The ecological effect of large-scale coastal natural and cultivated seaweed litter decay processes: An overview and perspective.

Seaweeds are important components of marine ecosystems and can form a large biomass in a few months. The decomposition of seaweed litter provides energy and material for primary producers and consumers and is an important link between material circulation and energy flow in the ecosystem. However, during the growth process, part of the seaweed is deposited on the sediment surface in the form of litter. Under the joint action of the environment and organisms, elements enriched in seaweed can be released back into the environment in a short time, causing pollution problems. The cultivation yield of seaweed worldwide reached 34.7 million tons in 2019, but the litter produced during the growth and harvest process has become a vital bottleneck that restricts the further improvement of production and sustainable development of the seaweed cultivation industry. Seaweed outbreaks worldwide occur frequently, producing a mass of litter and resulting in environmental pollution on coasts and economic losses, which have negative effects on coastal ecosystems. The objective of this review is to discuss the decomposition process and ecological environmental effects of seaweed litter from the aspects of the research progress on seaweed litter; the impact of seaweed litter on the environment; and its interaction with organisms. Understanding the decomposition process and environmental impact of seaweed litter can provide theoretical support for coastal environmental protection, seaweed resource conservation and sustainable development of the seaweed cultivation industry worldwide. This review suggests that in the process of large-scale seaweed cultivation and seaweed outbreaks, ageing or falling litter should be cleared in a timely manner, mature seaweed should be harvested in stages, and dried seaweed produced after harvest and washed up on shore should be handled properly to ensure the benefits of environmental protection provided by seaweed growth and sustainable seaweed resource development.

Biomass decomposition and heavy metal release from seaweed litter, Gracilaria lemaneiformis, and secondary pollution evaluation.

The seaweed Gracilaria lemaneiformis can bioremediate heavy metals and improve the environmental quality of mariculture zones. However, the seaweed litter that is produced in the growth and harvest processes becomes one of the important bottlenecks and causes secondary pollution that restricts the development of sustainable seaweed cultivation. Seaweeds exist widely in the coastal areas of the world and are cultivated on a large scale in Asia, but their decomposition process is rarely studied. Experiments that compared decaying dry (dead) and fresh (falling and dying) Gracilaria were conducted to quantify the differences in decomposition rates and heavy metal release in different physiological states. The heavy metals in the seawater and sediment were investigated. The litterbag technique under controlled laboratory conditions was used. The results indicated that the decomposition rates (k) and decay times in 50% (t50%) and 95% (t95%) values varied between dry and fresh Gracilaria. Fresh Gracilaria exhibited a weight loss rate of 15%, and the dry weight loss was 44%. The variations in MAIs (accumulation index of metals) and MR (release rate of metals) between the dry and fresh Gracilaria litters differed significantly, which provides evidence that metals are released back into the environment from Gracilaria litters. The contacted sediments could accelerate the heavy metal release from Gracilaria. Based on our estimates obtained from a 45 d experiment, at least 27.5％ of Cd, 16％ of Cu, 60.1％ of Pb, 72.3％ of Zn, 49.4％ of Fe, 38.6％ of Mn, 68.1％ of Cr, and 67.5％ of Ni present in the fresh Gracilaria and 37.4％ of Cd, 46.2％ of Cu, 77.7％ of Pb, 53.7％ of Zn, 42.7％ of Fe, 67.2％ of Mn, 75.1％ of Cr, and 73.5％ of Ni present in the dried Gracilaria were released back into the water when the biomass was left to decay. This study simulates and underscores that Gracilaria has an strong effect on the heavy metal cycles in marine environments and offers a theoretical basis for the development of sustainable seaweed industries in mariculture zones.

Protection of Siganus oramin, rabbitfish, from heavy metal toxicity by the selenium-enriched seaweed Gracilaria lemaneiformis.

