Analyzing Adaptation Mechanisms in Artificial Transplantation of Galaxea fascicularis

Analyzing Adaptation Mechanisms in Artificial Transplantation of Galaxea fascicularis | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Analyzing Adaptation Mechanisms in Artificial Transplantation of Galaxea fascicularis He Zhao, Hongmin Wang, Jingzhao Ke, Junling Zhang, Yushan Li, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4792475/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Jun, 2025 Read the published version in Marine Biology → Version 1 posted 5 You are reading this latest preprint version Abstract Coral reefs are among Earth's most biologically diverse and ecologically crucial ecosystems but face severe threats from climate change and human activities. Coral transplantation has become a key strategy for reef restoration. This study focused on transplanting Galaxea fascicularis at northern Wuzhizhou Island, Hainan, assessing physiological characteristics and metabolomic differences between transplanted and parent corals at 1, 6, and 12 months post-transplantation. Findings revealed that transplanted coral survival rates declined rapidly during the first 6 months but then stabilized. An algal bloom in May 2023 increased turbidity, dissolved inorganic nitrogen (DIN), and partial pressure of p CO 2 , negatively impacting coral photosynthesis and calcification and increasing physiological stress. From months 6 to 12, environmental conditions improved, with temperature and salinity aligning closely with natural conditions, dissolved oxygen levels recovering, turbidity decreasing significantly, and Ω arag reaching moderate levels, facilitating stable coral growth and calcification. By 12 months, transplanted corals reached sexual maturity, with notable increases in protein and lipid content. Metabolomic analysis showed that during the short-term (1 month) and mid-term (6 months) post-transplantation periods, the arachidonic acid metabolic pathway was upregulated while the glycerophosphate metabolic pathway was downregulated, enabling corals to cope with environmental stress and resource redistribution. By 12 months, oxidative phosphorylation was upregulated to meet reproductive energy demands. Results demonstrate that G. fascicularis can adapt well to restoration environments and achieve sexual maturity quickly, making it a suitable candidate for reef restoration. Galaxea fascicularis Artificial Transplantation Coral Restoration Metabolomics Adaptation Mechanism Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction Coral reefs are among Earth's most diverse and ecologically crucial ecosystems (Castro-Sanguino et al. 2021 ), providing essential habitats, coastal protection, fisheries, and tourism benefits (Massei et al. 2023 ). They face severe threats from climate change, pollution, overfishing, and other disturbances (Niz et al. 2023 ), leading to widespread coral bleaching and mortality (Andradi-Brown et al. 2020 ). Coral transplantation is a promising conservation strategy, involving the relocation of healthy coral fragments or larvae to degraded reefs to restore ecological functions (Ferse et al. 2021 ; Lang et al., 2024). Success depends on environmental conditions and coral species (Williams et al. 2019 ). Post-transplantation, corals undergo significant physiological and metabolic changes affecting survival and adaptation (Uribe-Castañeda et al. 2018 ). Understanding these mechanisms is essential for improving transplantation techniques and restoration outcomes. Metabolomics is an advanced scientific approach that examines the physiological state and response mechanisms of organisms through a comprehensive analysis of their small molecular metabolites (Hu et al. 2020 ). This technique is instrumental in unveiling the adaptive mechanisms of corals to environmental fluctuations. For instance, Roach et al. (2001) have demonstrated that metabolomic disparities in naturally bleached corals remain consistent over time, identifying betaine lipids as key biomarkers of coral bleaching. Similarly, Zhao et al. ( 2024 ) have explored the metabolomic heterogeneity of Acropora hyacinthus under both transplantation and natural conditions, revealing that differential metabolites can drive phenotypic variations, directing more energy towards growth rather than developmental processes. By investigating the metabolomic variations among corals, we can pinpoint individuals with superior adaptability and restoration potential, thereby refining coral transplantation methodologies and bolstering the restoration and preservation of coral reef ecosystems (Schmidt-Roach et al. 2020 ). Current research predominantly focuses on the transplantation of branching corals, with relatively limited attention given to massive corals. Boström-Einarsson et al. ( 2020 ) have noted that coral reef restoration projects have been implemented in at least 56 countries, utilizing 229 species from 72 coral genera, encompassing approximately 25% of reef-building coral species. However, these projects primarily target fast-growing branching coral species, which, despite their rapid growth, are highly sensitive to environmental disturbances. In contrast, the more resilient but slower-growing massive corals are frequently overlooked (Guest et al. 2023 ). The slow growth rate of massive corals often results in restoration outcomes that are visually unimpressive and fail to meet immediate expectations, leading to their neglect (Marshall and Baird 2000 ; Hein et al. 2020 ). Additionally, the unique structural characteristics of massive corals pose significant challenges during transplantation, further contributing to the lack of research focus on these species (Edwards and Clark 1999 ). Nevertheless, massive corals play an indispensable role in coral reef ecosystems. Their robust and durable structures provide critical physical support to the reefs, creating habitats and shelter for diverse marine organisms, thereby enhancing biodiversity (Glassom et al. 2004 ). Investigating the metabolomic changes in massive corals during transplantation and assessing their long-term performance post-transplantation is essential for evaluating transplantation success, adaptive responses, and long-term survival. In the northern part of Wuzhizhou Island, Hainan, we conducted an experiment involving the transplantation of massive corals. We selected Galaxea fascicularis , the dominant coral species of Wuzhizhou Island, and transplanted it onto our custom-designed restoration device, the "Trapezoidal Reef". Samples were collected at three key intervals, 1 month, 6 months, and 12 months post-transplantation, to analyze the physiological characteristics and metabolomic differences between transplanted and parent G. fascicularis . This study aimed to elucidate the adaptive changes occurring during the transplantation process, providing fundamental scientific data for future transplantation efforts of G. fascicularis and contributing to the broader field of coral reef restoration. 2. Materials and methods 2.1 Experimental Site and Sample Collection In April 2023, we established a coral reef restoration site in northern Wuzhizhou by installing 12 trapezoidal reef devices (Fig. 1 a). Each device featured eight designated transplantation points for massive corals (Fig. 1 b). Divers meticulously collected and tagged fragments of G. fascicularis from the natural reef area. Using hammers and chisels, small coral fragments were carefully extracted and transplanted onto the trapezoidal reefs within the restoration area, resulting in a total of 96 transplanted fragments of G. fascicularis . To assess the transplantation outcomes, we collected samples at 1 month (May 2023), 6 months (October 2023), and 12 months (April 2024) post-transplantation. At each time point, six tagged parent corals and six transplanted corals were retrieved. The collected samples were divided into two portions: one part was immediately preserved in liquid nitrogen for subsequent UPLC-MS/MS analysis, while the other part was used for measuring physiological indicators. 2.2 Measurement of Coral Physiological Indicators Upon collection, coral samples were immediately transported to the laboratory for analysis. Throughout the experiment, the maximum quantum efficiency of coral (Fv/Fm) was measured using the MINI-PAM-II chlorophyll fluorometer (Walz, Germany). To assess zooxanthellae density, coral surfaces were washed with a Waterpik dental cleaner (Water-pik, WP70-EC, USA). A specific volume of the wash solution (10 mL) was measured and counted using a hemocytometer, with the process repeated eight times to obtain an average value. This method follows the protocol of Li et al. ( 2006 ). Chlorophyll a was extracted using an Addison reagent kit and quantified with a spectrophotometer, as per Fitt et al. ( 2000 ). To determine coral tissue content, samples were dried at 60℃ for 24 hours to obtain dry weight and subsequently ashed at 400℃ in a muffle furnace for 4 hours to measure ash weight. The difference in mass pre- and post-ashing represents the coral tissue content. Coral surface area was measured using the aluminum foil method (Stimson 1997 ). Carbohydrate content was determined according to Dubois et al. (1956). Ground coral samples were extracted with phenol and concentrated sulfuric acid, and their absorbance was measured at A490 using a spectrophotometer. Absorbance values were converted to concentration using a glucose standard curve. Protein extraction followed the method of Smith et al. ( 2017 ). Ground coral samples were processed with a BCA reagent kit, and their absorbance was measured at A562. Concentrations were calculated using a protein standard curve. Lipid extraction adhered to the protocol by Rodrigues and Grottoli ( 2007 ). Approximately 1 g of wet coral sample was extracted with CM solution (methanol/chloroform 1:2, v/v). About 20 mL of CM extract was added to 1 g of coral and incubated in the dark for 24 hours. The organic phase was then evaporated and dried in an oxygen-free environment to determine lipid content. All data were standardized to the coral surface area. 2.3 Metabolome Analysis 2.3.1 Metabolite Extraction A solid sample of 10 mg was accurately weighed, and metabolites were extracted using 400 µL of methanol (4:1, v/v) solution containing 0.02 mg/mL L-2-chlorophenylalanine as an internal standard. The mixture was allowed to settle at -10℃ and then processed with a high-throughput tissue crusher Wonbio-96c at 50 Hz for 6 minutes, followed by ultrasound at 40 kHz for 30 minutes at 5℃. Samples were then placed at -20℃ for 30 minutes to precipitate proteins. After centrifugation at 13,000 g at 4℃ for 15 minutes, the supernatant was carefully transferred to sample vials for LC-MS/MS analysis. 2.3.2 Quality Control (QC) Sample As part of the system conditioning and QC process, a pooled QC sample was prepared by mixing equal volumes from all samples. The QC samples were processed and analyzed in the same manner as the analytical samples. This pooled QC sample served as a representative of the entire sample set and was injected at regular intervals (every six samples) to ensure and monitor the stability and reliability of the analysis. 2.3.3 UHPLC-MS/MS Analysis The UHPLC-MS analysis was conducted using the UHPLC-Q Exactive HF-X system from Thermo Fisher Scientific. Chromatographic separation was achieved using an HSS T3 column (100 mm × 2.1 mm i.d., 1.8 µm), with 2 µL of each sample injected for MS detection. The mobile phases consisted of 0.1% formic acid in water (95:5, v/v) (solvent A) and 0.1% formic acid in acetonitrile: isopropanol (47.5:47.5:5, v/v) (solvent B). The solvent gradient was programmed as follows: from 0 to 3.5 min, 0% B to 24.5% B (0.4 mL/min); from 3.5 to 5 min, 24.5% B to 65% B (0.4 mL/min); from 5 to 5.5 min, 65% B to 100% B (0.4 mL/min); from 5.5 to 7.4 min, 100% B (0.4 mL/min to 0.6 mL/min); from 7.4 to 7.6 min, 100% B to 51.5% B (0.6 mL/min); from 7.6 to 7.8 min, 51.5% B to 0% B (0.6 mL/min to 0.5 mL/min); from 7.8 to 9 min, 0% B (0.5 mL/min to 0.4 mL/min); and from 9 to 10 min, 0% B (0.4 mL/min) to equilibrate the system. The sample injection volume was 2 µL, with a flow rate set at 0.4 mL/min. The column temperature was maintained at 40℃, and all samples were stored at 4℃ during the analysis period. 