Comparison of growth and nutrient uptake capacities of three dominant species of Qinhuangdao green tides

Since 2015, green tides have been blooming in offshore waters of Qinhuangdao, with serious impacts on the local ecological environment and tourism. Ulva australis, Bryopsis plumosa, and U. prolifera are the dominant species of Qinhuangdao green tides, following a sequential succession pattern. Ulva prolifera is the dominant species,with the highest biomass and the greatest influence on the local ecological environment. To study the reason of green tide dominant species succession and U. profilera became the dominant species with the largest biomass,we compared and analyzed the growth and nutrient uptake capacity of the three algae. The results showed that temperature significantly affects the growth of the three species. Within the temperature range of the experimental setup, the optimum temperature for the growth of U. australis, B. plumosa and U. profilera is10℃, 15℃, and 20–25℃, respectively. Combined with the temperature variation trend during green tide bloom development, we believe that temperature is the key environmental factor for the succession of the dominant species. Ulva prolifera has a higher growth rate than U. australis and B. plumosa under the same nitrate,ammonium, and phosphate levels. Significant differences in the maximum absorption rate(R_(max)) and R_(max)/Ks(the relationship between uptake rate and substrate concentration) values indicated that U. prolifera had an apparent competitive advantage over U. australis and B. plumosa regarding nutrient uptake. Therefore, the strong growth and nutrient uptake capacities of U. prolifera might be the main reason for becoming the dominant species with the largest biomass in Qinhuangdao green tides.

The contribution of attached Ulva prolifera on Pyropia aquaculture rafts to green tides in the Yellow Sea

Green tides caused by the unusual accumulation of high floating Ulva prolifera have occurred regularly in the Yellow Sea since 2007.The primary source of the Yellow Sea green tides is the attached algae on the Pyropia aquaculture rafts in the Subei Shoal.Ulva prolifera and Blidingia(Italic)sp.are the main species observed on Pyropia aquaculture rafts in the Subei Shoal.We found that U.prolifera has strong buoyancy and a rapid growth rate,which may explain why it is the dominant species of green tides that occur in the China's sea area of the Yellow Sea.The growth rate of floating U.prolifera was about 20%–31%d–1,which was much higher than Blidingia(Italic)sp.There were about 1.7×104 t of attached algae on the Pyropia aquaculture rafts in May 2012.We found that 39%of attached algae could float when the tide rose in the Subei Shoal,and U.prolifera accounted for 63%of the floating algae.Our analysis estimated that about 4000 t of attached U.prolifera floated into the surrounding waters of the Subei Shoal during the recycling period of aquaculture rafts.These results suggest that the initial floating biomass of large-scale green tides in the Yellow Sea is determined by the U.prolifera biomass attached to Pyropia aquaculture rafts,further impacting the scale of the green tide。

Morphology and Molecular Identification of Ulva Forming Green Tides in Qingdao,China

Green tides are caused by the proliferation of chlorophytes under suitable hydrographic conditions.These blooms lead to environmental degradation and negatively impact the waters and seagrass beds,as well as fishing and other recreational activities in the bay.A comprehensive ecological understanding of the bloom dynamics,including the origin and persistence,is needed to foster management decisions.The algae in the great majority of green tide blooms usually belong to two genera of Ulvophyceae,Ulva and Enteromorpha.Ulva has been observed more often in recent years.In China,green tides occurred for the first time in the middle area of the Yellow Sea in 2007,and a large-scale algae blooming broke out in the middle and southern areas of the Yellow Sea in late May 2008.We identified them as Ulva prolifera by comparative analysis of the rDNA internal transcribed spacer 1 (ITS1),5.8S and ITS2 sequences in combination with microscopic observation.Morphological differences were found between the free-floating algae and the attached thalli.Various reproduction patterns of the free-floating algae include sexual,asexual and vegetative propagations,which played important roles in the long-term green tide persistence in China.The ITS sequences of the blooming algae were identical to those of the samples from the Lianyungang sea area but were different from the attached samples from the Qingdao sea area.The results infer that the blooms are originated from other sea areas rather than from the local attached populations.