Seaweed is an inherently important entity in marine ecosystems. It is not only consumed by aquatic animals but also improves environmental quality in the mariculture. Seaweed is also part of the diet of human beings. The purpose of the present study was to evaluate the antagonism of selenium (Se)-enriched Gracilaria lemaneiformis against heavy metals, specifically, the potential of dietary Se-enriched Gracilaria to protect against heavy metal toxicity in rabbitfish (Siganus oramin). Growth rate, heavy metal (Se, Cd, Pb, Cu, Zn and Cr) concentrations, malondialdehyde (MDA), metallothionein (MT), and the activity of the antioxidants, glutathione peroxidase (GPX), catalase (CAT) and superoxide dismutase (SOD) were all assessed. The results showed that the total organic and inorganic Se concentration for the 250 mg L-1 Se-enriched Gracilaria was significantly higher than those of the 50 and 10 mg L-1 treatments after 3 days of enrichment. The mean total Se concentrations in Gracilaria were 42.5 µg g-1 in the 250 mg L-1 treatment, 13.5 µg g-1 in the 50 mg L-1 treatment and 2.5 µg g-1 in the 10 mg L-1 treatment, respectively. Organic Se accounts for 80-82% of total Se in Se-enriched Gracilaria. The Se concentration of rabbitfish fed Se-enriched Gracilaria was significantly higher than control. Furthermore, Se increased Cu and Zn absorption, and enhanced MT generation, and improved GPX, CAT, and SOD antioxidant activity, and decreased MDA concentrations and lipid peroxidation levels, all antagonistic to Cd, Pb and Cr. The effects of Se-enriched Gracilaria on waterborne Cd, Pb and Cr-induced toxicity occurred via both enzymatic and non-enzymatic antioxidative mechanisms in rabbitfish. Selenium had synergistic effects on Zn and Cu in rabbitfish. For the 50 mg L-1 Se-enriched Gracilaria treatment, the Se, Cu, Zn, and antagonistic Cd, Pb, Cr, and the antioxidant enzymes CAT, SOD, GPX activities, and MT concentrations in rabbitfish were higher than that with the 250 mg L-1 and 10 mg L-1 Se-enriched Gracilaria treatments. The 50 mg L-1 Se treatment of Gracilaria was deemed to be the optimum concentration to promote growth of rabbitfish. Therefore, the obtained results suggest Se-enriched Gracilaria can antagonize heavy metal toxicity, and is an advisable Se supplement to improve the edible safety of cultured animals.

Protection of dietary selenium-enriched seaweed Gracilaria lemaneiformis against cadmium toxicity to abalone Haliotis discus hannai.

Seaweed Gracilaria lemaneiformis is the main dietary source of the abalone mariculture industry in China. In this study, we examined the protection of selenium (Se)-enriched G. lemaneiformis against cadmium (Cd) toxicity in the abalone, Haliotis discus hannai, using various indices including metal concentration (Se and Cd), growth rate, GPx enzymatic antioxidants, and metallothionein (MT) concentration over a period of 28 days of exposure. The growth rates and Se contents increased significantly in abalones fed with Se-enriched G. lemaneiformis, while the toxicity of Cd was reduced. Seven to 12 days Cd exposure to the Se-enriched G. lemaneiformis not only affected GPx activity but the MT levels fluctuated irregularly. MT concentrations increased after 3 days exposure and then gradually decreased to the control level after Day 7. There were statistically significant positive correlations between MT levels, GPx activity and Se concentrations, and negative relationships between MT levels, GPx activity and Cd levels in abalones. These findings suggest that Se-enriched Gracilaria protects abalone against Cd toxicity. The possible mechanism is the induction of MT with a concomitant increased capacity of GPx enzymatic antioxidants.

Heavy Metal Contamination in the Cultivated Oyster Crassostrea rivularis and Associated Health Risks from a Typical Mariculture Zone in the South China Sea.

With the rapid development of mariculture in potentially contaminated regions in China and the world, food safety, is a growing concern. To evaluate heavy metals and their associated health risks in the cultivated oyster Crassostrea rivularis, the concentrations of heavy metals (Cd, Cr, Pb, Zn) in oysters and water/sediment were examined in a typical mariculture environment (Kaozhou Bay, South China Sea). Trends in the seasonal dynamics of heavy metals in oysters revealed a potential synergistic effects among the concentrations of Cd, Cr, and Zn; trends associated with Pb were less clear, although the ability of oysters to bioaccumulate and depurate Pb was excellent. Bioconcentration factors (BCF) indicated that C. rivularis has a strong ability to accumulate heavy metals, and the BCF was the highest for Zn (2.32 × 105), followed by Cd (6.84 × 104), Pb (2.77 × 104) and Cr (1.23 × 103) through the four seasons. Results showed that Cd concentrations in oysters could pose a risk to human health (HQ > 1). This study, therefore, suggests that there are potential human health risks due to heavy metal exposure through the consumption of C. rivularis from mariculture zones in South China Sea.

Selenium-Containing Polysaccharide-Protein Complex in Se-Enriched Ulva fasciata Induces Mitochondria-Mediated Apoptosis in A549 Human Lung Cancer Cells.

