Marine animal forests (MAFs) are benthic ecosystems characterised by biogenic three-dimensional structures formed by suspension feeders such as corals, gorgonians, sponges and bivalves. They comprise highly diversified communities among the most productive in the world's oceans. However, MAFs are in decline due to global and local stressors that threaten the survival and growth of their foundational species and associated biodiversity. Innovative and scalable interventions are needed to address the degradation of MAFs and increase their resilience under global change. Surprisingly, few studies have considered trophic interactions and heterotrophic feeding of MAF suspension feeders as an integral component of MAF conservation. Yet, trophic interactions are important for nutrient cycling, energy flow within the food web, biodiversity, carbon sequestration, and MAF stability. This comprehensive review describes trophic interactions at all levels of ecological organisation in tropical, temperate, and cold-water MAFs. It examines the strengths and weaknesses of available tools for estimating the heterotrophic capacities of the foundational species in MAFs. It then discusses the threats that climate change poses to heterotrophic processes. Finally, it presents strategies for improving trophic interactions and heterotrophy, which can help to maintain the health and resilience of MAFs.
The distribution of symbiotic scleractinian corals is driven, in part, by light availability, as host energy demands are partially met through translocation of photosynthate. Physiological plasticity in response to environmental conditions, such as light, enables the expansion of resilient phenotypes in the face of changing environmental conditions. Here we compared the physiology, morphology, and taxonomy of the host and endosymbionts of individual Madracis pharensis corals exposed to dramatically different light conditions based on colony orientation on the surface of a shipwreck at 30 m depth in the Bay of Haifa, Israel. We found significant differences in symbiont species consortia, photophysiology, and stable isotopes, suggesting that these corals can adjust multiple aspects of host and symbiont physiology in response to light availability. These results highlight the potential of corals to switch to a predominantly heterotrophic diet when light availability and/or symbiont densities are too low to sustain sufficient photosynthesis, which may provide resilience for corals in the face of climate change.
Global warming endangers reef-building corals as they lose their photosynthetic symbionts, which limits their ability to feed autotrophically. Consequently, heterotrophy, the capture of zooplankton, can become crucial for the energy budget of heat-stressed corals. However, it is difficult to assess the extent of the heterotrophic contribution in corals, as well as the dynamics of nutrient exchange between the host and its symbionts. In this pioneering study, we employed a suite of isotopic markers, including 13C- and 15N bulk tissue isotope measurements, compound-specific isotope analysis of amino acids (CSIA-AAs), and 13C- and 15N-labeled food incubations, to investigate nutrient acquisition and allocation in the coral Stylophora pistillata under controlled and heat-induced bleaching conditions. Bulk isotope values and inorganic carbon assimilation remained unchanged in the bleached corals compared to the control corals, overall indicating undisturbed autotrophic activity of the symbionts under heat stress. However, CSIA-AAs showed an increased dependence on heterotrophy for amino acid synthesis in both the host and the symbionts despite reduced assimilation of 15N-labeled food. Overall, these results suggest that although S. pistillata reduces its assimilation of heterotrophic food under heat stress, the acquisition of amino acids by the coral host and symbionts still relies on heterotrophy. This study emphasizes the importance of using multiple indicators to gain a comprehensive understanding of coral nutrition. It shows that coral dependence on heterotrophy is not only associated with a decline in autotrophic availability. Rather, it demonstrates the ability of S. pistillata to adapt its utilization of food sources to the prevailing environmental conditions.IMPORTANCEThis work highlights that every isotopic marker displays a piece of different information concerning the diet of the model coral S. pistillata. By combining all markers, we observed that although S. pistillata exhibited reduced heterotrophic assimilation under heat stress, amino acid acquisition and synthesis remained dependent on heterotrophy. The findings emphasize the adaptability of corals in utilizing different food sources, which is vital for their resilience and recovery in changing environmental conditions. This research underscores the complexity of coral symbiosis and highlights the need for multiple indicators to understand dietary dynamics comprehensively.
