Largest Macroalgal Blooms Research Articles

Diel metabolism of Yellow Sea green tide algae alters bacterial community composition under in situ seawater acidification of coastal areas

Ocean acidification in coastal seawaters is a complex process, with coastal pH being affected by numerous factors including watershed and biological processes that also support metabolically diverse bacterial communities. The world's largest macroalgal blooms have occurred consecutively in the Yellow Sea over the last 13 years. In particular, algal mats formed by Yellow Sea green tides (YSGT) significantly influence coastal environments. Herein, we hypothesized that 1) inorganic carbonate chemistry in coastal areas is altered by diel metabolism of these giant algal mats and that 2) bacterial community composition in diffusive boundary layers might be altered along diel cycles due to algal mat metabolism. In situ studies indicated that algal mat metabolism led to changes in diel pH and CO2 in affected seawaters. Such metabolic activities could intensify diel pH fluctuations in algal mat diffusive boundary layers, as noted by pH fluctuations of 0.22 ± 0.01 units, and pCO2 fluctuations of 214.62 ± 29.37 μatm per day. In contrast, pH fluctuations of 0.11 ± 0.02 units and pCO2 fluctuations of 79.02 ± 42.70 μatm were noted in unaffected areas. Furthermore, the bacterial community composition associated with diffusive algal boundary layers, including those of ambient bacteria and epiphytic bacteria, exhibited diel changes, while endophytic bacterial communities were relatively stable. Flavobacteriaceae were particularly highly abundant taxa in the ambient and epiphytic bacterial communities and exhibited increased abundances at night but sharp decreases in abundances during daytime. Flavobacteriaceae are heterotrophic taxa that could contribute to coastal area acidification at night due to the transformation of organic carbon to inorganic carbon. These results provide new insights to understand the variability in coastal ocean acidification via harmful algal blooms while providing a framework for evaluating the effects of YSGT on costal carbon cycling.

Comparing performance of inter-sensor NDVI for the detection of floating macroalgal blooms in the Yellow Sea

There are many complains against macroalgal bloom (MAB) from all over the world for its high negative economic impacts. Early and precise detection of MAB occurance can reduce huge economic loss in tourism and marine business. Thus, this study aims to detect the MAB in the Yellow Sea (YS) by comparing performance of Normalized Difference Vegetation Index (NDVI) between Geostationary Ocean Color Imager (GOCI) and Landsat-8 sensors from 2014 to 2019. The largest MAB during this period was identified from Landsat-8 sensors along the coast of Sheyang and Qingdao of China in 2018. The locations of MABs were almost nearby areas during the begining phase of MABs in consecutive years. In successive years huge and dense MABs reached the brink of shoreline of Qingdao. In Landsat-8 images in most cases larger total areas of MABs were detected including very small pathches which could not be detected from GOCI due to low spatial resolution. The findings of this study will be helpful for scientists to choose between GOCI and Landsat-8 based on the objectives of their study.

Molecular cloning and transcriptional regulation of two γ-carbonic anhydrase genes in the green macroalga Ulva prolifera.

Ulva prolifera O.F. Müller (Ulvophyceae, Chlorophyta) is well known as a typical green-tide forming macroalga which hascaused the world's largest macroalgal blooms in the Yellow Sea of China. In this study, two full-length γ-carbonic anhydrase (γ-CA) genes (UpγCA1 and UpγCA2) were cloned from U. prolifera. UpγCA1 has three conserved histidine residues, which act as an active site for binding a zinc metal ion. In UpγCA2, two of the three histidine residues were replaced by serine and arginine, respectively. The two γ-CA genes are clustered together with other γ-CAs in Chlorophyta with strong support value (100% bootstrap) in maximum likelihood (ML) phylogenetic tree. Quantitative real-time PCR (qRT-PCR) analysis showed that stressful environmental conditions markedly inhibited transcription levels of these two γ-CA genes. Low pH value (pH 7.5) significantly increased transcription level of UpγCA2 not UpγCA1 at 12h, whereas high pH value (pH 8.5) significantly inhibited the transcription of these two γ-CA genes at 6h. These findings enhanced our understanding on transcriptional regulation of γ-CA genes in response to environmental factors in U. prolifera.

Analysis of the interannual variation characteristics of the northernmost drift position of the green tide in the Yellow Sea.

The green tide in the Yellow Sea is the world's largest macroalgal bloom. The maximum area affected by green tide can reach tens of thousands of square kilometers. Previous studies have shown that green tide drifts northward during the outbreak, yet the location of its northernmost drift and the characteristics of interannual variations have not been explored in detail. In this paper, we use the Moderate Resolution Imaging Spectrometer (MODIS) data, GaoFen-1 (GF-1) satellite data, unmanned aerial vehicle (UAV) aerial data, and field trips for extraction and monitoring of green tide and then analyze the emergence, development, and dissipation of the green tide in Rongcheng coastal waters and the change of the northernmost drift position and its related factors from the year of 2013 to 2018. The results show that green tide have drifted to the Rongcheng coastal area over the years and the northernmost drift position can reach 37.662° N in 2014. Interannual difference of the northernmost position of the drift of the green tide is obvious, it is mainly affected by the wind, and there is no certain connection with the maximum coverage area, the coverage area when reaching the northernmost position, and the existence days of the green tide. These results can help to understand the drift range and dissipation process of green tide in the Yellow Sea.