2.3.4 MS Conditions: The MS data were collected using a Thermo UHPLC-Q Exactive HF-X Mass Spectrometer equipped with an electrospray ionization (ESI) source, operating in both positive and negative ion modes. The optimal conditions for the mass spectrometer were set as follows: heater temperature at 425℃, capillary temperature at 325℃, sheath gas flow rate at 50 arb, aux gas flow rate at 13 arb, and ion-spray voltage floating (ISVF) at -3,500 V for negative mode and 3,500 V for positive mode. The normalized collision energy was set at 20-40-60 V rolling for MS/MS. The full MS resolution was configured at 60,000, while the MS/MS resolution was set at 7,500. Data acquisition was performed in Data Dependent Acquisition (DDA) mode, with detection conducted over a mass range of 70 − 1,050 m/z. 2.3.5 Data Preprocessing and Annotation Upon completion of MS detection, the raw LC/MS data were preprocessed using Progenesis QI software. A three-dimensional data matrix in CSV format was exported, containing details such as sample identifiers, metabolite names, and mass spectral response intensities. Internal standard peaks and known false positive peaks, including noise, column bleed, and derivatized reagent peaks, were removed from the data matrix. The data were then further processed to eliminate redundancies and pooled based on peak intensities. Metabolite identification was conducted using databases such as HMDB ( http://www.hmdb.ca/ ), Metlin ( https://metlin.scripps.edu/ ), and the Majorbio Database. The resulting data were uploaded to the Majorbio cloud platform ( https://cloud.majorbio.com ) for comprehensive analysis. Metabolic features detected in at least 80% of any sample set were retained after filtration. For samples where metabolite levels fell below the lower limit of quantitation, minimum metabolite values were imputed. Each metabolic feature was normalized by sum to mitigate errors introduced during sample preparation and due to instrument variability. The response intensity of sample MS peaks was normalized using the sum normalization method, producing a standardized data matrix. Variables with a relative standard deviation (RSD) exceeding 30% in QC samples were excluded. A log10 transformation was then applied to the final data matrix for further analysis. 2.3.6 Differential Metabolite Analysis Following data preprocessing, variance analysis was conducted on the matrix file. Principal component analysis (PCA) and orthogonal partial least squares discriminant analysis (OPLS-DA) were performed using the R package "ropls" (Version 1.6.2). A 7-cycle cross-validation was employed to assess the model's stability and reliability. Additionally, Student's t-test and fold difference analysis were conducted to identify significant differences between groups. Significantly different metabolites were identified based on the variable importance in projection (VIP) obtained from the OPLS-DA model, alongside the p-value from the Student's t-test. Metabolites with a VIP value greater than 1 and a p-value less than 0.05 were deemed statistically significant and selected for further analysis. Differential metabolites between the groups were summarized and mapped to their corresponding biochemical pathways using metabolic enrichment and pathway analysis, based on database searches such as KEGG ( http://www.genome.jp/kegg/ ). The identified metabolites were categorized according to the pathways they are involved in or the functions they perform. Enrichment analysis was conducted to determine whether a group of metabolites was overrepresented in a particular functional node. This analysis extended the annotation from individual metabolites to groups of metabolites. The statistical significance of enriched pathways was determined using Fisher's exact test, implemented with the "scipy.stats" Python package ( https://docs.scipy.org/doc/scipy/ ). 2.4 Water Quality Sample Collection To monitor key seawater parameters, an AQUAlogger 310TY turbidity meter, OS310 CTD, and YSI water quality analyzer were deployed to capture data on turbidity, temperature (WT), salinity (SAL), dissolved oxygen (DO), depth (WD), and pH. Seawater samples were collected from the bottom near the reef base in each sampling area using water sampling bottles. These samples were meticulously filtered through 0.45-µm glass fiber membranes and acidified with dilute hydrochloric acid. Subsequently, 100 mL of the filtrate was stored in reagent bottles. The concentrations of dissolved inorganic nutrients, including NH 4 + , NO 3 - , NO 2 - , SiO 3 2- , and PO 4 3- , were quantified using a SEAL AA3 automatic nutrient analyzer. Additionally, these samples were analyzed for dissolved inorganic carbon (DIC), partial pressure of carbon dioxide ( p CO 2 ), and aragonite saturation (Ω arag ) using the advanced CO 2 system software by Pierrot et al. ( 2006 ). 2.5 Data Analysis Physiological data were expressed as mean ± S.D., with the exception of metabolic group data. Statistical analyses were performed using Origin 2024b. Initially, the data were assessed to determine if they met the prerequisites for hypothesis testing, which included checking for normality using the Shapiro-Wilk test and testing for homogeneity of variance using Levene's test. For data that satisfied these conditions, an independent-sample t-test was conducted. The significance level was set at P < 0.05. 3. Results 3.1 Water Quality Monitoring In May 2023, environmental factors were meticulously measured and analyzed in both the natural and restoration areas of northern Wuzhizhou Island, Hainan ( Table S1 ). A significant event during this period was a large-scale algal bloom, which profoundly impacted various environmental indicators. This bloom led to marked increases in turbidity, dissolved inorganic nitrogen (DIN), and p CO 2 in the restoration area. By October 2023, and continuing into April 2024, these environmental factors began to stabilize, with most indicators in the restoration area approaching levels observed in the natural area, such as water temperature and salinity. However, notable differences remained; the natural area maintained higher levels of DO and Ω arag at both time points. These elevated levels of DO and lower turbidity in the natural area facilitated enhanced coral calcification and photosynthesis. In April 2024, the restoration area showed a significant increase in DO levels, which reached comparable heights to those of the natural area. This improvement supported enhanced coral respiration and metabolic activities, indicating a positive shift in the environmental conditions within the restoration site, fostering better conditions for coral health and growth. 3.2 Survival Rate and Physiological Indicators of Transplanted Corals The survival rate of transplanted G. fascicularis corals exhibited a notable decline from 100–64.58% within the first 6 months post-transplantation, with no further coral deaths observed between 6 and 12 months (Fig. 2 a). A critical factor contributing to coral mortality was the use of plastic ties for attachment. By the 12th month, corals that had shed their ties demonstrated a significantly higher survival rate (Fig. 2 b), while those with intact ties experienced nearly 100% mortality (Fig. 2 c). Surviving corals exhibited normal gonadal development, capable of producing mature sperm (Fig. 2 d) and eggs (Fig. 2 e). Physiological assessments indicated that the Fv/Fm of transplanted G. fascicularis was significantly lower than that of parent corals after 1 month. However, by the 6th month, the Fv/Fm values of transplanted corals recovered, showing no significant difference compared to parent corals (Fig. 2 f). As the transplantation period extended, both zooxanthellae density and coral biomass showed steady accumulation, reaching peak levels in both transplanted and parent corals by the 12th month (Fig. 2 g, i). Throughout the study, there were no significant differences in chlorophyll a and carbohydrate content among the groups (Fig. 2 h, j). By the 12th month, both transplanted and parent G. fascicularis reached sexual maturity, exhibiting significantly higher protein and lipid content compared to earlier time points (Fig. 2 k, l). 3.2 Comparative Analysis of Samples According to PLS-DA, the metabolic profiles of transplanted corals progressively aligned with those of parent corals over time, indicating a gradual acclimatization (Fig. 3 a, d, g). The distance between transplanted and parent coral samples decreased, demonstrating this convergence in metabolic patterns. Concurrently, the number of differential metabolites declined from 422 at 1 month to 343 at 6 months, and finally to 265 at 12 months (Fig. 3 b, e, h). This reduction in differential metabolites suggested that the physiological activities of transplanted corals were increasingly mirroring those of the parent corals. Specific metabolites showed significant changes during this period. Hydroxyethyl methacrylic acid was notably upregulated in the transplanted corals at 1 month (VIP = 5.75, P < 0.001) (Fig. 3 c). At 6 months, neamine was significantly upregulated (VIP = 6.48, P < 0.001) (Fig. 3 f). By 12 months, 15-keto-prostaglandin F2a exhibited significant upregulation in the transplanted corals (VIP = 5.00, P < 0.001) (Fig. 3 i). These findings highlighted the dynamic metabolic adjustments that transplanted corals underwent to adapt to their new environment, ultimately achieving physiological states similar to their parent counterparts. 3.3 KEGG Analysis KEGG pathway analysis revealed significant enrichment patterns among the upregulated and downregulated differential metabolites at various post-transplantation time points. At 1 month and 6 months post-transplantation, upregulated differential metabolites were predominantly enriched in the arachidonic acid metabolic pathway (Fig. 4 a, b). This indicated an early metabolic response to transplantation stress and adaptation processes. However, by 12 months post-transplantation, the upregulated differential metabolites showed a shift in pathway enrichment, no longer concentrating in the arachidonic acid metabolic pathway. Instead, they were significantly enriched in the oxidative phosphorylation pathway (Fig. 4 c). This transition suggested a shift in the metabolic focus of the transplanted corals towards enhanced energy production to support growth and reproductive processes. Conversely, at all three time points, 1 month, 6 months, and 12 months post-transplantation, the downregulated differential metabolites were consistently enriched in the glycerophospholipid metabolic pathway (Fig. 4 a, b, c). This consistent downregulation implied a potential redistribution of resources away from membrane lipid synthesis and maintenance, possibly reallocating metabolic efforts towards other critical functions necessary for adaptation and survival. These KEGG analysis results provided a comprehensive view of the dynamic metabolic adjustments occurring in transplanted corals, highlighting key pathways involved in their acclimation and long-term adaptation processes. 3.4 Analysis of Key Metabolic Pathways and Metabolite Content Glycerophospholipid metabolism was significantly downregulated across all three sampling periods. Within this pathway, only lysophosphatidylcholine (LPC) exhibited significant changes in expression patterns over time. The arachidonic acid pathway showed significant upregulation at 1 and 6 months post-transplantation, but no notable changes were observed at 12 months. Specifically, metabolites such as 20-Hydroxyeicosatetraenoic acid (20-HETE), 11,12-DiHETrE (11,12-DHET), and 8,9-DiHETrE (8,9-DHET) displayed significant expression changes at the 1- and 6-month marks. Lecithin emerged as a pivotal metabolite linking glycerophospholipid metabolism and the arachidonic acid pathway (Fig. 5 a). In glycerophospholipid metabolism, lecithin is catalyzed by the enzymes lecithin-cholesterol acyltransferase (LCAT) and secretory phospholipase A2 (sPLA2) to produce LPC. Subsequently, LPC is transformed into other metabolites, such as choline, phosphocholine, and sGPC, through various pathways. Within the arachidonic acid pathway, lecithin is converted into arachidonate under the catalytic action of sPLA2. Arachidonate is then metabolized to produce 20-HETE through the action of long-chain fatty acid omega-monooxygenase (CYP4A), phylloquinone omega-hydroxylase (CYP4F), and long-chain fatty acid omega-monooxygenase (CYP4U), while 11,12-DHET and 8,9-DHET are generated via more complex metabolic routes. Significant differences in LPC content were observed between transplanted and parent corals at 1, 6, and 12 months, with transplanted corals consistently showing downregulated expression (Fig. 5 b, c, d). For 20-HETE, significant differences were noted at 1 and 6 months, with upregulated expression in transplanted corals (Fig. 5 e, f). Similarly, 11,12-DHET and 8,9-DHET content differed significantly between transplanted and parent corals at 1 and 6 months, with both metabolites showing upregulated expression in the transplanted corals (Fig. 5 g, h, i, j). These findings underscored the intricate metabolic adjustments and adaptations occurring in transplanted corals, highlighting the critical roles of glycerophospholipid and arachidonic acid pathways in their physiological acclimation. 4. Discussion Most current coral transplantation efforts focus on asexual reproduction by transplanting coral fragments to restore damaged reefs (Patterson et al. 2016 ). This study further confirmed the asexual reproduction potential of G. fascicularis . By employing appropriate transplantation techniques and optimizing environmental conditions, G. fascicularis not only adapted swiftly to the new environment but also achieved a commendable survival rate. Beyond validating the efficacy of asexual reproduction, our study revealed that G. fascicularis could attain sexual maturity within a short transplantation period, specifically within 1 year. This rapid maturation suggested that future restoration efforts could enhance the genetic diversity and overall health of coral communities (Castillo et al. 2024 ). The ability of G. fascicularis to reproduce both asexually and sexually in a relatively short timeframe provided a dual approach to reef restoration, offering a robust strategy for improving resilience and adaptive capacity in coral ecosystems. These findings underscored the potential for integrated coral restoration strategies that leverage both asexual and sexual reproduction to rebuild and sustain diverse, resilient coral populations. This dual reproductive capability of G. fascicularis could play a pivotal role in future coral restoration projects, contributing to the long-term health and stability of reef ecosystems. 