Extreme weather events are increasing in frequency and magnitude. Consequently, it is important to understand their effects and remediation. Resilience reflects the ability of an ecosystem to absorb change, which is important for understanding ecological dynamics and trajectories. To describe the impact of a powerful storm on coral reef structural complexity, we used novel computational tools and detailed 3D reconstructions captured at three time points over three years. Our data-set Reefs4D of 21 co-registered image-based models enabled us to calculate the differences at seven sites over time and is released with the paper. We employed six geometrical metrics, two of which are new algorithms for calculating fractal dimension of reefs in full 3D. We conducted a multivariate analysis to reveal which sites were affected the most and their relative recovery. We also explored the changes in fractal dimension per size category using our cube-counting algorithm. Three metrics showed a significant difference between time points, i.e., decline and subsequent recovery in structural complexity. The multivariate analysis and the results per size category showed a similar trend. Coral reef resilience has been the subject of seminal studies in ecology. We add important information to the discussion by focusing on 3D structure through image-based modeling. The full picture shows resilience in structural complexity, suggesting that the reef has not gone through a catastrophic phase shift. Our novel analysis framework is widely transferable and useful for research, monitoring, and management.
As impacts on the Mediterranean Sea are expected to grow in the future, especially with climate change, habitat degradation, and displacement of native species by non-indigenous species (NIS), the investigation of significant alterations to trophic levels in this diverse marine habitat is important. Analysis of stable isotopes from targeted consumers has previously been shown to reliably reflect that of primary producers, thus enabling us to describe and highlight potential shifts in the food web of a particular ecosystem. In this study, we used δ 13 C values of essential amino acids (AA) in order to examine the dietary composition of established non-native, Lessepsian fish migrants in the Eastern Mediterranean Sea compared to that of the same fish species from their original population in the Gulf of Aqaba, Red Sea. Our data show that a clear variance in carbon isotopic signatures exists in food sources consumed by the same species between the different environments, with the exception of the classic herbivore, Siganus rivulatus (Forsskål & Niebuhr, 1775), whose very similar isotopic patterns reflect the algal source they predominantly consume in both locations. With the results of this research, we propose that Lessepsian fishes with the ability to maintain their nutritional patterns, though not necessarily that of their original food source, will acclimatize better in their new habitat. Consequences of flourishing Lessepsian fish populations include a further tropicalization of the Eastern Mediterranean Sea and the likely restructuring of local food webs.
Global warming raises seawater temperatures and creates changes which have been found to affect the movement of large migrating marine species. Understanding the thermal niches of marine species could prove essential to anticipate how the future climate will alter migrations, and how conservation efforts will have to change accordingly. Orot Rabin power station in Hadera, Israel uses seawater to cool its turbine and releases the warm water back into the Mediterranean Sea. As a result, a marine area with artificially elevated temperatures is created around the effluent. Every winter in the past two decades, this area attracts sharks of two species, Carcharhinus obscurus and Carcharhinus plumbeus, presumably to spend the cold months at a higher temperature. This study concentrated on this point of artificial heat dissipation, which maintains a wide gradient of surface temperatures and allowed us to examine the temperature preferences of these species when given a larger range than what is naturally found in the sea. Between 2016 and 2018, 16 sharks were tagged with acoustic tags, 3 of which had temperature sensors, and 2 were additionally tagged with pop-up archival tags also logging temperature data. Results show that the sharks stayed in the elevated temperature, while the ambient sea was cold during the winter, spending several months in the heated area. Both species displayed a similar preferred range, spending 90 percent of their time at a temperature between 21.8 °C and 26.1 °C while the surrounding sea was 15.5–25.5 °C. Considering this chosen thermal niche and the rise in water temperature, it appears that for the past 40 years, the Eastern shores of the Mediterranean have become more suitable for these species, especially during transitional seasons. The question that arises, however, is whether these shark populations will benefit from the expanding range of preferable temperatures, or whether their proximity to shorelines will put them at greater risk in terms of human activities such as fishing and pollution.