The great Atlantic Sargassum belt.

Pelagic Sargassum is abundant in the Sargasso Sea, but a recurrent great Atlantic Sargassum belt (GASB) has been observed in satellite imagery since 2011, often extending from West Africa to the Gulf of Mexico. In June 2018, the 8850-kilometer GASB contained >20 million metric tons of Sargassum biomass. The spatial distribution of the GASB is mostly driven by ocean circulation. The bloom of 2011 might be a result of Amazon River discharge in previous years, but recent increases and interannual variability after 2011 appear to be driven by upwelling off West Africa during boreal winter and by Amazon River discharge during spring and summer, indicating a possible regime shift and raising the possibility that recurrent blooms in the tropical Atlantic and Caribbean Sea may become the new norm.

The biggest bloom

Marine Ecology![Figure][1] Underwater photograph taken near Key Largo, Florida, of a Sargassum mat from below CREDIT: CHRIS GUG/ALAMY STOCK PHOTO Floating mats of Sargassum seaweed in the center of the North Atlantic were first reported by Christopher Columbus in the 15th century. These mats, although abundant, have until recently been limited and discontinuous. However, Wang et al. report that, since 2011, the mats have increased in density and aerial extent to generate a 8850-kilometer-long belt that extends from West Africa to the Caribbean Sea and Gulf of Mexico (see the Perspective by Gower and King). This represents the world's largest macroalgal bloom. Such recurrent blooms may become the new normal. Science , this issue p. [83][2]; see also p. [27][3] [1]: pending:yes [2]: /lookup/doi/10.1126/science.aaw7912 [3]: /lookup/doi/10.1126/science.aay0989

Spatiotemporal Patterns and Morphological Characteristics of Ulva prolifera Distribution in the Yellow Sea, China in 2016–2018

The world’s largest macroalgal blooms, Ulva prolifera, have appeared in the Yellow Sea every summer on different scales since 2007, causing great harm to the regional marine economy. In this study, the Normalized Difference of Vegetation Index (NDVI) index was used to extract the green tide of Ulva prolifera from MODIS images in the Yellow Sea in 2016–2018, to investigate its spatiotemporal patterns and to calculate its occurrence probability. Using the standard deviational ellipse (SDE), the morphological characteristics of the green tide, including directionality and regularity, were analyzed. The results showed that the largest distribution and coverage areas occurred in 2016, with 57,384 km2 and 2906 km2, respectively and that the total affected region during three years was 163,162 km2. The green tide drifted northward and died out near Qingdao, Shandong Province, which was found to be a high-risk region. The coast of Jiangsu Province was believed to be the source of Ulva prolifera, but it was probably not the only one. The regularity of the boundary shape of the distribution showed a change that was opposite to the variation of scale. Several sharp increases were found in the parameters of the SDE in all three years. In conclusion, the overall situation of Ulva prolifera was still severe in recent years, and the sea area near Qingdao became the worst hit area of the green tide event. It was also shown that the sea surface wind played an important part in its migration and morphological changes.

Annual patterns of macroalgal blooms in the Yellow Sea during 2007-2017.

The world’s largest macroalgal blooms caused by Ulva prolifera have occurred in the Yellow Sea for 11 consecutive years. The area covered by blooms has been approximately 500 km2 in previous years, while in 2017, the maximum area decreased significantly to 312 km2. In this study, we concluded that species competition between Ulva and Sargassum (fast rise of the golden tides), extreme high sea surface temperature and harvest for floating Ulva macroalgae were the three critical factors influencing the sharp reduction in covered area for blooms in 2017. In addition, analysis of annual variations of Pyropia aquaculture area in the Southern Yellow Sea over the past two decades revealed that a great expansion in “Sansha” regions was mainly responsible for the initial blooms in 2007, and that this expansion supported the great biomass of the blooms in following years. Based on these findings, we suggest comprehensive utilization of the macroalgal blooms is a feasible way to control them.