Genome-wide SNP markers provided insights into the reproductive strategy and genetic diversity of the green tide causative species Ulva prolifera in China

Ulva prolifera is the causative species of the annually occurring large-scale green tides in China since 2007.Its specific biological features on reproductivity strategies,as well as intra-species genetic diversity,are still largely unknown,especially at the genome level,despite their importance in understanding the formation and outbreak of massive green tides.In the present study,the restriction site-associated DNA genotyping approach(2b-RAD)was adopted to identify the genome-wide single-nucleotide polymorphisms(SNPs)of 54 individual thalli including samples collected from Subei Shoal in 2019 and Qingdao coast from 2019 to 2021.SNPs genotype results revealed that most of the thalli in 2019 and 2020 were haploid gametophytes,while only half of the thalli were gametophytes in 2021,indicating flexibility in the reproductive strategies for the formation of the green tides among different years and the dominance of asexual and vegetative reproductive mode for the floating period.Besides,population analysis was conducted,and it revealed a very low genetic diversity among samples from Subei Shoal and the Qingdao coast in the same year and a higher divergence among samples in different years.The results showed the efficiency of 2b-RAD in the exploration of SNPs in U.prolifera and provided the first genome-wide scale evidence for the origin of the large-scale green tides on the Qingdao coast.This study improved our understanding of the reproductive strategy and genetic diversity of the green tide causative species and will help further reveal the biological causes of the green tide in China.

Analysis on the causes of massive stranding of Yellow Sea green tide on Lianyungang and Rizhao coasts in 2022

In 2022,Yellow Sea green tide caused by Ulva prolifera reached a historic minimum and the coastal areas of Shandong Peninsula were less affected.However,the largest amount of seaweed biomass has been washed ashore on Lianyungang and Rizhao coasts since 2015.We studied the development pattern of Yellow Sea green tide in 2022,and analyzed the key environmental factors on the growth and drifting,then discussed the possible reasons that resulted in the massive stranding of green tide biomass in Lianyungang and Rizhao.Results show under the combined influence of the east to southeast winds and currents with shoreward anomalies,green tide drifted to the coastal waters between Shandong and Jiangsu provinces and the distribution areas located westward compared with previous years(2008–2021).Floating U.prolifera rafts from the coastal waters of Binhai and Sheyang drifted continuously into the coastal waters of Lianyungang and Rizhao,providing important supplements for Yellow Sea green tide.Because green tide in 2022 distributed close to the coastal waters,the abundant nutrients might support their continuous high growth rate.In addition,the amount of rainfall around Shandong Peninsula from late June to early July were significantly higher than in previous years,which might promote the development of green tide to some extent.

Eutrophication of Jiangsu Coastal Water and Its Role in the Formation of Green Tide

Since 2007,the large-scale green tide caused by Ulva prolifera(U.prolifera)have occurred as a recurrent phenomenon in the southern Yellow Sea of China.Field surveys and satellite remote sensing showed that the small scattered patches of green tide algae were first observed along the Porphyra agriculture area of the Subei Shoal in late April.In this study,we attempted to identify the role of eutrophication in the origin of the green tide in the Subei Shoal and its adjacent area.Subei Shoal and its adjacent area are characterized by rich nutrients,especially NO_(3)^(-)-N,NH_(4)^(+)-N,PO_(4)^(3-)-P,and other bioavailable components(such as urea-N and amino acids).In the spring of 2017,the average concentrations of NO_(3)^(-)-N were 19.01±11.01μmolL^(-1),accounting for 86.68%of the dis-solved inorganic nitrogen(DIN).In addition,the average concentration of NH4^(+)-N was 2.51±1.60μmolL^(-1).PO_(4)^(3-)-P had an average concentration of 0.14±0.13μmolL-1.The average concentrations of urea-N and total hydrolyzed amino acids(THAA)were 1.73±1.36μmolL^(-1)and 1.33±0.80μmolL^(-1),respectively.Rich nutritive substances play a key role in the rapid production of U.prolifera and make the Jiangsu coastal water an incubator for green tide.

Estimation and verification of green tide biomass based on UAV remote sensing

Since 2007,the Yellow Sea green tide has broken out every summer,causing great harm to the environment and society.Although satellite remote sensing(RS)has been used in biomass research,there are several shortcomings,such as mixed pixels,atmospheric interference,and difficult field validation.The biomass of green tide has been lacking a high-precision estimation method.In this study,high-resolution unmanned aerial vehicle(UAV)RS was used to quantitatively map the biomass of green tides.By utilizing experimental data from previous studies,a robust relationship was established to link biomass to the red-green-blue floating algae index(RGB-FAI).Then,the lab-based model for green tide biomass from visible images taken by the UAV camera was developed and validated by field measurements.Re sults show that the accurate and cost-effective method is able to estimate the green tide biomass and its changes in given local waters of the near and far seas.The study provided an effective complement to the traditional satellite RS,as well as high-precision quantitative techniques for decision-making in disaster management.