Sedimentary marine habitats are the largest ecosystem on our planet in terms of area. Marine sediment microbiota govern most of the benthic biological processes and therefore are responsible for much of the global biogeochemical activity. Sediment microbiota respond, even rapidly, to natural change in environmental conditions as well as disturbances of anthropogenic sources. The latter greatly impact the continental shelf. Characterization and monitoring of the sediment microbiota may serve as an important tool for assessing environmental health and indicate changes in the marine ecosystem. This study examined the suitability of marine sediment microbiota as a bioindicator for environmental health in the eastern Mediterranean Sea. Integration of information from Bacteria, Archaea, and Eukaryota enabled robust assessment of environmental factors controlling sediment microbiota composition: seafloor-depth (here representing sediment grain size and total organic carbon), core depth, and season (11%, 4.2%, and 2.5% of the variance, respectively). Furthermore, inter- and intrakingdom cooccurrence patterns indicate that ecological filtration as well as stochastic processes may control sediment microbiota assembly. The results show that the sediment microbiota was robust over 3 years of sampling, in terms of both representation of region (outside the model sites) and robustness of microbial markers. Furthermore, anthropogenic disturbance was reflected by significant transformations in sediment microbiota. We therefore propose sediment microbiota analysis as a sensitive approach to detect disturbances, which is applicable for long-term monitoring of marine environmental health. IMPORTANCE Analysis of data, curated over 3 years of sediment sampling, improves our understanding of microbiota assembly in marine sediment. Furthermore, we demonstrate the importance of cross-kingdom integration of information in the study of microbial community ecology. Finally, the urgent need to propose an applicable approach for environmental health monitoring is addressed here by establishment of sediment microbiota as a robust and sensitive model.
Two species of carcharhinid sharks aggregate every winter at the warm water effluent of a coastal power plant on the Israeli Mediterranean coast. The two species (Carcharhinus obscurus and Carcharhinus plumbeus) cooccur in a highly confined area for several months every year and are highly associated with the area in and around the hot water effluent. Niche partitioning has recently been suggested as a mechanism that enables the coexistence of similar shark species by resource partitioning, spatial partitioning, and temporal partitioning. In this study, we used acoustic telemetry to study the individual diel movement and activity patterns within this enclosed area and examined the differences between the two species sharing it. Although this location only reaches a maximum depth of 7.5 m, we found both species perform a diel vertical movement, rising closer to the surface at night and moving deeper during daytime. Furthermore, the two shark species swam at different depths both day and night, with C. obscurus swimming in the upper column, about 2 m shallower than C. plumbeus. The very small scale of movement, which nearly equals the sharks’ body length, suggests movement patterns might be conserved at the species level. Moreover, spatiotemporal differences between the two species may reflect a mean of interspecific partitioning that occurs even in a highly confined and shallow habitat.
Black corals (Antipatharia) are among the most common and diverse taxa in mesophotic and aphotic rocky habitats. Studies of their ecology and biology are limited mostly because of the technical challenges involved in deep-water work. Here, we describe taxonomic traits, habitat, and nutrition of 2 species from the upper mesophotic zone of the northern Red Sea, Gulf of Aqaba, Israel. Our study describes a broader distribution of the branched Antipathes griggi, which to date has been primarily found in the Hawaiian Archipelago. Additionally, a new record is reported regarding the coiled Stichopathes spp. discovered at the upper mesophotic zone. The branched A. griggi were more abundant than the coiled Stichopathes spp. in all explored sites, and differences in light and water flow conditions significantly affected their distribution. Both species lacked symbiotic algae (Symbiodiniaceae). Analysis of the amino acid compound-specific stable isotope signature (AA-CSIA) revealed that both antipatharian species show a high trophic position relative to known mixotrophic and heterotrophic corals. Although both species inhabit the same area and are planktivorous, their carbon signature suggests they have different carbon sources. Our findings imply that antipatharians have a vital role in the trophic dynamics of mesophotic coral ecosystems, a unique habitat whose ecology remains largely unexplored.
Coral bleaching, the breakdown of the coral-Symbiodiniaceae association has been identified as a major cause of coral reef decline worldwide. When symbiont functions are compromised, corals receive fewer photosynthetic products from their symbionts and suffer significant starvation along with changes in nutrient cycling. Not all coral species are equally susceptible to bleaching, but despite intensive research, our understanding of the causes for coral bleaching remains incomplete. Here, we investigated nutrient exchange between host and symbionts of two coral- Symbiodiniaceae associations that are differentially susceptible to bleaching when maintained under heterotrophy in the dark. We followed the fate of heterotrophic nutrients using bulk isotope and compound-specific (amino acid) isotope analyses. We showed that symbiont starvation is a major cause of symbiotic breakdown in the dark. While Oculina patagonica transferred almost all heterotrophically-acquired amino acids within two weeks in the dark to its symbionts and did not bleach, Turbinaria reniformis, transferred only 2 amino acids to its symbionts after 4 weeks in the dark, and experienced significant bleaching. These results pave the way for future studies on the role of nutrition in coral stress response and the importance of maintaining a healthy symbiont population to avoid coral bleaching.
