quality cultured pearls though selected lines of donor oysters capable of producing pearls with a green overtone. Selection of appropriate donor phenotypes, incorporation of pigmentation traits into a pearl oyster breeding programme and production of lines with desirable colours will be developed for oyster aquaculture in French Polynesia.

Pearl oyster aquaculture is a major activity in French Polynesia atoll lagoons. After the economic decline that characterized the last decade, concerns recently rose about discarded installations and materials that supported aquaculture practices and by facilities abandoned after they had to close their activities. In May 2013, a first inventory of the type and amount of pearl farms derelict gear (PFDG) was achieved on 47 sites in Ahe lagoon. Surveys were conducted within and outside the boundaries of aquaculture concessions. Twenty types of PFDG littered the lagoon floor and the water column. The most impacted areas were near abandoned grafting houses with up to nine types of PFDG. Forty-five percent of the sites were impacted, including outside concessions. While management authorities are fully aware of the problem, this first assessment is a wake-up call to stimulate the cleaning of lagoons, enhance awareness among farmers, and identify potential ecological consequences on lagoon ecosystems.

Individual Pinctada margaritifera molluscs were collected from the Takapoto atoll (Tuamotu Archipelago, French Polynesia) and used to produce ten first generation full-sib families in a hatchery system, following artifi- cial breeding protocols. After three years of culture, these progenies were transferred to Rangiroa atoll (Tuamotu Archipelago, French Polynesia) and tested for their potential as graft donors. A large-scale grafting experiment of 1500 grafts was conducted, in which a single professional grafter used ten individual donor oysters from each of the ten families, grafting 15 recipient oysters from each donor. The recipient oysters were all obtained from wild spat collec- tion in Ahe (Tuamotu Archipelago, French Polynesia). After 18 months of culture, 874 pearls were harvested. Highly significant donor family effects were found for nucleus retention, nacre thickness, nacre weight, pearl colour darkness and visually-perceived colour (bodycolor and overtone), pearl shape categories, surface defects and lustre, the last two of which are components of the Tahitian classification grade. No significant difference was recorded between the ten G1 families for the absence or presence of rings. The progenies could be ranked from "best" (i.e., the donor whose grafts produced the greatest number of grade A pearls) to the "worst". Some progenies had extreme characteristics: family B presented the greatest number of pearls with lustre (98%) and a high proportion of dark gray to black with green overtone pearls (70%). These results have important implications for the selective breeding of donor pearl oysters: it may be possible to reach a point where specific donor lines whose grafts produce pearls with specific quality traits could be identified and maintained as specific breeding lines.

11 pages; International audience

The pearl industry in French Polynesia is based on exploitation of natural stocks of the black-lip pearl oyster (Pinctada margaritifera); it generates an annual turnover of 90 million euros. Improvements in pearl quality need genetic studies to improve the populations. This pearl oyster is a protandric species, where the sex-ratio normally is biased towards males. There is an increasing interest in gender control to find the mechanisms to augment female proportions for management purposes. This review summarizes information on exogenous and endogenous factors regulating gender in this and other bivalves and concludes that P. margaritifera is a protandric hermaphrodite, developing as a male during the first two years and without evidence of an effect from abiotic and biotic factors on gender during this phase. Later, pearl oysters progressively change to females, reaching a sex ratio close to 1:1 in specimens >8 years; at this stage, gender is apparently influenced by environmental parameters, but particularly by stress. Future research should seek to accurately determine the effect of temperature and food on sex ratios. Studies should be performed to characterize genes responsible for expression of gender. The use of hormones is a path that might be explored to influence the gender of pearl oysters.

International audience; Animals use sensory stimuli either to assess and select habitats, mates or food, as well as for communication. The present study aimed to understand the behavioural processes enabling several Chaetodon species (butterflyfishes) to locate one of their food sources (epibionts present on pearl oyster shells) at Rangiroa atoll (French Polynesia). Among the five species tested, our 2-channel choice flume chamber experiments identified three species that were attracted to their food source by chemical stimuli. HPLC experiments showed that pearl oysters and epibionts have specific and unique chemical fingerprints, either one or nine specific peaks, respectively. Overall, chemical stimuli are emitted by both epibionts (used direct ly by Chaetodon auriga, C. lunula and C. citrinellus) and live pearl oysters (used indirectly by C. auriga and C. lunula) to locate t heir food source. Biosynthesis of these chemical stimuli could be used to artificially attract butterflyfishes to pearl oyster rearing stations in order to increase the natural cleaning of pearl oyster shells and thus reduce one large cost for this aquaculture.

A workshop organized in French Polynesia in November 2004 allowed reviewing the current methods to model the three-dimensional hydrodynamic circulation in semi-enclosed atoll lagoons for aquaculture applications. Mollusk (e.g. pearl oyster, clam) aquaculture is a major source of income for South Pacific countries such as French Polynesia or Cook Islands. This aquaculture now requires a better understanding of circulation patterns to improve the spatial use of the lagoons, especially to define the best area to set larvae collectors. The pelagic larval duration of the relevant species (< 20 days) and the size of the semi-closed lagoons (few hundreds of km 2) drive the specifications of the model in terms of the spatial and temporal scale. It is considered that, in contrast with fish, mollusk larvae movements are limited and that their cycle occurs completely in the lagoon, without an oceanic stage. Atolls where aquaculture is productive are generally well-bounded, or semi-closed, without significant large and deep openings to the ocean. Nevertheless part of the lagoon circulation is driven by oceanic water inputs through the rim, ocean swells, tides and winds. Therefore, boundary conditions of the lagoon system are defined by the spatial structure of a very shallow rim (exposition and number of hoas), the deep ocean swell climate, tides and wind regimes. To obtain a realistic 3D numerical model of lagoon circulation with adequate forcing, it is thus necessary to connect in an interdisciplinary way a variety of methods (models, remote sensing and in situ data collection) to accurately represent the different components of the lagoon system and its specific boundary conditions. We review here the current methods and tools used to address these different components for a hypothetical atoll of the Tuamotu Archipelago (French Polynesia), representative of the semi-closed lagoons of the South Pacific Ocean. We hope this paper will serve as a guide for similar studies elsewhere and we provide guidelines in terms of costs for all the different stages involved. (c) 2006 Elsevier Ltd. All rights reserved.