Source of propagules of the fouling green macroalgae in the Subei Shoal,China

Since 2007,large-scale green tides dominated by Ulva prolifera consecutively bloomed in the Yellow Sea and caused great economic losses.The fouling U.prolifera on the Pyropia yezoensis aquaculture rafts in the Subei Shoal was regarded as the major source of the floating biomass.However,it was still unclear about the seed source of fouling green macroalgae attached on the rafts.In this study,the field surveys and the indoor experiments were conducted to reveal the source of propagules of the fouling green macroalgae on the rafts and to study the anti-fouling material for P.yezoensis aquaculture rafts which could possibly be a feasible strategy to control the green tides in the Yellow Sea.The results showed that（1） micro-propagules of several green macroalgal species,including U.prolifera,U.linza,U.compressa,U.flexuosa,and Blidingia sp.coexisted in the waters and sediments in the Subei Shoal and their proportion remarkably changed over time;（2） the bamboo poles with peeling treatment could significantly reduce the amount of U.prolifera micro-propagules attached.This study confirmed that the micro-propagules distributed in the Subei Shoal area were the precursors of the green tides,and provided a feasible method to control the Yellow Sea large-scale green tides at the beginning.

Acute toxicity of live and decomposing green alga Ulva (Enteromorpha) prolifera to abalone Haliotis discus hannai

From 2007 to 2009, large-scale blooms of green algae (the so-called "green tides") occurred every summer in the Yellow Sea, China. In June 2008, huge amounts of floating green algae accumulated along the coast of Qingdao and led to mass mortality of cultured abalone and sea cucumber. However, the mechanism for the mass mortality of cultured animals remains undetermined. This study examined the toxic effects of Ulva (Enteromorpha) prolifera, the causative species of green tides in the Yellow Sea during the last three years. The acute toxicity of fresh culture medium and decomposing algal effluent of U. prolifera to the cultured abalone Haliotis discus hannai were tested. It was found that both fresh culture medium and decomposing algal effluent had toxic effects to abalone, and decomposing algal effluent was more toxic than fresh culture medium. The acute toxicity of decomposing algal effluent could be attributed to the ammonia and sulfide presented in the effluent, as well as the hypoxia caused by the decomposition process.

An automatic detection of green tide using multi-windows with their adaptive threshold from Landsat TM/ETM plus image

Since the atmospheric correction is a necessary preprocessing step of remote sensing image before detecting green tide, the introduced error directly affects the detection precision. Therefore, the detection method of green tide is presented from Landsat TM/ETM plus image which needs not the atmospheric correction. In order to achieve an automatic detection of green tide, a linear relationship（y =0.723 x＋0.504） between detection threshold y and subtraction x（x=λnir–λred） is found from the comparing Landsat TM/ETM plus image with the field surveys.Using this relationship, green tide patches can be detected automatically from Landsat TM/ETM plus image.Considering there is brightness difference between different regions in an image, the image will be divided into a plurality of windows（sub-images） with a same size firstly, and then each window will be detected using an adaptive detection threshold determined according to the discovered linear relationship. It is found that big errors will appear in some windows, such as those covered by clouds seriously. To solve this problem, the moving step k of windows is proposed to be less than the window width n. Using this mechanism, most pixels will be detected[n/k]×[n/k] times except the boundary pixels, then every pixel will be assigned the final class（green tide or sea water） according to majority rule voting strategy. It can be seen from the experiments, the proposed detection method using multi-windows and their adaptive thresholds can detect green tide from Landsat TM/ETM plus image automatically. Meanwhile, it avoids the reliance on the accurate atmospheric correction.

The green tide in Yingkou,China in summer 2021 was caused by a subtropical alga——Ulva meridionalis(Ulvophyceae,Chlorophyta)

The large-scale green tide caused by Ulva has occurred successively in the Yellow Sea since 2007,and new events of green tide also continued to appear in nearby sea areas,indicating an undergoing rapid development of occurrence patterns for harmful macroalgal blooms(HMBs)along coastal China.In August 2021,a green tide occurred for the first time in Bayuquan sea area of Yingkou city,Liaoning Province in the Bohai Sea.In this study,morphological and molecular approaches were used to identify the causative species as U.meridionalis,an alien subtropical alga previously found to dominate green tides in the South China Sea.According to the hydrological data of Bayuquan in summer 2021,combined with morphological and developmental observations for this alga,we hypothesized that the disturbance caused by the typhoon In-Fa might have detached the local U.meridionalis from substrates,especially for those thalli with poorly developed holdfasts,and the ensuing wave-free period with unusually high temperature,which fell in the reported optimum growth temperature for U.meridionalis,might have provided the favorable conditions for the final bloom of the floating seaweeds.This is the first report on the bloom of subtropical U.meridionalis in the north temperate sea zone,indicating that the ecological risk of causing green tides in the future by this rapidly spreading species deserves high attention.