4.1 Survival Rate and Physiological Changes of Transplanted Corals Despite potential manual operation errors, we maintained transplanted G. fascicularis fragments at approximately 6 cm × 6 cm, while parent colonies exceeded 30 cm in diameter. Unpublished experiments in Wuzhizhou revealed that fragments from larger A. hyacinthus and Acropora microphthalma parent colonies could not reach sexual maturity within 2 years. Similarly, Zhao et al. ( 2024 ) have found that transplanted A. hyacinthus prioritizes growth over reproductive development. Ligson et al. (2021) have reported that Acropora verweyi requires 4 years to achieve sexual maturity. In contrast, G. fascicularis demonstrated remarkable adaptability by developing gonads while ensuring both survival and growth. Wei et al. ( 2023 ) have observed that large, wild-collected adult G. fascicularis shows normal gonadal development after 2 years in an artificial environment, consistent with our findings where transplanted G. fascicularis reached sexual maturity within 1 year. Environmental factors such as light, tides, and water flow in Wuzhizhou’s natural environment likely accelerated maturity compared to controlled indoor conditions. Studies on massive corals such as Favites colemani and Favites abdita (Guest et al. 2023 ) show that over 50% survive nursery acclimatization, with only a 10–14% decrease in survival 4 years post-transplantation. After 6 years, over 90% of the transplanted corals reach reproductive maturity. These findings suggest that restoring reproductive maturity in large coral populations within 10 years is feasible. The results of this study indicated that transplanting coral fragments from natural colonies could achieve similar reproductive maturity outcomes at a faster rate and lower cost compared to traditional methods. G. fascicularis , with its strong adaptability and rapid reproductive development, presented a viable candidate for effective coral reef restoration efforts. In this study, we observed that the survival rate of transplanted corals decreased rapidly within the first 6 months but remained stable between the 6th and 12th months. This trend is commonly seen in coral restoration efforts and suggests that once transplanted corals acclimate to the restoration environment, their physiological state can remain relatively stable (Ligson et al. 2021). Initially, environmental conditions such as a large-scale algal bloom in May 2023 increased turbidity, adversely affecting light penetration and photosynthesis (Palomar et al. 2009 ). High concentrations of DIN and phosphate (PO 4 3- ) promoted algal growth, further degrading water quality (Zhao et al. 2013 ). Elevated p CO 2 led to water acidification, impacting coral calcification and reducing survival rates (Nakamura et al. 2018 ). These stresses resulted in a decrease in Fv/Fm and zooxanthellae density, contributing to lower survival rates (Wall et al. 2018 ). From the 6th to 12th months, the stabilization of environmental conditions supported consistent coral survival. Favorable changes in temperature and salinity, comparable to those in the natural area, facilitated coral growth (Beck et al. 2022 ). Improved levels of DO and reduced turbidity in April 2024 restored coral photosynthetic capacity (Carlson et al. 2022 ). Enhanced Ω arag levels supported calcification processes (Martinez et al. 2019 ). As a result, the transplanted corals exhibited significantly higher protein and lipid content compared to natural corals after 12 months (Kaposi et al. 2023 ; Keister et al. 2023 ; Jung et al. 2021 ). Achieving sexual maturity, a crucial indicator of transplantation success, showed that transplanted corals could complete their life cycle and reproduce. Coral eggs and sperm, rich in lipids required for energy and cell structure, benefited from the improved conditions (Padilla-Gamiño et al. 2011 ). In the restoration area, the trapezoidal reef provided a relatively stable and suitable growth environment, optimizing light conditions, which likely contributed to the rapid growth and development of transplanted corals (Gomez-Campo et al. 2024 ). G. fascicularis demonstrated strong adaptability, making it an ideal candidate for coral restoration due to its ability to acclimate within 6 months. Our field investigation revealed that traditional binding methods might significantly contribute to the mortality of transplanted corals. Although plastic ties are inexpensive and easy to use (Tortolero-Langarica et al. 2019 ), they inflicted severe physical damage on G. fascicularis . These ties covered the corals' large calices, obstructing their respiration and feeding, which could ultimately lead to death. While some G. fascicularis managed to recover from this trauma, the overall recovery rate remained low. The energy expended on trauma recovery slows growth and increases survival pressure (Kaufman et al. 2021 ). In contrast, corals that detached from the ties due to growth breaking the ties, plastic degradation, or ocean currents exhibited higher survival rates and successfully attached to the restoration devices. These corals matured, producing sperm or eggs, which underscored the necessity of improving current binding methods to minimize physical harm and enhance survival rates. This finding highlighted the need for alternative attachment techniques that do not impede the natural physiological processes of the corals. 4.2 Synergistic Effects of Arachidonic Acid Metabolism and Glycerophospholipid Metabolism in Short-term (1 month) and Mid-term (6 months) Adaptation of Transplanted Corals In this study, we focused on the significant metabolic changes in arachidonic acid and glycerophospholipid pathways during the coral transplantation process. Transplanted G. fascicularis underwent a series of metabolic adjustments to adapt to the new environmental conditions. At 1 and 6 months post-transplantation, the arachidonic acid metabolic pathway was upregulated, while the glycerophospholipid metabolic pathway was significantly downregulated. These two pathways are biochemically interlinked, and alterations in one can induce corresponding changes in the other (Hanna and Hafez 2018 ). Glycerophospholipids are essential components of cell membranes and are hydrolyzed by PLA2 to produce arachidonic acid and lysophospholipids (Sikorskaya 2023 ). This process not only involves membrane phospholipid metabolism and remodeling but also impacts numerous signaling pathways. Our findings revealed a significant increase in arachidonic acid levels in G. fascicularis samples 1 month post-transplantation, with elevated levels persisting at 6 months. This finding underscored the crucial role of the arachidonic acid metabolic pathway in coral adaptation. Arachidonic acid metabolism facilitates stress response, inflammation regulation, antioxidation, and cell protection (Safuan et al. 2021 ). During the initial transplantation stage, it generates bioactive molecules like prostaglandins, which aid in coral tissue repair and defense against pathogens (Agalias et al. 2020). Additionally, increased levels of 20-HETE were observed at 1 and 6 months post-transplantation. 20-HETE modulates the severity and duration of inflammatory responses, supporting coral tissue survival and health (Lock et al. 2020). Moreover, the metabolites 11,12-DHET and 8,9-DHET, products of arachidonic acid metabolism, act as antioxidants, playing vital roles in responding to environmental and oxidative stress (Chhonker et al. 2018 ). These metabolites enhance antioxidant capacity by activating NF-κB and Nrf2 signaling pathways, thereby increasing the expression of cellular antioxidant genes (Zhang et al. 2022 ). This interplay between arachidonic acid and glycerophospholipid metabolism is critical for the short-term and mid-term adaptation of transplanted corals, facilitating their survival and acclimation in new environments. On the other hand, the downregulation of glycerophospholipid metabolism plays a crucial role in cell membrane remodeling and the adjustment of energy metabolism in transplanted corals. Experimental results indicate that during the initial phase of transplantation, the synthesis and degradation of glycerophospholipids, such as phosphatidylcholine and phosphatidylethanolamine, proceed rapidly to address cell membrane damage and facilitate reconstruction (Imbs 2013 ). LPC is a significant intermediate in the glycerophospholipid metabolic pathway, possessing various physiological functions (Stien et al. 2020 ). One month post-transplantation, corals begin to acclimate to the new environmental conditions, and the initial intense stress response starts to subside. As cellular oxidative stress levels decrease, the demand for LPC as an antioxidant molecule also diminishes (Tang et al. 2019 ). Consequently, LPC expression levels begin to downregulate. Cell membrane remodeling, which is critical during the early stages of transplantation, involves a substantial amount of LPC for repair and restructuring (Smith et al. 2009 ). Once the cell membrane structure stabilizes after the initial month, the need for LPC decreases, leading to its downregulation. This reduction in LPC can also enhance the flexibility of the cell membrane, thereby maintaining its structural stability (Fuller and Rand 2001 ). To achieve efficient metabolism in the new environment, cells adjust and optimize metabolic pathways to minimize unnecessary energy expenditure (Matthews et al. 2020 ). The generation and metabolism of LPC are energy-consuming processes. As cells gradually adapt to the new environment and regain stability, reducing LPC production conserves energy, redirecting resources to other vital physiological functions (Sousa et al. 2020 ). This adjustment helps restore metabolic balance as the cells acclimate to their new surroundings. The upregulation of arachidonic acid metabolism and the downregulation of glycerophospholipid metabolism in transplanted coral cells resulted from their synergistic response to environmental stress and optimization of resource allocation. In the early and mid-stages of transplantation, the arachidonic acid metabolic pathway offered robust antioxidant and inflammation-regulating capabilities, aiding cells in coping with environmental changes and repairing damage (Safuan et al. 2021 ). Concurrently, the downregulation of glycerophospholipid metabolism reflected a resource-conservation strategy after cell membrane stabilization. This adjustment optimized the overall metabolic state by reducing the energy expenditure associated with phospholipid synthesis and degradation (Sousa et al. 2020 ). These metabolic adjustments are critical for corals to achieve long-term stable survival in their new environment. Future research can delve deeper into the responses and interactions of these metabolic pathways under varying environmental stresses, aiming to uncover more profound mechanisms. Such insights will provide valuable theoretical support and practical guidance for coral conservation and restoration strategies. 4.3 Changes in Metabolic Patterns of Transplanted Corals 12 Months After Transplantation Natural corals, due to their long-term adaptation to stable environments, have evolved efficient energy management and utilization mechanisms (Hein et al. 2020 ). This efficiency allows them to meet the energy demands of sexual maturity without significant metabolic adjustments (Randall et al. 2020 ). In contrast, transplanted corals must undergo metabolic changes during their initial reproductive phase to meet the high energy demands of sexual maturity. By 12 months post-transplantation, the transplanted G. fascicularis appeared to have fully adapted to their new environment and reached sexual maturity. During this period, corals experience significant metabolic changes to prioritize reproductive activities (Guest et al. 2014 ). This involves reallocating resources from other physiological functions, such as growth and repair, to reproductive processes (Edwards and Clark 1999 ). This resource reallocation necessitates adjustments in cellular metabolism to ensure reproductive activities receive prioritized energy supply (Chan et al. 2019 ). Experimental data indicate that glycerophospholipid metabolism plays a crucial role in energy storage and allocation in transplanted corals (Boulotte et al. 2023 ). After 12 months post-transplantation, significant adjustments in glycerophospholipid metabolism were observed, optimizing metabolic pathways to reduce unnecessary energy consumption and allocate more energy to reproductive activities and growth (Wu et al. 2022 ). These metabolic adjustments enable transplanted corals to effectively utilize energy to support their sexual maturity and reproductive activities (Haydon et al. 2021 ). Twelve months post-transplantation, no significant changes were observed in the arachidonic acid metabolic pathway, indicating that the transplanted corals reached a stable state and no longer required additional energy investment in this pathway. The performance of the transplanted corals in the arachidonic acid metabolic pathway was consistent with that of natural corals at this stage. The study results showed that as corals gradually stabilized and reached sexual maturity, the demand for LPC decreased, consistent