Globally, species are migrating in an attempt to track optimal isotherms as climate change increasingly warms existing habitats. Stony corals are severely threatened by anthropogenic warming, which has resulted in repeated mass bleaching and mortality events. Since corals are sessile as adults and with a relatively old age of sexual maturity, they are slow to latitudinally migrate, but corals may also migrate vertically to deeper, cooler reefs. Herein we describe vertical migration of the Mediterranean coral Oculina patagonica from less than 10 m depth to > 30 m. We suggest that this range shift is a response to rapidly warming sea surface temperatures on the Israeli Mediterranean coastline. In contrast to the vast latitudinal distance required to track temperature change, this species has migrated deeper where summer water temperatures are up to 2 °C cooler. Comparisons of physiology, morphology, trophic position, symbiont type, and photochemistry between deep and shallow conspecifics revealed only a few depth-specific differences. At this study site, shallow colonies typically inhabit low light environments (caves, crevices) and have a facultative relationship with photosymbionts. We suggest that this existing phenotype aided colonization of the mesophotic zone. This observation highlights the potential for other marine species to vertically migrate.
The association between corals and photosynthetic dinoflagellates is one of the most well-known nutritional symbioses, but nowadays it is threatened by global changes. Nutritional exchanges are critical to understanding the performance of this symbiosis under stress conditions. Here, compound-specific δ15N and δ13C values of amino acids (δ15NAA and δ13CAA) were assessed in autotrophic, mixotrophic and heterotrophic holobionts as diagnostic tools to follow nutritional interactions between the partners. Contrary to what was expected, heterotrophy was mainly traced through the δ15N of the symbiont’s amino acids (AAs), suggesting that symbionts directly profit from host heterotrophy. The trophic index (TP) ranged from 1.1 to 2.3 from autotrophic to heterotrophic symbionts. In addition, changes in TP across conditions were more significant in the symbionts than in the host. The similar δ13C-AAs signatures of host and symbionts further suggests that symbiont-derived photosynthates are the main source of carbon for AAs synthesis. Symbionts, therefore, appear to be a key component in the AAs biosynthetic pathways, and might, via this obligatory function, play an essential role in the capacity of corals to withstand environmental stress. These novel findings highlight important aspects of the nutritional exchanges in the coral–dinoflagellates symbiosis. In addition, they feature δ15NAA as a useful tool for studies regarding the nutritional exchanges within the coral–symbiodiniaceae symbiosis.
As the devastating impacts of global climate change and local anthropogenic stressors on shallow-water coral reefs are expected to rise, mesophotic coral ecosystems have increasingly been regarded as potential lifeboats for coral survival, providing a source of propagules to replenish shallower reefs. Yet, there is still limited knowledge of the capacity for coral larvae to adjust to light intensities that change with depth. This study elucidates the mechanisms underlying plasticity during early life stages of the coral Porites astreoides that enable survival across broad depth gradients. We examined physiological and morphological variations in larvae from shallow (8–10 m) and mesophotic (45 m) reefs in Bermuda, and evaluated differences in survival, settlement patterns and size among recruits depending on light conditions using a reciprocal ex situ transplantation experiment. Larvae released from mesophotic adults were found to have significantly lower respiration rates and were significantly larger than those derived from shallow adults, indicating higher content of energetic resources and suggesting a greater dispersal potential for mesophotic larvae compared to their shallow counterparts. Additionally, larvae released from mesophotic adults experienced higher settlement success and larger initial spat size compared to larvae from shallow adults, demonstrating a potential connection between parental origin, offspring quality, and recruitment success. Although both shallow and mesophotic larvae exhibited the capacity to survive and settle under reciprocal light conditions, all larvae had higher survival under mesophotic light conditions regardless of parental origin, suggesting that conditions experienced under low light may enable longer larval life, further extending the dispersal period. These results indicate that larvae from mesophotic Porites astreoides colonies are likely capable of reseeding shallow reefs in Bermuda, thereby supporting the Deep Reef Refugia Hypothesis.