Tempo-spatial distribution of Ulva spp.micro-propagules in the Yellow Sea during and after green tide in 2019

Large-scale green tides occur frequently in summer in the Yellow Sea in recent decade.Micropropagules,providing“seeds”for the seaweeds,play an important role in the formation of green tide.We studied the distribution pattern and genetic composition of the Ulva micro-propagules in the Yellow Sea during and after the green tide in 2019.In April,a great amount of Ulva micro-propagules was detected in the Subei Shoal,with the average density of 908 inds./L,being similar to that of previous studies.U.prolifera was found out at a low percentage(4%),and 25%of them were positive when tested by the specific marker for the floating ecotype.The percentage of U.prolifera was obviously lower than that in the same season from 2009 to 2016.Therefore,the quantity of U.prolifera micro-propagules sharply decreased.Significant variation in composition of the micro-propagules in Subei radial sand ridges was revealed in the background of the alternation of macroalgal tides in the Yellow Sea in the latest years.During the green tide in June and July,Ulva micro-propagules were detected from some nearshore sites where the floating seaweeds distributed,and U.prolifera was the dominate species.In September,when the green tide ended,no Ulva micro-propagule was detected from the open sea area of the Yellow Sea.It is indicated that the green algae micro-propagules were unlikely to exist in the open water of the Yellow Sea for a long time according to the large-scale surveys.The mechanism of preserving micro-propagules in Subei radial sand ridges,and the destination of U.prolifera micro-propagules in the open sea area of the Yellow Sea after green tide deserves further study.

Pyrolytic characteristics and kinetics of the marine green tide macroalgae, Enteromorpha prolifera

The marine macroalgae Enteromorpha prolifera was one of the main algal genera that occurred in the widespread green tides in Qingdao, China, during the summers of 2007, 2008 and 2010. It is thus a plentiful source of biomass and could be used as a biofuel. In this study, the pyrolytic characteristics and kinetics of E. prolifera were investigated using thermogravimetric analysis (TGA) method. Cornstalk and sawdust were used as comparisons. Pyrolytic characteristics were studied using TG-DTG (thermogravimetry-derivative thermogravimetry) curves. Three stages in the pyrolytic process were determined: dehydration, dramatic weight loss and slow weight loss. E. prolifera was pyrolyzed at a lower initial temperature than the two terrestrial biomass forms. The apparent activation energy values for the three types of biomass were calculated and the mechanism functions were determined using 16 different mechanism functions, frequently used in thermal kinetics analysis. Activation energy values varied with mechanism function and the range of activation energy values for E. prolifera, cornstalk, and sawdust were 25-50 kJ/mol, 60-90 kJ/mol and 120-155 kJ/mol, respectively. This indicates that E. prolifera has low thermal stability for pyrolysis and good combustion characteristics.

Antioxidant system responses in two co-occurring green-tide algae under stress conditions

Green tides have occurred every year from 2007 to 2014 in the Yellow Sea. Ulva prolifera （Mtiller） J. Agardh has been identified as the bloom-forming alga, co-occurring with U. intestinalis. We observed distinct strategies for both algal species during green tides. U.prolifera exhibited a high abundance initially and then decreased dramatically, while U. intestinalis persisted throughout. The antioxidant system responses of these two macroalgae were compared in the late phase of a green tide （in-situ） and after laboratory acclimation. Lipid peroxidation and antioxidant system responses differed significantly between the two. Malondialdehyde and hydrogen peroxide contents increased significantly in-situ in U. prolifera, but not in U. intestinalis. In U. prolifera, we observed a significant decrease in total antioxidant ability （T-AOC）, antioxidant enzymes （SOD and Apx）, and non-enzyme antioxidants （GSH and AsA） in-situ. U. intestinalis showed the same pattern of T-AOC and SOD, but its Gpx, Apx, and GSH responses did not differ significantly. The results suggest that U. prolifera was more susceptible than U. intestinalis to the harsh environmental changes during the late phase of a Yellow Sea green tide. The boom and bust strategy exhibited by U. prolifera and the persistence of U. intestinalis can be explained by differences in enzyme activity and antioxidant systems.