The role of selenium (Se) and Ulva fasciata as potent cancer chemopreventive and chemotherapeutic agents has been supported by epidemiological, preclinical, and clinical studies. In this study, Se-containing polysaccharide-protein complex (Se-PPC), a novel organoselenium compound, a Se-containing polysaccharide-protein complex in Se-enriched Ulva fasciata, is a potent anti-proliferative agent against human lung cancer A549 cells. Se-PPC markedly inhibited the growth of cancer cells via induction of apoptosis which was accompanied by the formation of apoptotic bodies, an increase in the population of apoptotic sub-G1 phase cells, upregulation of p53, and activation of caspase-3 in A549 cells. Further investigation on intracellular mechanisms indicated that cytochrome C was released from mitochondria into cytosol in A549 cells after Se-PPC treatment. Se-PPC induced depletion of mitochondrial membrane potential (ΔΨm) in A549 cells through regulating the expression of anti-apoptotic (Bcl-2, Bcl-XL) and pro-apoptotic (Bax, Bid) proteins, resulting in disruption of the activation of caspase-9. This is the first report to demonstrate the cytotoxic effect of Se-PPC on human cancer cells and to provide a possible mechanism for this activity. Thus, Se-PPC is a promising novel organoselenium compound with potential to treat human cancers.

The accumulation and release characteristics of heavy metals on the cultivation environment in Gracilaria litters during decay process.

Gracilaria bioremediates heavy metals (Cd, Cr, Pb, Ni, Cu, Zn, Fe, and Mn) and improves water quality in mariculture zones. However, Gracilaria litter produced during the growth and harvest process has become a critical bottleneck problem that limits the sustainable development of the Gracilaria cultivation industry. Experiments of decaying dried (dead) and frozen fresh (falling and dying) G. lemaneiformis and G. lichenosdies were carried out using the litterbag technique under laboratory-controlled and in situ conditions. The results showed that decay rates (k), decomposed time in 50 % (t50) and in 95 % (t95) varied between dried and frozen fresh Gracilaria and were different between G. lemaneiformis and G. lichenosdies. All Gracilaria samples showed an 80 %-90 % weight loss in 15-45 d. The variation in MAIs (accumulation index of metals) between the dried and frozen fresh Gracilaria litters differed significantly and provided evidence that metals could be imported or exported from litter to the environment. Based on our estimates from the 15-45 d experiment, the decay of Gracilaria can release and adsorb heavy metals. The enrichment of Fe, Pb, and Mn was more significant than the release, but the release of Cr, Zn, Cd, Pb, Cu, and Ni was more significant than the enrichment. Heavy metals in Gracilaria litters were accumulated and released simultaneously during decay. The present study simulated and underscores that Gracilaria cultivation intensely influences heavy metals recycled in marine environments It provides a theoretical basis for seaweed management for the sustainable development of the seaweed industry in the mariculture zone.

Bioaccumulation and release of heavy metals during growth and decomposition of cultivated Gracilaria lemaneiformis.

Seaweeds are important primary producers and bioremediation materials, but its litter produced during growth and harvest is one of the restrictions to the sustainable development of seaweed cultivation. In this study, we conducted field investigation and indoor experiments to analyze the bioaccumulation and release of metals in Gracilaria lemaneiformis during the growth and decaying. The investigation revealed the 3.5 × 105 t (wet weight) G. lemaneiformis from a 1500 ha cultivation area bioaccumulated 1925-2353 kg Zn, 233.5-251 kg Cu, 70.5-80.5 kg Pb and 25.5-47 kg Cd, indicating that G. lemaneiformis is a good metals remover. The growth and decaying period of G. lemaneiformis releases, absorbs or adsorbs metals. It has the function of a "heavy metal pool", simultaneously accumulate and release metals. G. lemaneiformis has a strong influence on heavy metals cycling in the seaweed cultivation ecosystem and provides a very good sample for biogeochemistry study for the globally seaweed sustainable development.

Potential bioremediation effects of seaweed Gracilaria lemaneiformis on heavy metals in coastal sediment from a typical mariculture zone.