Coral recruitment represents a key element for coral reef persistence and resilience in the face of environmental disturbances. Studying coral recruitment patterns is fundamental for assessing reef health and implementing appropriate management strategies in an era of climate change. The FluorIS system has been developed to acquire high resolution, wide field-of-view (FOV) in situ images of coral recruits fluorescence and has proven successful in shallow reef environments. However, up to now, its applicability to mesophotic coral ecosystems remains unknown due to the complexity of the system and the limited time available when working at mesophotic depth. In this study we optimized the FluorIS system by utilizing a single infrared-converted camera instead of the bulkier regular dual-camera system, substantially reducing the system complexity and significantly decreasing the acquisition time to an average of 10 s for a set of 3 images. Moreover, the speed-FluorIS system is much more economical, decreasing the cost of the full set-up by roughly 40% compared to the original dual-camera system. We tested the utility of the speed-FluorIS by surveying coral recruits across shallow and mesophotic reefs of the Red Sea (Gulf of Eilat) and Bermuda, two of the most northerly reefs in the world with markedly different substrate and topography, and demonstrate that the modified system enables fast imaging of fluorescence to study coral recruitment patterns over a broader range of depths and reef topographies than previous fluorescence methods. Our single-camera system represents a valuable, non-invasive and rapid underwater tool which will help standardize surveys and long-term monitoring of coral recruits, contributing to our understanding of these vital and delicate early life stages of corals.
Amino acids compound-specific nitrogen stable isotope analysis (AA-CSIA) is an emerging tool in ecology for understanding trophic system dynamics. While it has been successfully used for several independent studies across a range of environments and study locations, researchers have encountered calculation issues for determining trophic position values. Most studies introduce modifications to the constants of trophic position equation calculations, but then fail to account for the equation variations when comparing across separate research studies. The broad acceptance of this approach is anchored in an underlying presumption that no addition of the exogenous nitrogen atom occurs in the different methods; and therefore, such variations should not affect the outcome. In this paper, we evaluate the use of the EZfaast amino acid derivatization kit (chloroformate) and compare it to the isotopic results of two other derivatization methods. We highlight new considerations for working with AA-CSIA that might account for some of the variations in the results and lead researchers to modify constants in the equation. This study concludes that developing unique constants per derivatization method is required to have more accurate cross-study comparisons of trophic positions.
The NATO SPS multi-year project ThreatDetect investigates an autonomous platform to secure marine infrastructures by reliably detecting divers and mines in real time. Our system combines acoustic remote detection with verification using pattern recognition on underwater imagery. For diver detection, we rely on active acoustics from a single transceiver, and analyze the acoustic reflections to detect and localize a target that fits the pattern of a diver. For mine detection, we segment sonar images from an autonomous underwater vehicle (AUV) to differentiate between background, highlight, and shadow. In case of detection, we steer the AUV’s trajectory so as to closely observe the target, and transmit segmented sonar images to a surface station via underwater acoustic communications. At the time of writing, the project is performing final technology tuning and integrated sea experiments.
The Indo-Pacific gorgonian coral Melithaea erythraea (Melithaeidae, previously Acabaria) was first recorded in the Mediterranean in 1999 in the harbor of the Hadera power station, Israel. This species is the only octocoral known to have invaded the Mediterranean Sea. In the past two decades, it has demonstrated a stable population in this harbor, and never found outside this location, not even on the adjacent natural rocky reefs. Then, during 2015, several specimens of M. erythraea were found on a natural substrate at Nahsholim, Israel, about 23 km north of the power station. This is the first evidence of this coral's existence beyond the power plant harbor. The number of colonies there suggests that the population is sustainable, but further study is needed. Although no genetic differences were found among specimens from Nahsholim, Hadera, and the Red Sea, their bacterial epibiota has undergone change following migration. The carbon source and trophic position are similar between the Red Sea and the Mediterranean, suggesting that there is no nutritional barrier to migration for the coral. The larval connectivity model supports the hypothesis that the planulae that have settled in Nahsholim originated from the Port of Hadera, although other arrival scenarios are also possible. The spread of this species suggests that the eastern Mediterranean is becoming increasingly suitable for migrating corals, joining the many other Indo-Pacific migrants that have already established populations there.
