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.     

The Ulva prolifera genome reveals the mechanism of green tides

The genome of green microalgae has rarely been reported.Ulva prolifera is a green microalga that has received much attention.Despite research articles about U.prolifera in recent years,we know very little about its genome.Therefore,the 87.9-Mb haploid genome(containing 10 311 protein-coding genes)of U.prolifera was studied,and the genome was compared with that of U.mutabilis,which is the only published Ulva species.Results showed that the two species are closely related.A phylogenetic tree was constructed among U.prolifera and other green algae available in GenBank,revealing the evolutionary status of U.prolifera in Chlorophyta.To understand why U.prolifer a could grow rapidly,we identified some genes related to growth,such as those involved in cell division,phosphorylation,and cell proliferation.In addition,genes related to stress resistance were found,which supports the notion that U.prolifera grows vigorously in nature.These results help to characterize green tides from a new perspective and reveal some important insight into the biology of U.prolifera.     

Seasonal variation and modal content of internal tides in the northern South China Sea

A nine-month mooring record was used to investigate seasonal variation and modal content of internal tides(ITs) on the continental slope in the northern South China Sea(SCS). Diurnal tides at this site show clear seasonal change with higher energy in winter than in spring and autumn, whereas semidiurnal tides show the opposite seasonal pattern. The consistency of ITs with barotropic tides within the Luzon Strait, which is the generation region of the ITs, implies that the seasonal variation of ITs depends on their astronomical forcing, even after extended propagation across the SCS basin. Diurnal tides also differ from semidiurnal tides in relation to modal content; they display signals of high modes while semidiurnal tides are dominated by low modes. Reflection of the diurnal tides on the continental slope serves as a reasonable explanation for their high modes. Both diurnal and semidiurnal tides are composed of a larger proportion of coherent components that have a regular 14-day spring-neap cycle. The coherent components are dominated by low modes and they show obvious seasonal variation, while the incoherent components are composed mainly of higher modes and they display intermittent characteristics.     

Coherent and incoherent internal tides in the southern South China Sea

Coherent and incoherent internal tides(CITs and ICITs) in the southern South China Sea were investigated from two sets of _18-month mooring current records. The CITs were mainly composed of diurnal Q _1, O _1, P _1 and K _1 and semidiurnal M_2. The observed diurnal internal tides(ITs) were more coherent than the semidiurnal constituents. Coherent diurnal variance accounted for approximately 58% of the diurnal motion, whereas semidiurnal tides contained a much smaller fraction(35%) of coherent motion. The ICITs mainly consisted of motion at non-tidal harmonic frequencies around the tidal frequency, and showed clear intermittency. The modal decomposition of CITs and ICITs showed that CITs were dominated by mode-1, whereas mode-1 and higher modes in ICITs signals showed comparable amplitudes. CITs and ICITs accounted for approximately 64% and 36% of the total kinetic energy of internal tides, respectively.     

Energetics and temporal variability of internal tides in Luzon Strait: a nonhydrostatic numerical simulation

A fully nonlinear,three-dimensional nonhydrostatic model driven by four principal tidal constituents(M2,S2,K1,and O1) is used to investigate the spatial-temporal characteristics and energetics of internal tides in Luzon Strait(LS).The model results show that,during spring(neap) tides,about 64(47) GW(1 GW=109 W) of barotropic tidal energy is consumed in LS,of which 59.0%(50.5%) is converted to baroclinic tides.About 22(11) GW of the derived baroclinic energy flux subsequently passes from LS,among which 50.9%(54.3%) flows westward into the South China Sea(SCS) and 45.0%(39.7%) eastward into the Pacific Ocean,and the remaining 16(13) GW is lost locally owing to dissipation and convection.It is revealed that generation areas of internal tides vary with the spring and neap tide,indicating different source areas for internal solitary waves in the northern SCS.The region around the Batan Islands is the most important generation region of internal tides during both spring and neap tides.In addition,the baroclinic tidal energy has pronounced seasonal variability.Both the total energy transferred from barotropic tides to baroclinic tides and the baroclinic energy flux flowing out of LS are the highest in summer and lowest in winter.     

Nonlinear interaction between astronomical tides and storm surges at wusong tidal station

There are obvious periodic oscillations in the observations of storm surges in the East China Sea. The storm surges are not only controlled by the wind stresses and isolated long wave caused by typhoons but also affected by the interaction between astronomical tides and storm surges. In the present paper we simulate the interaction between tides and storm surges by using a two dimensional numerical model. In our numerical experiments we use the data of the storm surge induced by Typhoon 8114. The calculations tally with the measured data well. The results indicate that the periodic oscillations occurring in the elevations of the surge are mainly caused by the interaction between the tide and the storm surge. The numerical experiments also indicate that the forecasting precision may be notably improved if the nonlinear interaction between tides and storm surges is taken into account.     

INTERNAL TIDES, SOLITARY WAVES AND BORES IN SHALLOW SEAS

Remote sensing and in situ observations of internal tides, solitary waves and bores in shallow water are briefly reviewed in this paper. The emphasis is laid on interpreting SAR images based on oceanographic measurements, and analyzing characteristics of internal waves in the China Seas. Direc-tions for future research are discussed.     

A TWO-DIMENSIONAL VORTICITY MODEL OF INTERNAL TIDES GENERATED ON THE CONTINENTAL SHELF/SLOPE

A two-dimensional spectral-difference mode (with vorticity and density equations) of internal tides isdeveloped for studying the genration and propagration of internal tides generated at the continentalshelf/slope. In general, internal tides propagate seaward in deep sea regions and shoreward on the shelf,and are dissipated rapidly. When the Vaisala frequency decreases vertically, waves may be mostly limited to thecontinental slope region. in deep sea region, motions may have either boam-like structure or modal structure,depending on the stratification strerigth and structure, whereas a modal structure may always exist onthe shelf. Waves show strong bottom intensification on the slope when strong stratification exists on thebottom. The barotropic tidal advection may affed the temporal character of internal tides at thecontinental slope, shelf break and shelf regions. but may have little influence on the energy density and energy flux of internal tides. ln the case of strong stratification, waverforms of     

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.     