with the optimization of the metabolic pathways they are involved in, ensuring efficient energy utilization and normal cellular function (Sun et al. 2023 ). Additionally, the oxidative phosphorylation pathway is significantly upregulated in transplanted corals, primarily to meet the high energy demands of sexual maturity and reproductive activities (Yang et al. 2024 ). During sexual maturity, corals require a substantial amount of ATP to support gamete formation, release, and fertilization, all of which are energy-intensive processes (Briggs et al. 2024 ). Experimental results indicated that in corals 12 months post-transplantation, the activity of the oxidative phosphorylation pathway was enhanced, leading to a significant increase in ATP production. This increased activity supported the division and development of reproductive cells, antioxidant defense, signal transduction, and hormone regulation, all of which require additional energy (Zhang et al. 2021 ). The upregulation of the oxidative phosphorylation pathway not only boosts ATP production but also enhances the expression of antioxidant enzymes, helping to neutralize reactive oxygen species (ROS) generated during reproduction, thereby protecting the health of reproductive cells (Braun 2020 ). Key metabolites in the oxidative phosphorylation pathway, such as riboflavin and ubiquinone-1, are significantly upregulated in transplanted corals. Riboflavin, a precursor to flavin mononucleotide and flavin adenine dinucleotide, serves as a coenzyme in the electron transport chain (Udhayabanu et al. 2017 ). Ubiquinone-1, a crucial electron carrier, enhances the oxidative phosphorylation process, thereby increasing ATP production to meet high energy demands (Tang et al. 2022 ). Both riboflavin and ubiquinone-1 possess antioxidant properties, which help enhance antioxidant defense mechanisms, reduce oxidative stress damage to cells, and protect cell health (Pobłocka-Olech et al. 2019 ; Moller et al. 1996 ). By adapting their metabolism, transplanted corals can sustain optimal physiological functions, thereby ensuring their health and survival in their new environment. Declarations Conflict of Interest The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Ethics approval This study is original research. The Research Ethics Committee has confirmed that no ethical approval is required. Funding This work were financially supported by Major Science and Technology plan of Hainan Province (ZDYF2023SHFZ173), the National Natural Science Foundation of China (42161144006 or 3511/21 and 42076108), the Innovative Talent Foundation of Hainan Province (KJRC2023C39). Author contributions He Zhao conceived the ideas and designed methodology; Hongmin Wang, Jingzhao Ke and Junling Zhang collected the data; Yushan Li, Xiangbo Liu and Wentao Zhu analysed the data; Aimin Wang and Xiubao Li led the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication. Acknowledgments The authors were grateful for constructive suggestions and technical support from the instructors and all the laboratory members. Data availability Data will be made available on request. References Agalias A, Mihopoulos N, Tsoukatou M, Marinos L, Vagias C, Harvala C, Roussis V (2000) New Prostaglandins from the Chemically Defended Soft Coral Plexaura nina. 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Remote Sens 5(1):415–431. https://doi.org/10.3390/rs5010415 Supplementary Files 20240724Supplementarymaterials.docx Cite Share Download PDF Status: Published Journal Publication published 05 Jun, 2025 Read the published version in Marine Biology → Version 1 posted Editorial decision: Revise and Resubmit 04 Feb, 2025 Reviewers agreed at journal 02 Oct, 2024 Reviewers invited by journal 03 Sep, 2024 Editor assigned by journal 25 Jul, 2024 First submitted to journal 23 Jul, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4792475","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":349179954,"identity":"c6d0a9e4-cd79-492a-8ca8-00768cee266c","order_by":0,"name":"He Zhao","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"He","middleName":"","lastName":"Zhao","suffix":""},{"id":349179955,"identity":"bf6d5525-1a00-4d17-b995-3b63418587d6","order_by":1,"name":"Hongmin Wang","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Hongmin","middleName":"","lastName":"Wang","suffix":""},{"id":349179956,"identity":"dd7ac67d-7f88-467f-84f3-c3f33800a80b","order_by":2,"name":"Jingzhao Ke","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Jingzhao","middleName":"","lastName":"Ke","suffix":""},{"id":349179957,"identity":"4f0b40c4-f0ee-4632-bb89-54f6b5b265c8","order_by":3,"name":"Junling Zhang","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Junling","middleName":"","lastName":"Zhang","suffix":""},{"id":349179958,"identity":"c51eefaf-efe3-4f44-b3d0-1d3a060ec955","order_by":4,"name":"Yushan Li","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Yushan","middleName":"","lastName":"Li","suffix":""},{"id":349179959,"identity":"dfc8d787-a427-48bd-9715-186aae7463af","order_by":5,"name":"Xiangbo Liu","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Xiangbo","middleName":"","lastName":"Liu","suffix":""},{"id":349179960,"identity":"2810e289-3a13-49d8-8265-ba7a516fd1c0","order_by":6,"name":"Wentao Zhu","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Wentao","middleName":"","lastName":"Zhu","suffix":""},{"id":349179961,"identity":"7a921196-34e0-44fc-992c-f313d48625e7","order_by":7,"name":"Aimin Wang","email":"","orcid":"","institution":"Hainan University","correspondingAuthor":false,"prefix":"","firstName":"Aimin","middleName":"","lastName":"Wang","suffix":""},{"id":349179962,"identity":"c61be3cb-6fd3-4e12-9f50-77b83dc2a6b5","order_by":8,"name":"Xiubao Li","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA90lEQVRIiWNgGAWjYDACZgY2IGnBwy/BwHAAKmZAjBYJHskZRGthgGhhMLiBEMGvxeA4j9mDHxUSMsa3mx8eLqi5k9jA3rxNgqHmDk4tks085oY9ZyR4zO4cMzg849izxAaeY2USDMee4dTCz8xjJsHbBtRyI8HgMA/b4cQGiRwzCcaGw7g9AtQi+ReoxXhG+ofDPP+AWuTf4NcCskUaZIuBRI7BYd42kC08+LVINrOVG8sA/SJxI6fgMG/fYeM2nrRii4RjuLUYnD+87eGbCht7/hnpmz/zfDss289+eOONDzW4tWDxHYhIIEHDKBgFo2AUjAJMAADsNU3qt3XCmgAAAABJRU5ErkJggg==","orcid":"","institution":"Hainan University","correspondingAuthor":true,"prefix":"","firstName":"Xiubao","middleName":"","lastName":"Li","suffix":""}],"badges":[],"createdAt":"2024-07-24 05:16:54","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4792475/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4792475/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00227-025-04659-8","type":"published","date":"2025-06-05T15:57:15+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":65934842,"identity":"bbd90d9a-1934-4870-8d6d-68937437a527","added_by":"auto","created_at":"2024-10-04 14:52:44","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1503431,"visible":true,"origin":"","legend":"\u003cp\u003eExperimental site and trapezoidal reef design diagram. (a: Location of the natural area and restoration area in Wuzhizhou; b: Design diagram and actual photo of the trapezoidal reef)\u003c/p\u003e","description":"","filename":"OnlineFIg.1.png","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/70fce358cdffffdf6e0e6fdc.png"},{"id":65934843,"identity":"f452a84b-1191-448c-ae34-e1748c8953e8","added_by":"auto","created_at":"2024-10-04 14:52:44","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":18806742,"visible":true,"origin":"","legend":"\u003cp\u003eSurvival rate and physiological indicators changes of transplanted \u003cem\u003eG. fascicularis. \u003c/em\u003e(a: Changes in survival rate of transplanted corals; b: Higher survival rate of corals with fallen ties after 12 months; c: Nearly complete mortality of corals with intact ties after 12 months; d: Male \u003cem\u003eG. fascicularis\u003c/em\u003e reaching sexual maturity after 12 months of transplantation; e: Female \u003cem\u003eG. fascicularis\u003c/em\u003e reaching sexual maturity after 12 months of transplantation; f: Fv/Fm; g: Zooxanthellae density; h: Chlorophyll a content; i: Biomass; j: Total carbohydrate content; k: Protein content; i: Lipid content. * indicates P\u0026lt;0.05; ** indicates P\u0026lt;0.01; *** indicates P\u0026lt;0.001)\u003c/p\u003e","description":"","filename":"OnlineFig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/7ca983df588006366692d281.png"},{"id":65934838,"identity":"405b4bab-a91a-46d7-9bb5-47379a714ad3","added_by":"auto","created_at":"2024-10-04 14:52:44","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":53672,"visible":true,"origin":"","legend":"\u003cp\u003ePLS-DA between samples and volcano maps of different periods. (a: PLS-DA of transplanted and parent corals 1 month after transplantation; b: Number of differential metabolites between transplanted and parent corals 1 month after transplantation; c: Heatmap and VIP values of differential metabolite expression between transplanted and parent corals 1 month after transplantation; d: PLS-DA of transplanted and parent corals 6 months after transplantation; e: Number of differential metabolites between transplanted and parent corals 6 months after transplantation; f: Heatmap and VIP values of differential metabolite expression between transplanted and parent corals 6 months after transplantation; g: PLS-DA of transplanted and parent corals 12 months after transplantation; h: Number of differential metabolites between transplanted and parent corals 12 months after transplantation; i: Heatmap and VIP values of differential metabolite expression between transplanted and parent corals 12 months after transplantation)\u003c/p\u003e","description":"","filename":"Onlinefloatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/6685c8150a93aec30077ff2f.png"},{"id":65936707,"identity":"091138a4-520f-4648-9f63-7557b4db86ff","added_by":"auto","created_at":"2024-10-04 15:08:44","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":421865,"visible":true,"origin":"","legend":"\u003cp\u003eKEGG enrichment pathway. (a: Enriched pathways of differential metabolites 1 month after transplantation; b: Enriched pathways of differential metabolites 6 months after transplantation; Figure C: Enriched pathways of differential metabolites 12 months after transplantation)\u003c/p\u003e","description":"","filename":"OnlineFig.4.png","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/7e7f36cbdf45aae21728b3ca.png"},{"id":65936060,"identity":"3f9751e5-07fc-49d0-bee3-38e753f7bdda","added_by":"auto","created_at":"2024-10-04 15:00:44","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":5190763,"visible":true,"origin":"","legend":"\u003cp\u003eKey metabolic pathways and metabolite content. (a: Arachidonic acid metabolic pathway (red) and glycerophospholipid metabolic pathway (blue); b: Comparison of LPC content between transplanted and parent corals at 1 month; c: Comparison of LPC content between transplanted and parent corals at 6 months; d: Comparison of LPC content between transplanted and parent corals at 12 months; e: Comparison of 20-HETE content between transplanted and parent corals at 1 month; f: Comparison of 20-HETE content between transplanted and parent corals at 12 months; g: Comparison of 11,12-DHET content between transplanted and parent corals at 1 month; h: Comparison of 11,12-DHET content between transplanted and parent corals at 6 months; i: Comparison of 8,9-DHET content between transplanted and parent corals at 1 month; j: Comparison of 8,9-DHET content between transplanted and parent corals at 6 months. * indicates P\u0026lt;0.05; ** indicates P\u0026lt;0.01; *** indicates P\u0026lt;0.001)\u003c/p\u003e","description":"","filename":"OnlineFIg.5.png","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/6ba1ca9fd03467c5536fb715.png"},{"id":84242536,"identity":"b6a71c7d-682d-435f-9e19-79321a1fd006","added_by":"auto","created_at":"2025-06-09 16:09:10","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":7040912,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/a58d471e-1472-4dba-9f22-4acb13db572b.pdf"},{"id":65936058,"identity":"04e8bbd0-d010-42d9-9f33-66b812384f8c","added_by":"auto","created_at":"2024-10-04 15:00:44","extension":"docx","order_by":9,"title":"","display":"","copyAsset":false,"role":"supplement","size":17299,"visible":true,"origin":"","legend":"","description":"","filename":"20240724Supplementarymaterials.docx","url":"https://assets-eu.researchsquare.com/files/rs-4792475/v1/a47fdc83e8b899544511a29b.docx"}],"financialInterests":"","formattedTitle":"Analyzing Adaptation Mechanisms in Artificial Transplantation of Galaxea fascicularis","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eCoral reefs are among Earth's most diverse and ecologically crucial ecosystems (Castro-Sanguino et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), providing essential habitats, coastal protection, fisheries, and tourism benefits (Massei et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). They face severe threats from climate change, pollution, overfishing, and other disturbances (Niz et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), leading to widespread coral bleaching and mortality (Andradi-Brown et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Coral transplantation is a promising conservation strategy, involving the relocation of healthy coral fragments or larvae to degraded reefs to restore ecological functions (Ferse et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Lang et al., 2024). Success depends on environmental conditions and coral species (Williams et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Post-transplantation, corals undergo significant physiological and metabolic changes affecting survival and adaptation (Uribe-Casta\u0026ntilde;eda et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Understanding these mechanisms is essential for improving transplantation techniques and restoration outcomes.