Variations in nano-and pico-eukaryotic phytoplankton assemblages in the Qinhuangdao green-tide area

Qinhuangdao coastal waters have been frequently hitting by nano-and pico-eukaryotic phytoplankton(NPEP)blooms and green tides(macroalgal blooms)in the recent decade.However,understanding about the impacts of environmental factors and the green tides on the NPEP assemblages in this area is limited.In this study,the composition of NPEP assemblages and their variations were analyzed via amplicon sequence variants(ASVs)assay based on amplicon high-throughput sequencing data with the 18S V4 region as a targeted gene in the Qinhuangdao green-tide area during the green tide.Consequently,average NPEP effective sequences and ASVs of 178000 and 200 were obtained from each sample,respectively.Although there were 25 classes,110 genera,and 97 species of NPEP were identified and annotated,the proportions of annotated ASVs at genus and species levels were only 44.7%and 17.8%,respectively.The NPEP communities had a seasonal succession from diatom-dominated to dinoflagellate-dominated.During the three investigations,Skeletonema,Karlodinium,and Gonyaulax were the most dominant genera in May,August,and September,respectively.Species diversity and the abundance of NPEP communities could be increased by a high content of dissolved organic nitrogen(DON)and dissolved organic phosphorus(DOP)but inhibited by low dissolved inorganic phosphorus content.The outbreak of green tides could alter the composition and content of nutrients and accelerate the succession of the NPEP communities from diatom-dominated to dinoflagellate-dominated under the background of a seasonal increase in seawater temperature.These results preliminarily revealed the impacts of the recurrent occurrences of green tides on the NPEP assemblages in the Qinhuangdao green-tide area exhibiting high DON content and dissolved inorganic nitrogen/phosphorus ratio.

Population differentiation in the dominant species(Ulva prolifera)of green tide in coastal waters of China

Since 2015,green tides with Ulva prolifera as the dominant species in the Qinhuangdao coastal waters have continued to occur.In this study,the relationship between green tides in Qinhuangdao and the Yellow Sea(setting sites in Rudong and Qingdao)was evaluated by genetic analyses of U.prolifera.Single nucleotide polymorphism(SNP)markers were used to analyze genetic diversity and genetic relationships among groups.Genetic differentiation was lower among floating U.prolifera populations in Rudong and Qingdao than in Qinhuangdao.The floating U.prolifera population had higher genetic diversity and polymorphism levels in Qingdao and Rudong than in Qinhuangdao.Physiological experiments showed that the growth rate and net buoyancy of floating U.prolifera were highest in Qinhuangdao and Qingdao,respectively,under the same environmental conditions(temperature and light).Overall,these findings showed that U.prolifera populations in the Qinhuangdao and Yellow Sea green tides(Rudong and Qingdao)differ significantly at the molecular and physiological levels.Therefore,the Qinhuangdao green tide is not correlated with the Yellow Sea green tide and has a different origin and development mode.This study provides insight into the mechanism underlying green tide blooms in coastal waters of China.

Research on Growth Characteristics of Green-Tide-Forming Green Algae under Stress Conditions

The cytological characteristics of major green-tide-forming green algae Ulva prolifera collected from Yellow Sea were studied through cutting segments, long time low temperature or dark treatments. After being dried in the shade and preserved at -20°C for 30 days, the U. prolifera was cultured at 4°C in sterilized seawater under 40 μmol photons m-2·s-1 light intensity for 120 days, results indicated that the plastid of U. prolifera continuously shrank with the extension of treatment, and most cells turned white and died, only a small amount of cells still contained a few of visible inclusions at the 120d of treatment. Then those samples were transferred to 20°C and 40 μmol photons m-2·s-1 condition for recovery cultivation, after about 10 days, some recovery cells were observed in the thallus, and those cells developed to young thallus gradually and released germ cells almost in the same time. After about 60 days of recovery cultivation, the newly-grown green thallus broke through the original dead thallus, and the germ cells also grew to new individual thallus. Before dark treatment, the U. prolifera cells were filled with plastid, contained visible starch grain and discernible cell outlines, while after 120 days of dark treatment, the plastid shrank and degraded together with the disappearance of cell inclusions, and the cell outlines also blurred, then those samples were transferred to optimal culture conditions at 20°C in 40 μmol photons m-2·s-1 light intensity, and 15 days later, newly-grown cells appeared on the almost dead thallus, these cells divided continuously and grew to young thallus, and those newly-grown thallus also generated active germ cells, which developed to new thallus that cytologically identical to the original thallus. Observation of chopped tissue of U. prolifera cultivated at 20°C, 40 μmol m-2·s-1 showed that the morphological upper part cells turned to germ cells first, those germ cells including gametophyte and sporophyte, which released later and grew to new individual thallus. These findings provided cytological evidences for how U. prolifera live through stress conditions such as low temperature, darkness, and also useful for understanding the mechanism of the occurrence of green tide.