Seaweeds are good bio-monitors of heavy metals pollution in coastal seawater. In the present study, the potential bioremediation effects of cultivated Gracilaria lemaneiformis on heavy metals in Nan'ao coastal sediment from a typical mariculture zone, South China were evaluated. Sediment samples were collected from five different zones (Gracilaria cultivation zone, G; Fish culture zone, F; Shellfish culture zone, S; Transition zone, T; Control zone, C) from December 2014 to July 2015. The concentrations of Cd, Pb, Cu, and Zn in the sediments were significantly different among the various types of mariculture areas. The concentrations varied widely: Cd (0.04-1.02) µg g-1; Cu (1.19-37.70) µg g-1; Pb (8.45-74.45) µg g-1; Zn (36.80-201.24) µg g-1. The lowest heavy metal concentrations in the sediment were occurred at Gracilaria cultivation zone, while higher concentrations occurred at control zones and fish culture zones. The pollution load index, principal components and cluster analysis showed that heavy metal concentrations were the highest at fish culture zone, while the concentrations were the lowest at Gracilaria cultivation zone, and Gracilaria cultivation affects the heavy metals in the sediments. Gracilaria had strong adsorption capacities for heavy metals from seawater, showing the highest heavy metal Bioconcentration Factors in May (higher seaweed biomass period). Consequently, the results suggested that Gracilaria cultivation influences the heavy metal concentrations in sediments from the typical coastal mariculture zone. Gracilaria cultivation has the potential to bioremediate heavy metals in the coastal sediments. Therefore, Gracilaria cultivation can add environmental advantages and ecological values to coastal mariculture zones.

The Beta-Diversity of Siganus fuscescens-Associated Microbial Communities From Different Habitats Increases With Body Weight.

Fish-associated microbial communities play important roles in host growth, health and disease in the symbiont ecosystem; however, their diversity patterns and underlying mechanisms in different body habitats remain poorly understood. Siganus fuscescens is one of the most important consumers of macroalgae and an excellent natural marine source of nutritional lipids for humans, and widely distributes in shallow coastal areas. Here we systematically studied the microbial communities of 108 wild S. fuscescens in four body habitats (i.e., skin, gill, stomach, and hindgut) and surrounding water. We found that the ß-diversity but not α-diversity of fish-associated microbial communities from each habitat significantly (p < 0.05) increased as body weight increased. Also, opportunistic pathogens and probiotics (e.g., Pseudomongs, Methylobacterium) appeared to be widely distributed in different body habitats, and many digestive bacteria (e.g., Clostridium) in the hindgut; the abundances of some core OTUs associated with digestive bacteria, "Anaerovorax" (OTU_6 and OTU_46724) and "Holdemania" (OTU_33295) in the hindgut increased as body weight increased. Additionally, the quantification of ecological processes indicated that heterogeneous selection was the major process (46-70%) governing the community assembly of fish microbiomes, whereas the undominated process (64%) was found to be more important for the water microbiome. The diversity pattern showed that ß-diversity (75%) of the metacommunity overweight the α-diversity (25%), confirming that the niche separation of microbial communities in different habitats and host selection were important to shape the fish-associated microbial community structure. This study enhances our mechanistic understanding of fish-associated microbial communities in different habitats, and has important implications for analyzing host-associated metacommunities.

Proteogenomic Analyses Revealed Favorable Metabolism Pattern Alterations in Rotifer Brachionus plicatilis Fed with Selenium-rich Chlorella.

Organoselenium have garnered attention because of their potential to be used as ingredients in new anti-aging and antioxidation medicines and food. Rotifers are frequently used as a model organism for aging research. In this study, we used Se-enriched Chlorella (Se- Chlorella), a novel organoselenium compound, to feed Brachionus plicatilis to establish a rotifer model with a prolonged lifespan. The results showed that the antioxidative effect in Se-enriched rotifer was associated with an increase in guaiacol peroxidase (GPX) and catalase (CAT). The authors then performed the first proteogenomic analysis of rotifers to understand their possible metabolic mechanisms. With the de novo assembly of RNA-Seq reads as the reference, we mapped the proteomic output generated by iTRAQ-based mass spectrometry. We found that the differentially expressed proteins were primarily involved in antireactive oxygen species (ROS) and antilipid peroxidation (LPO), selenocompound metabolism, glycolysis, and amino acid metabolisms. Furthermore, the ROS level of rotifers was diminished after Se- Chlorella feeding, indicating that Se- Chlorella could help rotifers to enhance their amino acid metabolism and shift the energy generating metabolism from tricarboxylic acid cycle to glycolysis, which leads to reduced ROS production. This is the first report to demonstrate the anti-aging effect of Se- Chlorella on rotifers and to provide a possible mechanism for this activity. Thus, Se- Chlorella is a promising novel organoselenium compound with the potential to prolong human lifespans.