First record of Ulva californica in the mainland of China: a single alien parthenogenetic population in discontinuous distribution

Journal of Oceanology and Limnology - Molecular investigations have raised concerns about the ecological risks of green tides caused by alien Ulva species in new habitats. The green tide-forming...     

A New Molecular Label Applied to the Study of the Yellow Sea Green Tide

From 2007 to 2017, large-scale green tides occurred every year in the Yellow Sea of China, and Ulva prolifera was the main species leading to the green tide. In this study, we used the Polymerase chain reaction and 3' Rapid-amplification of c DNA ends technique to amplify the nr DNA-LSU and IGS sequences in U. prolifera and one species of Blidingia. These techniques showed 3259 bp of nrD NA-LSU and 3388 bp of IGS in U. prolifera and 3282 bp nr DNA-LSU and 3059 bp IGS in Blidingia sp. At the same time, tandem repeats, short dyads, palindromic and multiple simple repeat sequences in the IGS sequence were found by analyzing the structure of the IGS sequence in U. prolifera and Blidingia sp. G + C contents of the IGS sequence in U. prolifera and Blidingia sp. were 52.42% and 53.09%, respectively. We divided the U. prolifera into two types according to the morphological characteristics. Although the specimens of U. prolifera from the Qingdao coastal area, Jiangsu coastal area and the Yellow Sea have different morphologies, their ITS and IGS sequences are almost identical. Therefore, the main species of the green tides in the Qingdao coastal area, Jiangsu coastal area and the Yellow Sea are the same and have the same origin.     

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

Journal of Oceanology and Limnology - Qinhuangdao coastal waters have been frequently hitting by nano- and pico-eukaryotic phytoplankton (NPEP) blooms and green tides (macroalgal blooms) in the...     

AN EXACT SOLUTION OF INTERNAL TIDES GENERATED AT CONTINENTAL SLOPE

By using a coordinate transformation,an exact solution of internal tides is obtained when the bot- tom slope is linear and the V(?)is(?)la frequency is constant.Consequently the dispersion relations of free waves are presented.Compared with Baines solution,the solution derived here is more consistent with experiment     

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

Journal of Oceanology and Limnology - Large-scale green tides occur frequently in summer in the Yellow Sea in recent decade. Micro-propagules, providing “seeds” for the seaweeds, play...     

A layered numerical model for simulating the generation and propagation of internal tides over continental slope III. Numerical experiments and simulation

The layered model in Part I was used to simulate the internal tide in a stratified, two layer, and rectangular sea area with step-like topography. The internal tide current velocities of the upper and lower layers and the interfacial elevations were computed and the effect of the upper layer water depth and density difference were studied. Numerical experiments verified that the model can correctly simulate internal tides. The model was also applied to the northwestern part of the South China Sea to simulate the internal tides there with real topography. The distributions of internal tide amplitude in interfaces were delineated. Contribution No. 3590 from the Institute of Oceanology, Chinese Academy of Sciences. Project 49676275 supported by NSFC and also supported by National Special Research Program 97-926-05-02.     

Global ocean tides from Geosat altimetry by Quasi-Harmonic analysis

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Bacterial diversity in surface water of the Yellow Sea during and after a green alga tide in 2008

From May to August 2008, a large "green tide", consisting of the alga Ulva (Enteromorpha) prolifera, occurred in the Yellow Sea, China, affecting the local marine ecosystem and human activities. We investigated the influence of the green tide on the microbial community in the surface seawater, at four sites from July to August 2008, using bacterial 16S rRNA gene clone libraries. We sequenced 228 clones of unique patterns identified by restriction fragment length polymorphism (RFLP) techniques. The results show that 228 sequenced clones fell into six bacterial phyla:Proteobacteria, Bacteroidetes, Cyanobacteria, Verrucomicrobia, Actinobacteria, and Planctomycetes. Alphaproteobacteria (33%), Gammaproteobacteria (25%), Bacteroidetes (23%) and Cyanobacteria (9%) dominated the assemblage. Comparison between samples collected in July (during the tide) and those collected in August (after the tide) showed that, in the microbial community, diversities of Alphaproteobacteria and Cyanobacteria increased after the tide, while those of Gammaproteobacteria and Bacteroidetes decreased. These results indicate that the green tide influenced the growth of some bacteria, and provide information for further studies on the interactions and relationships between U. prolifera and the bacterial community. This study suggests that microbial community analysis is a good approach to monitoring green tides.     

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.     

A layered numerical model for simulating the generation and propagation of internal tides over continental slope

A layered three-dimensional nonlinear numerical model was constructed to simulate the generation and propagation of internal tides over the continental slope. The simulation was split into external mode computation (EMC) and internal mode computation (IMC) to minimize the computational load. IMC was carried out once after EMC was implemented N times. As to EMC, a semi-implicit numerical scheme was applied in such a way that the pressure gradient terms and the velocity divergence terms were discretized semi-implicitly, but the other terms were discretized explicitly. Eulerian-Lagrangian explicit discretization was applied to the convective terms simultaneously. As a result, the stability of EMC did not depend on the wave celerity and time step was not limited by the CFL condition. More than that, use of the conjugate gradient accelerated Jacobi method further improved the computational efficiency of the model. Contribution No. 3588 from the Institute of Oceanology, Chinese Academy of Sciences. Project 49676275 supported by NSFC and National Special Research Program 97-925-05-02.