The Overwintering Capability of Ulva prolifera Spores and Gametes in the Yellow Sea,China

Green tides caused by the bloom of Ulva prolifera have affected the Yellow Sea annually since 2007.To explore the intrinsic causes of successive blooms,field investigations and indoor simulation experiments were conducted using micro-propagules of U.prolifera.From January to February 2014,micro-propagules of various green algae,including U.prolifera,Ulva flexuosa,and Ulva linza,were prevalent in the coastal waters of Rudong,Lianyungang,and Qingdao.However,in March 2014,micro-propagules of U.prolifera were not detected in the coastal waters of Lianyungang and Qingdao.Neither the spores nor gametes of U.prolifera can germinate at temperatures below 5℃.At 2℃and 5℃,U.prolifera spores and gametes could survive for 1-3 months at a light intensity range of 0-30μmol m^(-2)s^(-1)and for less than 1 month if the light intensity exceeded 30μmol m^(-2)s^(-1).By integrating indoor experiments and field investigations,this study found that U.prolifera spores and gametes could overwinter in the Rudong coastal area,which is an important factor for the annual emergence of recurring green tides in the Yellow Sea.

Increased diversity and environmental threat of harmful algal blooms in the Southern Yellow Sea,China

Harmful algal blooms(HABs)in the Southern Yellow Sea(SYS)have shown a trend of increasing diversity and detrimental ef fects.While the Bohai Sea,East China Sea,and South China Sea have experienced a high incidence of HABs since the 1980s,the Yellow Sea provides a relatively healthy ecological environment in which fewer HABs have been documented before the 21s t century.Yet largescale blooms of the green macroalga Ulva prolifera(so-called“green tides”)have occurred annually since 2007 in the Yellow Sea.Six people were poisoned and one person died in Lianyungang in 2008 due to ingestion of algal toxins.Moreover,the Yellow Sea experienced co-occurrence of harmful red tides,green tides,and golden tides in 2017.This combination of events,rare worldwide,indicates the potential for further deterioration of the marine environment in the Yellow Sea,which may be related to climate change,aquaculture,and other human activities.Using the SYS as an example,we collected data of the frequency and scale of HABs over the years,as well as that of marine algal toxins,and analyzed the trend in the diversity of HABs in the SYS,to explore the causes and impacts of HABs,as well as the interrelationships among dif ferent types of HABs,including harmful red tides,green tides,and golden tides.We also attempted to improve our understanding of HAB evolution under the influence of global climate change and intensified human activities.

Effect of temperature,salinity and irradiance on growth and photosynthesis of Ulva prolifera

Intensive Pyropia aquaculture in the coast of southwestern Yellow Sea and its subsequent waste, including disposed Ulva prolifera, was speculated to be one of the major sources for the large-scale green tide proceeding in the Yellow Sea since 2007. It was, however, unclear how the detached U. prolifera responded and resumed growing after they detached from its original habitat. In this study, we investigated the growth and photosynthetic response of the detached U. prolifera to various temperature, salinity and irradiance in the laboratory. The photosynthetic rate of the detached U. prolifera was significantly higher at moderate temperature levels（14–27℃）and high salinity（26–32）, with optimum at 23℃ and 32. Both low（14℃） and highest temperature（40℃）, as well as low salinity（8） had adverse effects on the photosynthesis. Compared with the other Ulva species, U. prolifera showed higher saturated irradiance and no significant photoinhibition at high irradiance, indicating the great tolerance of U. prolifera to the high irradiance. The dense branch and complex structure of floating mats could help protect the thalli and reduce photoinhibition in field. Furthermore, temperature exerted a stronger influence on the growth rate of the detached U. prolifera compared to salinity. Overall, the high growth rate of this detached U. prolifera（10.6%–16.7% d^–1） at a wide range of temperature（5–32℃） and salinity（14–32） implied its blooming tendency with fluctuated salinity and temperature during floating. The environmental parameters in the southwestern Yellow Sea at the beginning of green tide were coincident with the optimal conditions for the detached U. prolifera.