\u003c/p\u003e \u003cp\u003eMetabolomics is an advanced scientific approach that examines the physiological state and response mechanisms of organisms through a comprehensive analysis of their small molecular metabolites (Hu et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This technique is instrumental in unveiling the adaptive mechanisms of corals to environmental fluctuations. For instance, Roach et al. (2001) have demonstrated that metabolomic disparities in naturally bleached corals remain consistent over time, identifying betaine lipids as key biomarkers of coral bleaching. Similarly, Zhao et al. (\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) have explored the metabolomic heterogeneity of Acropora hyacinthus under both transplantation and natural conditions, revealing that differential metabolites can drive phenotypic variations, directing more energy towards growth rather than developmental processes. By investigating the metabolomic variations among corals, we can pinpoint individuals with superior adaptability and restoration potential, thereby refining coral transplantation methodologies and bolstering the restoration and preservation of coral reef ecosystems (Schmidt-Roach et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCurrent research predominantly focuses on the transplantation of branching corals, with relatively limited attention given to massive corals. Bostr\u0026ouml;m-Einarsson et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) have noted that coral reef restoration projects have been implemented in at least 56 countries, utilizing 229 species from 72 coral genera, encompassing approximately 25% of reef-building coral species. However, these projects primarily target fast-growing branching coral species, which, despite their rapid growth, are highly sensitive to environmental disturbances. In contrast, the more resilient but slower-growing massive corals are frequently overlooked (Guest et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The slow growth rate of massive corals often results in restoration outcomes that are visually unimpressive and fail to meet immediate expectations, leading to their neglect (Marshall and Baird \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Hein et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Additionally, the unique structural characteristics of massive corals pose significant challenges during transplantation, further contributing to the lack of research focus on these species (Edwards and Clark \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). Nevertheless, massive corals play an indispensable role in coral reef ecosystems. Their robust and durable structures provide critical physical support to the reefs, creating habitats and shelter for diverse marine organisms, thereby enhancing biodiversity (Glassom et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInvestigating the metabolomic changes in massive corals during transplantation and assessing their long-term performance post-transplantation is essential for evaluating transplantation success, adaptive responses, and long-term survival. In the northern part of Wuzhizhou Island, Hainan, we conducted an experiment involving the transplantation of massive corals. We selected \u003cem\u003eGalaxea fascicularis\u003c/em\u003e, the dominant coral species of Wuzhizhou Island, and transplanted it onto our custom-designed restoration device, the \"Trapezoidal Reef\". Samples were collected at three key intervals, 1 month, 6 months, and 12 months post-transplantation, to analyze the physiological characteristics and metabolomic differences between transplanted and parent \u003cem\u003eG. fascicularis\u003c/em\u003e. This study aimed to elucidate the adaptive changes occurring during the transplantation process, providing fundamental scientific data for future transplantation efforts of \u003cem\u003eG. fascicularis\u003c/em\u003e and contributing to the broader field of coral reef restoration.\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Experimental Site and Sample Collection\u003c/h2\u003e \u003cp\u003eIn April 2023, we established a coral reef restoration site in northern Wuzhizhou by installing 12 trapezoidal reef devices (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). Each device featured eight designated transplantation points for massive corals (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Divers meticulously collected and tagged fragments of \u003cem\u003eG. fascicularis\u003c/em\u003e from the natural reef area. Using hammers and chisels, small coral fragments were carefully extracted and transplanted onto the trapezoidal reefs within the restoration area, resulting in a total of 96 transplanted fragments of \u003cem\u003eG. fascicularis\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eTo assess the transplantation outcomes, we collected samples at 1 month (May 2023), 6 months (October 2023), and 12 months (April 2024) post-transplantation. At each time point, six tagged parent corals and six transplanted corals were retrieved. The collected samples were divided into two portions: one part was immediately preserved in liquid nitrogen for subsequent UPLC-MS/MS analysis, while the other part was used for measuring physiological indicators.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Measurement of Coral Physiological Indicators\u003c/h2\u003e \u003cp\u003eUpon collection, coral samples were immediately transported to the laboratory for analysis. Throughout the experiment, the maximum quantum efficiency of coral (Fv/Fm) was measured using the MINI-PAM-II chlorophyll fluorometer (Walz, Germany). To assess zooxanthellae density, coral surfaces were washed with a Waterpik dental cleaner (Water-pik, WP70-EC, USA). A specific volume of the wash solution (10 mL) was measured and counted using a hemocytometer, with the process repeated eight times to obtain an average value. This method follows the protocol of Li et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eChlorophyll a was extracted using an Addison reagent kit and quantified with a spectrophotometer, as per Fitt et al. (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). To determine coral tissue content, samples were dried at 60℃ for 24 hours to obtain dry weight and subsequently ashed at 400℃ in a muffle furnace for 4 hours to measure ash weight. The difference in mass pre- and post-ashing represents the coral tissue content. Coral surface area was measured using the aluminum foil method (Stimson \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1997\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCarbohydrate content was determined according to Dubois et al. (1956). Ground coral samples were extracted with phenol and concentrated sulfuric acid, and their absorbance was measured at A490 using a spectrophotometer. Absorbance values were converted to concentration using a glucose standard curve. Protein extraction followed the method of Smith et al. (\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Ground coral samples were processed with a BCA reagent kit, and their absorbance was measured at A562. Concentrations were calculated using a protein standard curve.\u003c/p\u003e \u003cp\u003eLipid extraction adhered to the protocol by Rodrigues and Grottoli (\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Approximately 1 g of wet coral sample was extracted with CM solution (methanol/chloroform 1:2, v/v). About 20 mL of CM extract was added to 1 g of coral and incubated in the dark for 24 hours. The organic phase was then evaporated and dried in an oxygen-free environment to determine lipid content. All data were standardized to the coral surface area.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Metabolome Analysis\u003c/h2\u003e \u003cdiv id=\"Sec6\" class=\"Section3\"\u003e \u003ch2\u003e2.3.1 Metabolite Extraction\u003c/h2\u003e \u003cp\u003eA solid sample of 10 mg was accurately weighed, and metabolites were extracted using 400 \u0026micro;L of methanol (4:1, v/v) solution containing 0.02 mg/mL L-2-chlorophenylalanine as an internal standard. The mixture was allowed to settle at -10℃ and then processed with a high-throughput tissue crusher Wonbio-96c at 50 Hz for 6 minutes, followed by ultrasound at 40 kHz for 30 minutes at 5℃. Samples were then placed at -20℃ for 30 minutes to precipitate proteins. After centrifugation at 13,000 g at 4℃ for 15 minutes, the supernatant was carefully transferred to sample vials for LC-MS/MS analysis.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section3\"\u003e \u003ch2\u003e2.3.2 Quality Control (QC) Sample\u003c/h2\u003e \u003cp\u003eAs part of the system conditioning and QC process, a pooled QC sample was prepared by mixing equal volumes from all samples. The QC samples were processed and analyzed in the same manner as the analytical samples. This pooled QC sample served as a representative of the entire sample set and was injected at regular intervals (every six samples) to ensure and monitor the stability and reliability of the analysis.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section3\"\u003e \u003ch2\u003e2.3.3 UHPLC-MS/MS Analysis\u003c/h2\u003e \u003cp\u003eThe UHPLC-MS analysis was conducted using the UHPLC-Q Exactive HF-X system from Thermo Fisher Scientific. Chromatographic separation was achieved using an HSS T3 column (100 mm \u0026times; 2.1 mm i.d., 1.8 \u0026micro;m), with 2 \u0026micro;L of each sample injected for MS detection. The mobile phases consisted of 0.1% formic acid in water (95:5, v/v) (solvent A) and 0.1% formic acid in acetonitrile: isopropanol (47.5:47.5:5, v/v) (solvent B). The solvent gradient was programmed as follows: from 0 to 3.5 min, 0% B to 24.5% B (0.4 mL/min); from 3.5 to 5 min, 24.5% B to 65% B (0.4 mL/min); from 5 to 5.5 min, 65% B to 100% B (0.4 mL/min); from 5.5 to 7.4 min, 100% B (0.4 mL/min to 0.6 mL/min); from 7.4 to 7.6 min, 100% B to 51.5% B (0.6 mL/min); from 7.6 to 7.8 min, 51.5% B to 0% B (0.6 mL/min to 0.5 mL/min); from 7.8 to 9 min, 0% B (0.5 mL/min to 0.4 mL/min); and from 9 to 10 min, 0% B (0.4 mL/min) to equilibrate the system. The sample injection volume was 2 \u0026micro;L, with a flow rate set at 0.4 mL/min. The column temperature was maintained at 40℃, and all samples were stored at 4℃ during the analysis period.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section3\"\u003e \u003ch2\u003e2.3.4 MS Conditions:\u003c/h2\u003e \u003cp\u003eThe MS data were collected using a Thermo UHPLC-Q Exactive HF-X Mass Spectrometer equipped with an electrospray ionization (ESI) source, operating in both positive and negative ion modes. The optimal conditions for the mass spectrometer were set as follows: heater temperature at 425℃, capillary temperature at 325℃, sheath gas flow rate at 50 arb, aux gas flow rate at 13 arb, and ion-spray voltage floating (ISVF) at -3,500 V for negative mode and 3,500 V for positive mode. The normalized collision energy was set at 20-40-60 V rolling for MS/MS. The full MS resolution was configured at 60,000, while the MS/MS resolution was set at 7,500. Data acquisition was performed in Data Dependent Acquisition (DDA) mode, with detection conducted over a mass range of 70\u0026thinsp;\u0026minus;\u0026thinsp;1,050 m/z.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section3\"\u003e \u003ch2\u003e2.3.5 Data Preprocessing and Annotation\u003c/h2\u003e \u003cp\u003eUpon completion of MS detection, the raw LC/MS data were preprocessed using Progenesis QI software. A three-dimensional data matrix in CSV format was exported, containing details such as sample identifiers, metabolite names, and mass spectral response intensities. Internal standard peaks and known false positive peaks, including noise, column bleed, and derivatized reagent peaks, were removed from the data matrix. The data were then further processed to eliminate redundancies and pooled based on peak intensities. Metabolite identification was conducted using databases such as HMDB (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.hmdb.ca/\u003c/span\u003e\u003cspan address=\"http://www.hmdb.ca/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), Metlin (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://metlin.scripps.edu/\u003c/span\u003e\u003cspan address=\"https://metlin.scripps.edu/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), and the Majorbio Database. The resulting data were uploaded to the Majorbio cloud platform (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://cloud.majorbio.com\u003c/span\u003e\u003cspan address=\"https://cloud.majorbio.com\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) for comprehensive analysis.