Energy sources of corals, ultimately sunlight and plankton availability, change dramatically from shallow to mesophotic (30–150 m) reefs. Depth-generalist corals, those that occupy both of these two distinct ecosystems, are adapted to cope with such extremely diverse conditions. In this study, we investigated the trophic strategy of the depth-generalist hermatypic coral Stylophora pistillata and the ability of mesophotic colonies to adapt to shallow reefs. We compared symbiont genera composition, photosynthetic traits and the holobiont trophic position and carbon sources, calculated from amino acids compound-specific stable isotope analysis (AA-CSIA), of shallow, mesophotic and translocated corals. This species harbors different Symbiodiniaceae genera at the two depths: Cladocopium goreaui (dominant in mesophotic colonies) and Symbiodinium microadriaticum (dominant in shallow colonies) with a limited change after transplantation. This allowed us to determine which traits stem from hosting different symbiont species compositions across the depth gradient. Calculation of holobiont trophic position based on amino acid δ 15 N revealed that heterotrophy represents the same portion of the total energy budget in both depths, in contrast to the dogma that predation is higher in corals growing in low light conditions. Photosynthesis is the major carbon source to corals growing at both depths, but the photosynthetic rate is higher in the shallow reef corals, implicating both higher energy consumption and higher predation rate in the shallow habitat. In the corals transplanted from deep to shallow reef, we observed extensive photo-acclimation by the Symbiodiniaceae cells, including substantial cellular morphological modifications, increased cellular chlorophyll a , lower antennae to photosystems ratios and carbon signature similar to the local shallow colonies. In contrast, non-photochemical quenching remains low and does not increase to cope with the high light regime of the shallow reef. Furthermore, host acclimation is much slower in these deep-to-shallow transplanted corals as evident from the lower trophic position and tissue density compared to the shallow-water corals, even after long-term transplantation (18 months). Our results suggest that while mesophotic reefs could serve as a potential refuge for shallow corals, the transition is complex, as even after a year and a half the acclimation is only partial.
Reef-building corals, the major producers of biogenic calcium carbonate, form skeletons in a plethora of morphological forms. Here we studied skeletal modifications of Stylophora pistillata (clade 4) colonies that dramatically adapt to different depths with decreasing light availability. They show notable transitions from spherical morphologies (shallow depths, 5 m deep) to flat and branching ones (at deep 60 m mesophotic depths). Such changes are typically ascribed to the algal photosymbiont physiological feedback with the coral host. We find typical fine-scale skeletal variability (such as extensive vs. weak development of rapid accretion deposits at shallow- and mesophotic depth morphotypes, respectively) that suggest underlying genetic modifications of biomineralization pathways of the coral host. To explain this, we conducted comparative morphology-based analyses, including microscopy, electron microscopy and spectroscopy coupled with a comprehensive transcriptomic analysis of S. pistillata samples originated from Gulf of Eilat (Red Sea, Israel), collected along a depth gradient from the sea surface (5 m deep) to mesophotic depths (up to 60m). Additional samples were experimentally transplanted from 5m to 60m and from 60m to 5 m. Interestingly, both morphologically and functionally, transplanted corals partly adapt the typical depth-specific properties. In mesophotic depths, we find that the organic matrix fraction is enriched in the coralla, results corroborated by overrepresentation of biomineralization "tool-kit" structural extracellular genes. These results provide insights into the molecular mechanisms of coral calcification and skeletal adaptation that repeatedly allowed this group to adapt to extreme environments now and in geological past.
Coral recruitment is a pivotal factor in coral reef stability and in recovery following substantial disturbances. Despite its immense importance, the study of coral recruitment has some major gaps, notably larval survival before and following settlement, mainly due to technical limitations, which stem from the difficulty in observing the minute larvae. To overcome the major limitation in coral recruitment studies, i. e. the in situ detection of recruits during their early stages, we designed a new detection set-up, composed of a fluorescence detection set-up, a grid-covered substrate, and a Geographic Information System tracking system. This set-up, enabling the identification of coral recruits soon after settlement, revealed that in the critical period of the first day, less than 45% of the settling corals may survive. The results also suggest that either coral larva select locations that may increase their survival chances or they experience dramatic mortality during the early hours of settlement, which induce a consistent pattern of spat distribution. Our study confirms an earlier speculation that the first 24 h post-settlement may determine the rates and spatial patterns of recruitment. The significant implications of these findings, and the implemented " detection set-up" for coral reef monitoring and management, are discussed.