\u003c/p\u003e \u003cp\u003eMetabolic features detected in at least 80% of any sample set were retained after filtration. For samples where metabolite levels fell below the lower limit of quantitation, minimum metabolite values were imputed. Each metabolic feature was normalized by sum to mitigate errors introduced during sample preparation and due to instrument variability. The response intensity of sample MS peaks was normalized using the sum normalization method, producing a standardized data matrix. Variables with a relative standard deviation (RSD) exceeding 30% in QC samples were excluded. A log10 transformation was then applied to the final data matrix for further analysis.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section3\"\u003e \u003ch2\u003e2.3.6 Differential Metabolite Analysis\u003c/h2\u003e \u003cp\u003eFollowing data preprocessing, variance analysis was conducted on the matrix file. Principal component analysis (PCA) and orthogonal partial least squares discriminant analysis (OPLS-DA) were performed using the R package \"ropls\" (Version 1.6.2). A 7-cycle cross-validation was employed to assess the model's stability and reliability. Additionally, Student's t-test and fold difference analysis were conducted to identify significant differences between groups. Significantly different metabolites were identified based on the variable importance in projection (VIP) obtained from the OPLS-DA model, alongside the p-value from the Student's t-test. Metabolites with a VIP value greater than 1 and a p-value less than 0.05 were deemed statistically significant and selected for further analysis.\u003c/p\u003e \u003cp\u003eDifferential metabolites between the groups were summarized and mapped to their corresponding biochemical pathways using metabolic enrichment and pathway analysis, based on database searches such as KEGG (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.genome.jp/kegg/\u003c/span\u003e\u003cspan address=\"http://www.genome.jp/kegg/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). The identified metabolites were categorized according to the pathways they are involved in or the functions they perform. Enrichment analysis was conducted to determine whether a group of metabolites was overrepresented in a particular functional node. This analysis extended the annotation from individual metabolites to groups of metabolites. The statistical significance of enriched pathways was determined using Fisher's exact test, implemented with the \"scipy.stats\" Python package (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://docs.scipy.org/doc/scipy/\u003c/span\u003e\u003cspan address=\"https://docs.scipy.org/doc/scipy/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Water Quality Sample Collection\u003c/h2\u003e \u003cp\u003eTo monitor key seawater parameters, an AQUAlogger 310TY turbidity meter, OS310 CTD, and YSI water quality analyzer were deployed to capture data on turbidity, temperature (WT), salinity (SAL), dissolved oxygen (DO), depth (WD), and pH. Seawater samples were collected from the bottom near the reef base in each sampling area using water sampling bottles. These samples were meticulously filtered through 0.45-\u0026micro;m glass fiber membranes and acidified with dilute hydrochloric acid. Subsequently, 100 mL of the filtrate was stored in reagent bottles.\u003c/p\u003e \u003cp\u003eThe concentrations of dissolved inorganic nutrients, including NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e, NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e-\u003c/sup\u003e, NO\u003csub\u003e2\u003c/sub\u003e\u003csup\u003e-\u003c/sup\u003e, SiO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e2-\u003c/sup\u003e, and PO\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e3-\u003c/sup\u003e, were quantified using a SEAL AA3 automatic nutrient analyzer. Additionally, these samples were analyzed for dissolved inorganic carbon (DIC), partial pressure of carbon dioxide (\u003cem\u003ep\u003c/em\u003eCO\u003csub\u003e2\u003c/sub\u003e), and aragonite saturation (Ω\u003csub\u003earag\u003c/sub\u003e) using the advanced CO\u003csub\u003e2\u003c/sub\u003e system software by Pierrot et al. (\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e2.5 Data Analysis\u003c/h2\u003e \u003cp\u003ePhysiological data were expressed as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;S.D., with the exception of metabolic group data. Statistical analyses were performed using Origin 2024b. Initially, the data were assessed to determine if they met the prerequisites for hypothesis testing, which included checking for normality using the Shapiro-Wilk test and testing for homogeneity of variance using Levene's test. For data that satisfied these conditions, an independent-sample t-test was conducted. The significance level was set at P\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Water Quality Monitoring\u003c/h2\u003e \u003cp\u003eIn May 2023, environmental factors were meticulously measured and analyzed in both the natural and restoration areas of northern Wuzhizhou Island, Hainan (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). A significant event during this period was a large-scale algal bloom, which profoundly impacted various environmental indicators. This bloom led to marked increases in turbidity, dissolved inorganic nitrogen (DIN), and \u003cem\u003ep\u003c/em\u003eCO\u003csub\u003e2\u003c/sub\u003e in the restoration area.\u003c/p\u003e \u003cp\u003eBy October 2023, and continuing into April 2024, these environmental factors began to stabilize, with most indicators in the restoration area approaching levels observed in the natural area, such as water temperature and salinity. However, notable differences remained; the natural area maintained higher levels of DO and Ω\u003csub\u003earag\u003c/sub\u003e at both time points. These elevated levels of DO and lower turbidity in the natural area facilitated enhanced coral calcification and photosynthesis.\u003c/p\u003e \u003cp\u003eIn April 2024, the restoration area showed a significant increase in DO levels, which reached comparable heights to those of the natural area. This improvement supported enhanced coral respiration and metabolic activities, indicating a positive shift in the environmental conditions within the restoration site, fostering better conditions for coral health and growth.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Survival Rate and Physiological Indicators of Transplanted Corals\u003c/h2\u003e \u003cp\u003eThe survival rate of transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e corals exhibited a notable decline from 100\u0026ndash;64.58% within the first 6 months post-transplantation, with no further coral deaths observed between 6 and 12 months (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). A critical factor contributing to coral mortality was the use of plastic ties for attachment. By the 12th month, corals that had shed their ties demonstrated a significantly higher survival rate (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb), while those with intact ties experienced nearly 100% mortality (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSurviving corals exhibited normal gonadal development, capable of producing mature sperm (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed) and eggs (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ee). Physiological assessments indicated that the Fv/Fm of transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e was significantly lower than that of parent corals after 1 month. However, by the 6th month, the Fv/Fm values of transplanted corals recovered, showing no significant difference compared to parent corals (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ef).\u003c/p\u003e \u003cp\u003eAs the transplantation period extended, both zooxanthellae density and coral biomass showed steady accumulation, reaching peak levels in both transplanted and parent corals by the 12th month (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eg, i). Throughout the study, there were no significant differences in chlorophyll a and carbohydrate content among the groups (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eh, j). By the 12th month, both transplanted and parent \u003cem\u003eG. fascicularis\u003c/em\u003e reached sexual maturity, exhibiting significantly higher protein and lipid content compared to earlier time points (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ek, l).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Comparative Analysis of Samples\u003c/h2\u003e \u003cp\u003eAccording to PLS-DA, the metabolic profiles of transplanted corals progressively aligned with those of parent corals over time, indicating a gradual acclimatization (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, d, g). The distance between transplanted and parent coral samples decreased, demonstrating this convergence in metabolic patterns. Concurrently, the number of differential metabolites declined from 422 at 1 month to 343 at 6 months, and finally to 265 at 12 months (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb, e, h). This reduction in differential metabolites suggested that the physiological activities of transplanted corals were increasingly mirroring those of the parent corals.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSpecific metabolites showed significant changes during this period. Hydroxyethyl methacrylic acid was notably upregulated in the transplanted corals at 1 month (VIP\u0026thinsp;=\u0026thinsp;5.75, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec). At 6 months, neamine was significantly upregulated (VIP\u0026thinsp;=\u0026thinsp;6.48, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef). By 12 months, 15-keto-prostaglandin F2a exhibited significant upregulation in the transplanted corals (VIP\u0026thinsp;=\u0026thinsp;5.00, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ei). These findings highlighted the dynamic metabolic adjustments that transplanted corals underwent to adapt to their new environment, ultimately achieving physiological states similar to their parent counterparts.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003e3.3 KEGG Analysis\u003c/h2\u003e \u003cp\u003eKEGG pathway analysis revealed significant enrichment patterns among the upregulated and downregulated differential metabolites at various post-transplantation time points. At 1 month and 6 months post-transplantation, upregulated differential metabolites were predominantly enriched in the arachidonic acid metabolic pathway (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, b). This indicated an early metabolic response to transplantation stress and adaptation processes.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eHowever, by 12 months post-transplantation, the upregulated differential metabolites showed a shift in pathway enrichment, no longer concentrating in the arachidonic acid metabolic pathway. Instead, they were significantly enriched in the oxidative phosphorylation pathway (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec). This transition suggested a shift in the metabolic focus of the transplanted corals towards enhanced energy production to support growth and reproductive processes.\u003c/p\u003e \u003cp\u003eConversely, at all three time points, 1 month, 6 months, and 12 months post-transplantation, the downregulated differential metabolites were consistently enriched in the glycerophospholipid metabolic pathway (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, b, c). This consistent downregulation implied a potential redistribution of resources away from membrane lipid synthesis and maintenance, possibly reallocating metabolic efforts towards other critical functions necessary for adaptation and survival. These KEGG analysis results provided a comprehensive view of the dynamic metabolic adjustments occurring in transplanted corals, highlighting key pathways involved in their acclimation and long-term adaptation processes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Analysis of Key Metabolic Pathways and Metabolite Content\u003c/h2\u003e \u003cp\u003eGlycerophospholipid metabolism was significantly downregulated across all three sampling periods. Within this pathway, only lysophosphatidylcholine (LPC) exhibited significant changes in expression patterns over time. The arachidonic acid pathway showed significant upregulation at 1 and 6 months post-transplantation, but no notable changes were observed at 12 months. Specifically, metabolites such as 20-Hydroxyeicosatetraenoic acid (20-HETE), 11,12-DiHETrE (11,12-DHET), and 8,9-DiHETrE (8,9-DHET) displayed significant expression changes at the 1- and 6-month marks.\u003c/p\u003e \u003cp\u003eLecithin emerged as a pivotal metabolite linking glycerophospholipid metabolism and the arachidonic acid pathway (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea). In glycerophospholipid metabolism, lecithin is catalyzed by the enzymes lecithin-cholesterol acyltransferase (LCAT) and secretory phospholipase A2 (sPLA2) to produce LPC. Subsequently, LPC is transformed into other metabolites, such as choline, phosphocholine, and sGPC, through various pathways. Within the arachidonic acid pathway, lecithin is converted into arachidonate under the catalytic action of sPLA2. Arachidonate is then metabolized to produce 20-HETE through the action of long-chain fatty acid omega-monooxygenase (CYP4A), phylloquinone omega-hydroxylase (CYP4F), and long-chain fatty acid omega-monooxygenase (CYP4U), while 11,12-DHET and 8,9-DHET are generated via more complex metabolic routes.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSignificant differences in LPC content were observed between transplanted and parent corals at 1, 6, and 12 months, with transplanted corals consistently showing downregulated expression (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb, c, d). For 20-HETE, significant differences were noted at 1 and 6 months, with upregulated expression in transplanted corals (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ee, f). Similarly, 11,12-DHET and 8,9-DHET content differed significantly between transplanted and parent corals at 1 and 6 months, with both metabolites showing upregulated expression in the transplanted corals (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eg, h, i, j). These findings underscored the intricate metabolic adjustments and adaptations occurring in transplanted corals, highlighting the critical roles of glycerophospholipid and arachidonic acid pathways in their physiological acclimation.\u003c/p\u003e \u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eMost current coral transplantation efforts focus on asexual reproduction by transplanting coral fragments to restore damaged reefs (Patterson et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). This study further confirmed the asexual reproduction potential of \u003cem\u003eG. fascicularis\u003c/em\u003e. By employing appropriate transplantation techniques and optimizing environmental conditions, \u003cem\u003eG. fascicularis\u003c/em\u003e not only adapted swiftly to the new environment but also achieved a commendable survival rate.\u003c/p\u003e \u003cp\u003eBeyond validating the efficacy of asexual reproduction, our study revealed that \u003cem\u003eG. fascicularis\u003c/em\u003e could attain sexual maturity within a short transplantation period, specifically within 1 year. This rapid maturation suggested that future restoration efforts could enhance the genetic diversity and overall health of coral communities (Castillo et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The ability of \u003cem\u003eG. fascicularis\u003c/em\u003e to reproduce both asexually and sexually in a relatively short timeframe provided a dual approach to reef restoration, offering a robust strategy for improving resilience and adaptive capacity in coral ecosystems. These findings underscored the potential for integrated coral restoration strategies that leverage both asexual and sexual reproduction to rebuild and sustain diverse, resilient coral populations. This dual reproductive capability of \u003cem\u003eG. fascicularis\u003c/em\u003e could play a pivotal role in future coral restoration projects, contributing to the long-term health and stability of reef ecosystems.\u003c/p\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Survival Rate and Physiological Changes of Transplanted Corals\u003c/h2\u003e \u003cp\u003eDespite potential manual operation errors, we maintained transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e fragments at approximately 6 cm \u0026times; 6 cm, while parent colonies exceeded 30 cm in diameter. Unpublished experiments in Wuzhizhou revealed that fragments from larger \u003cem\u003eA. hyacinthus\u003c/em\u003e and \u003cem\u003eAcropora microphthalma\u003c/em\u003e parent colonies could not reach sexual maturity within 2 years. Similarly, Zhao et al. (\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) have found that transplanted \u003cem\u003eA. hyacinthus\u003c/em\u003e prioritizes growth over reproductive development. Ligson et al. (2021) have reported that \u003cem\u003eAcropora verweyi\u003c/em\u003e requires 4 years to achieve sexual maturity.\u003c/p\u003e \u003cp\u003eIn contrast, \u003cem\u003eG. fascicularis\u003c/em\u003e demonstrated remarkable adaptability by developing gonads while ensuring both survival and growth. Wei et al. (\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) have observed that large, wild-collected adult \u003cem\u003eG. fascicularis\u003c/em\u003e shows normal gonadal development after 2 years in an artificial environment, consistent with our findings where transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e reached sexual maturity within 1 year. Environmental factors such as light, tides, and water flow in Wuzhizhou\u0026rsquo;s natural environment likely accelerated maturity compared to controlled indoor conditions.\u003c/p\u003e \u003cp\u003eStudies on massive corals such as \u003cem\u003eFavites colemani\u003c/em\u003e and \u003cem\u003eFavites abdita\u003c/em\u003e (Guest et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) show that over 50% survive nursery acclimatization, with only a 10\u0026ndash;14% decrease in survival 4 years post-transplantation. After 6 years, over 90% of the transplanted corals reach reproductive maturity. These findings suggest that restoring reproductive maturity in large coral populations within 10 years is feasible.\u003c/p\u003e \u003cp\u003eThe results of this study indicated that transplanting coral fragments from natural colonies could achieve similar reproductive maturity outcomes at a faster rate and lower cost compared to traditional methods. \u003cem\u003eG. fascicularis\u003c/em\u003e, with its strong adaptability and rapid reproductive development, presented a viable candidate for effective coral reef restoration efforts.\u003c/p\u003e \u003cp\u003eIn this study, we observed that the survival rate of transplanted corals decreased rapidly within the first 6 months but remained stable between the 6th and 12th months. This trend is commonly seen in coral restoration efforts and suggests that once transplanted corals acclimate to the restoration environment, their physiological state can remain relatively stable (Ligson et al. 2021). Initially, environmental conditions such as a large-scale algal bloom in May 2023 increased turbidity, adversely affecting light penetration and photosynthesis (Palomar et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). High concentrations of DIN and phosphate (PO\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e3-\u003c/sup\u003e) promoted algal growth, further degrading water quality (Zhao et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Elevated \u003cem\u003ep\u003c/em\u003eCO\u003csub\u003e2\u003c/sub\u003e led to water acidification, impacting coral calcification and reducing survival rates (Nakamura et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). These stresses resulted in a decrease in Fv/Fm and zooxanthellae density, contributing to lower survival rates (Wall et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFrom the 6th to 12th months, the stabilization of environmental conditions supported consistent coral survival. Favorable changes in temperature and salinity, comparable to those in the natural area, facilitated coral growth (Beck et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Improved levels of DO and reduced turbidity in April 2024 restored coral photosynthetic capacity (Carlson et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Enhanced Ω\u003csub\u003earag\u003c/sub\u003e levels supported calcification processes (Martinez et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). As a result, the transplanted corals exhibited significantly higher protein and lipid content compared to natural corals after 12 months (Kaposi et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Keister et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Jung et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Achieving sexual maturity, a crucial indicator of transplantation success, showed that transplanted corals could complete their life cycle and reproduce. Coral eggs and sperm, rich in lipids required for energy and cell structure, benefited from the improved conditions (Padilla-Gami\u0026ntilde;o et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the restoration area, the trapezoidal reef provided a relatively stable and suitable growth environment, optimizing light conditions, which likely contributed to the rapid growth and development of transplanted corals (Gomez-Campo et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). \u003cem\u003eG. fascicularis\u003c/em\u003e demonstrated strong adaptability, making it an ideal candidate for coral restoration due to its ability to acclimate within 6 months.\u003c/p\u003e \u003cp\u003eOur field investigation revealed that traditional binding methods might significantly contribute to the mortality of transplanted corals. Although plastic ties are inexpensive and easy to use (Tortolero-Langarica et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), they inflicted severe physical damage on \u003cem\u003eG. fascicularis\u003c/em\u003e. These ties covered the corals' large calices, obstructing their respiration and feeding, which could ultimately lead to death. While some \u003cem\u003eG. fascicularis\u003c/em\u003e managed to recover from this trauma, the overall recovery rate remained low. The energy expended on trauma recovery slows growth and increases survival pressure (Kaufman et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn contrast, corals that detached from the ties due to growth breaking the ties, plastic degradation, or ocean currents exhibited higher survival rates and successfully attached to the restoration devices. These corals matured, producing sperm or eggs, which underscored the necessity of improving current binding methods to minimize physical harm and enhance survival rates. This finding highlighted the need for alternative attachment techniques that do not impede the natural physiological processes of the corals.\u003c/p\u003e \u003cp\u003e \u003cb\u003e4.2 Synergistic Effects of Arachidonic Acid Metabolism and Glycerophospholipid Metabolism in Short-term (1 month) and Mid-term (6 months) Adaptation of Transplanted Corals\u003c/b\u003e \u003c/p\u003e \u003cp\u003eIn this study, we focused on the significant metabolic changes in arachidonic acid and glycerophospholipid pathways during the coral transplantation process. Transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e underwent a series of metabolic adjustments to adapt to the new environmental conditions. At 1 and 6 months post-transplantation, the arachidonic acid metabolic pathway was upregulated, while the glycerophospholipid metabolic pathway was significantly downregulated. These two pathways are biochemically interlinked, and alterations in one can induce corresponding changes in the other (Hanna and Hafez \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Glycerophospholipids are essential components of cell membranes and are hydrolyzed by PLA2 to produce arachidonic acid and lysophospholipids (Sikorskaya \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This process not only involves membrane phospholipid metabolism and remodeling but also impacts numerous signaling pathways.\u003c/p\u003e \u003cp\u003eOur findings revealed a significant increase in arachidonic acid levels in \u003cem\u003eG. fascicularis\u003c/em\u003e samples 1 month post-transplantation, with elevated levels persisting at 6 months. This finding underscored the crucial role of the arachidonic acid metabolic pathway in coral adaptation. Arachidonic acid metabolism facilitates stress response, inflammation regulation, antioxidation, and cell protection (Safuan et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). During the initial transplantation stage, it generates bioactive molecules like prostaglandins, which aid in coral tissue repair and defense against pathogens (Agalias et al. 2020). Additionally, increased levels of 20-HETE were observed at 1 and 6 months post-transplantation. 20-HETE modulates the severity and duration of inflammatory responses, supporting coral tissue survival and health (Lock et al. 2020).\u003c/p\u003e \u003cp\u003eMoreover, the metabolites 11,12-DHET and 8,9-DHET, products of arachidonic acid metabolism, act as antioxidants, playing vital roles in responding to environmental and oxidative stress (Chhonker et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). These metabolites enhance antioxidant capacity by activating NF-κB and Nrf2 signaling pathways, thereby increasing the expression of cellular antioxidant genes (Zhang et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This interplay between arachidonic acid and glycerophospholipid metabolism is critical for the short-term and mid-term adaptation of transplanted corals, facilitating their survival and acclimation in new environments.\u003c/p\u003e \u003cp\u003eOn the other hand, the downregulation of glycerophospholipid metabolism plays a crucial role in cell membrane remodeling and the adjustment of energy metabolism in transplanted corals. Experimental results indicate that during the initial phase of transplantation, the synthesis and degradation of glycerophospholipids, such as phosphatidylcholine and phosphatidylethanolamine, proceed rapidly to address cell membrane damage and facilitate reconstruction (Imbs \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). LPC is a significant intermediate in the glycerophospholipid metabolic pathway, possessing various physiological functions (Stien et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne month post-transplantation, corals begin to acclimate to the new environmental conditions, and the initial intense stress response starts to subside. As cellular oxidative stress levels decrease, the demand for LPC as an antioxidant molecule also diminishes (Tang et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Consequently, LPC expression levels begin to downregulate. Cell membrane remodeling, which is critical during the early stages of transplantation, involves a substantial amount of LPC for repair and restructuring (Smith et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Once the cell membrane structure stabilizes after the initial month, the need for LPC decreases, leading to its downregulation. This reduction in LPC can also enhance the flexibility of the cell membrane, thereby maintaining its structural stability (Fuller and Rand \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo achieve efficient metabolism in the new environment, cells adjust and optimize metabolic pathways to minimize unnecessary energy expenditure (Matthews et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The generation and metabolism of LPC are energy-consuming processes. As cells gradually adapt to the new environment and regain stability, reducing LPC production conserves energy, redirecting resources to other vital physiological functions (Sousa et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This adjustment helps restore metabolic balance as the cells acclimate to their new surroundings.\u003c/p\u003e \u003cp\u003eThe upregulation of arachidonic acid metabolism and the downregulation of glycerophospholipid metabolism in transplanted coral cells resulted from their synergistic response to environmental stress and optimization of resource allocation. In the early and mid-stages of transplantation, the arachidonic acid metabolic pathway offered robust antioxidant and inflammation-regulating capabilities, aiding cells in coping with environmental changes and repairing damage (Safuan et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Concurrently, the downregulation of glycerophospholipid metabolism reflected a resource-conservation strategy after cell membrane stabilization. This adjustment optimized the overall metabolic state by reducing the energy expenditure associated with phospholipid synthesis and degradation (Sousa et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThese metabolic adjustments are critical for corals to achieve long-term stable survival in their new environment. Future research can delve deeper into the responses and interactions of these metabolic pathways under varying environmental stresses, aiming to uncover more profound mechanisms. Such insights will provide valuable theoretical support and practical guidance for coral conservation and restoration strategies.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003e4.3 Changes in Metabolic Patterns of Transplanted Corals 12 Months After Transplantation\u003c/h2\u003e \u003cp\u003eNatural corals, due to their long-term adaptation to stable environments, have evolved efficient energy management and utilization mechanisms (Hein et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This efficiency allows them to meet the energy demands of sexual maturity without significant metabolic adjustments (Randall et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In contrast, transplanted corals must undergo metabolic changes during their initial reproductive phase to meet the high energy demands of sexual maturity. By 12 months post-transplantation, the transplanted \u003cem\u003eG. fascicularis\u003c/em\u003e appeared to have fully adapted to their new environment and reached sexual maturity. During this period, corals experience significant metabolic changes to prioritize reproductive activities (Guest et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This involves reallocating resources from other physiological functions, such as growth and repair, to reproductive processes (Edwards and Clark \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). This resource reallocation necessitates adjustments in cellular metabolism to ensure reproductive activities receive prioritized energy supply (Chan et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eExperimental data indicate that glycerophospholipid metabolism plays a crucial role in energy storage and allocation in transplanted corals (Boulotte et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). After 12 months post-transplantation, significant adjustments in glycerophospholipid metabolism were observed, optimizing metabolic pathways to reduce unnecessary energy consumption and allocate more energy to reproductive activities and growth (Wu et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These metabolic adjustments enable transplanted corals to effectively utilize energy to support their sexual maturity and reproductive activities (Haydon et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTwelve months post-transplantation, no significant changes were observed in the arachidonic acid metabolic pathway, indicating that the transplanted corals reached a stable state and no longer required additional energy investment in this pathway. The performance of the transplanted corals in the arachidonic acid metabolic pathway was consistent with that of natural corals at this stage. The study results showed that as corals gradually stabilized and reached sexual maturity, the demand for LPC decreased, consistent with the optimization of the metabolic pathways they are involved in, ensuring efficient energy utilization and normal cellular function (Sun et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAdditionally, the oxidative phosphorylation pathway is significantly upregulated in transplanted corals, primarily to meet the high energy demands of sexual maturity and reproductive activities (Yang et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). During sexual maturity, corals require a substantial amount of ATP to support gamete formation, release, and fertilization, all of which are energy-intensive processes (Briggs et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Experimental results indicated that in corals 12 months post-transplantation, the activity of the oxidative phosphorylation pathway was enhanced, leading to a significant increase in ATP production. This increased activity supported the division and development of reproductive cells, antioxidant defense, signal transduction, and hormone regulation, all of which require additional energy (Zhang et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe upregulation of the oxidative phosphorylation pathway not only boosts ATP production but also enhances the expression of antioxidant enzymes, helping to neutralize reactive oxygen species (ROS) generated during reproduction, thereby protecting the health of reproductive cells (Braun \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Key metabolites in the oxidative phosphorylation pathway, such as riboflavin and ubiquinone-1, are significantly upregulated in transplanted corals. Riboflavin, a precursor to flavin mononucleotide and flavin adenine dinucleotide, serves as a coenzyme in the electron transport chain (Udhayabanu et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Ubiquinone-1, a crucial electron carrier, enhances the oxidative phosphorylation process, thereby increasing ATP production to meet high energy demands (Tang et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Both riboflavin and ubiquinone-1 possess antioxidant properties, which help enhance antioxidant defense mechanisms, reduce oxidative stress damage to cells, and protect cell health (Pobłocka-Olech et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Moller et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). By adapting their metabolism, transplanted corals can sustain optimal physiological functions, thereby ensuring their health and survival in their new environment.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of Interest\u003c/h2\u003e \u003cp\u003eThe authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.\u003c/p\u003e \u003ch2\u003eEthics approval\u003c/h2\u003e \u003cp\u003eThis study is original research. The Research Ethics Committee has confirmed that no ethical approval is required.\u003c/p\u003e \u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work were financially supported by Major Science and Technology plan of Hainan Province (ZDYF2023SHFZ173), the National Natural Science Foundation of China (42161144006 or 3511/21 and 42076108), the Innovative Talent Foundation of Hainan Province (KJRC2023C39).\u003c/p\u003e\u003ch2\u003eAuthor contributions\u003c/h2\u003e \u003cp\u003eHe Zhao conceived the ideas and designed methodology; Hongmin Wang, Jingzhao Ke and Junling Zhang collected the data; Yushan Li, Xiangbo Liu and Wentao Zhu analysed the data; Aimin Wang and Xiubao Li led the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication.\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eThe authors were grateful for constructive suggestions and technical support from the instructors and all the laboratory members.\u003c/p\u003e\u003ch2\u003eData availability\u003c/h2\u003e \u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAgalias A, Mihopoulos N, Tsoukatou M, Marinos L, Vagias C, Harvala C, Roussis V (2000) New Prostaglandins from the Chemically Defended Soft Coral Plexaura nina. 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Remote Sens 5(1):415\u0026ndash;431. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/rs5010415\u003c/span\u003e\u003cspan address=\"10.3390/rs5010415\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Galaxea fascicularis, Artificial Transplantation, Coral Restoration, Metabolomics, Adaptation Mechanism","lastPublishedDoi":"10.21203/rs.3.rs-4792475/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4792475/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eCoral reefs are among Earth's most biologically diverse and ecologically crucial ecosystems but face severe threats from climate change and human activities. Coral transplantation has become a key strategy for reef restoration. This study focused on transplanting \u003cem\u003eGalaxea fascicularis\u003c/em\u003e at northern Wuzhizhou Island, Hainan, assessing physiological characteristics and metabolomic differences between transplanted and parent corals at 1, 6, and 12 months post-transplantation. Findings revealed that transplanted coral survival rates declined rapidly during the first 6 months but then stabilized. An algal bloom in May 2023 increased turbidity, dissolved inorganic nitrogen (DIN), and partial pressure of \u003cem\u003ep\u003c/em\u003eCO\u003csub\u003e2\u003c/sub\u003e, negatively impacting coral photosynthesis and calcification and increasing physiological stress. From months 6 to 12, environmental conditions improved, with temperature and salinity aligning closely with natural conditions, dissolved oxygen levels recovering, turbidity decreasing significantly, and Ω\u003csub\u003earag\u003c/sub\u003e reaching moderate levels, facilitating stable coral growth and calcification. By 12 months, transplanted corals reached sexual maturity, with notable increases in protein and lipid content. Metabolomic analysis showed that during the short-term (1 month) and mid-term (6 months) post-transplantation periods, the arachidonic acid metabolic pathway was upregulated while the glycerophosphate metabolic pathway was downregulated, enabling corals to cope with environmental stress and resource redistribution. By 12 months, oxidative phosphorylation was upregulated to meet reproductive energy demands. Results demonstrate that \u003cem\u003eG. fascicularis\u003c/em\u003e can adapt well to restoration environments and achieve sexual maturity quickly, making it a suitable candidate for reef restoration.\u003c/p\u003e","manuscriptTitle":"Analyzing Adaptation Mechanisms in Artificial Transplantation of Galaxea fascicularis","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-10-04 14:52:38","doi":"10.21203/rs.3.rs-4792475/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revise and Resubmit","date":"2025-02-04T23:58:28+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2024-10-02T13:45:04+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-09-03T23:53:24+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-07-25T09:16:03+00:00","index":"","fulltext":""},{"type":"submitted","content":"Marine Biology","date":"2024-07-24T01:16:43+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"marine-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mabi","sideBox":"Learn more about [Marine Biology](https://www.springer.com/journal/227)","snPcode":"227","submissionUrl":"https://submission.nature.com/new-submission/227/3","title":"Marine Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"40a1d166-b1d3-4258-8b83-c109a646e77a","owner":[],"postedDate":"October 4th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-06-09T16:00:37+00:00","versionOfRecord":{"articleIdentity":"rs-4792475","link":"https://doi.org/10.1007/s00227-025-04659-8","journal":{"identity":"marine-biology","isVorOnly":false,"title":"Marine Biology"},"publishedOn":"2025-06-05 15:57:15","publishedOnDateReadable":"June 5th, 2025"},"versionCreatedAt":"2024-10-04 14:52:38","video":"","vorDoi":"10.1007/s00227-025-04659-8","vorDoiUrl":"https://doi.org/10.1007/s00227-025-04659-8","workflowStages":[]},"version":"v1","identity":"rs-4792475","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4792475","identity":"rs-4792475","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}