Spatio-temporal patterns of Ulva prolifera blooms and the corresponding influence on chlorophyll-a concentration in the Southern Yellow Sea, China

The world's largest macroalgal blooms (MABs) caused by the Ulva prolifera outbreaks have occurred every summer since 2007 in the Southern Yellow Sea, China. Accumulating evidence showed that MABs may deteriorate the regional marine environment and influence the growth of some primary producers such as phytoplankton. In this study, we investigated the spatio-temporal patterns of U. prolifera green tides and chlorophyll-a concentration in the Southern Yellow Sea in 2015 using satellite images obtained from HJ-1 CCD, MODIS, and GOCI. The correlation between the distributions of U. prolifera abundance and chlorophyll-a concentration was analyzed quantitatively by setting up a series of 5 × 5 km experimental grids, and we also discussed the possible mechanisms about the influence of U. prolifera blooms on the other floating microalgae. The results showed that the development of U. prolifera blooms in the Southern Yellow Sea in 2015 could be featured as “appearance – development – outbreak – decline – disappearance”, while the concentration of chlorophyll-a showed “increase – sharp decline – slow recovery – stabilization” from April to August. We also found that the concentration of chlorophyll-a had the following relationships with U. prolifera temporally: (1) the concentration of chlorophyll-a increased with the growth of U. prolifera from April to mid-May; (2) the chlorophyll-a concentration decreased sharply with the dramatically increased coverage of U. prolifera in June; and (3) the chlorophyll-a concentration slowly recovered and finally stabilized as U. prolifera decreased in July. Generally, there was a negative correlation between the occurrence of U. prolifera and chlorophyll-a concentration in the Southern Yellow Sea, China. Our results showed that the outbreak of U. prolifera does have a certain impact on the growth and reproduction of planktonic microalgae, and it suggests that U. prolifera blooms have potentially altered the ecological balance in the coastal waters of the Southern Yellow Sea.

Variations of dominant free-floating Ulva species in the source area for the world’s largest macroalgal blooms, China: Differences of ecological tolerance

Species composition and seasonal variations of free-floating Ulva species were investigated in the source area of the world’s largest macroalgal blooms during 2009–2015. Based on a combination of a morphological analysis and sequences of nuclear-encoded ITS and 5S rDNA spacer regions, the dominant species in the free-floating Ulva community at the early stage of green tides were Ulva compressa, Ulva flexuosa, and Ulva linza. The first appearance of Ulva prolifera on the sea surface was in mid-May and it dominated the floating Ulva community in June from 2009 to 2011. From 2012–2015, U. prolifera was not only the first species to appear on the sea surface but also the dominant species during the whole early stage of green tides. To explain the successional mechanism, the effects of environmental factors on the growth of four Ulva species were examined in the laboratory under different combinations of light intensity and temperature. It was found that the highest growth rate of U. prolifera was 44.9%/d, which was much higher than the other three Ulva species. The strong tolerance of U. prolifera to extreme conditions also helps it survive and bloom in the Yellow Sea.

Complete mitochondrial genome of Ulva prolifera, the dominant species of green macroalgal blooms in Yellow Sea, China

Ulva prolifera (U. prolifera), a green macroalgae, is widely known as the dominant species of the world’s largest macroalgal blooms in the Yellow Sea, China. In this study, we sequenced and annotated the complete mitochondrial genome of U. prolifera (GenBank accession number: KU161104). The genome consists of circular chromosomes of 61 962 bp and encodes a total of 26 protein-coding genes include nine ribosomal protein genes, five atp genes, three cox genes, eight nad genes and cob gene. Phylogenetic analysis showed U. prolifera clustered into Ulvo phyceae clade and had close genetic relationship with algae Ulva fasciata.

Complete mitochondrial genome of Ulva linza, one of the causal species of green macroalgal blooms in Yellow Sea, China

Ulva linza, a green macroalgae, is one of the causal species of the world’s largest macroalgal blooms in the Yellow Sea, China. In this study, we sequenced and annotated the complete mitochondrial genome of U.linza (GenBank accession no. KU189740). The genome consists of circular chromosomes of 70 858 bp and encodes a total of 28 protein-coding genes including eight rps genes, three rpl genes, five atp genes, three cox genes, eight nad genes and cob gene. Phylogenetic analysis showed U. linza clustered into Ulvophyceae clade and had close genetic relationship with algae Ulva prolifera.

Interpreting the progressive eutrophication behind the world’s largest macroalgal blooms with water quality and ocean color data

World’s largest macroalgal blooms (MAB) caused by Ulva prolifera outbreak every summer in the Yellow Sea since 2007, which bring potential damage to regional marine environment and economics. Assessing the evolution of historic eutrophication status is of importance for exploring the reasons for the blooms. In this study, we used an area-weighted nutrient pollution index (AWCPI-NP) and ocean color data to assess temporal changes in eutrophication status in the Yellow Sea basin. Results show that significant shifts occurred from the pre-MAB phase (2001–2006) to MAB phase (2007–2012): the average AWCPI-NP was higher than that in the pre-MAB phase (2001–2006) by about 45 %; chlorophyll a concentration, an indicator of eutrophication, increased by about 15 % in the region of MAB. Meanwhile, in the Jiangsu Shoal, the origin place of the floating macroalgae, the annual in situ nutrient concentrations increased rapidly from 2000 to 2011. The result obtained through our approach highlights that the MAB in the Yellow Sea might be primarily linked to the progressive eutrophication. In order to reduce this hazard, policy makers should include this new knowledge for an effective countermeasure planning.

Variations of morphology and photosynthetic performances of Ulva prolifera during the whole green tide blooming process in the Yellow Sea

Since 2007, the world's largest macroalgal blooms have occurred along the coastal area of the Yellow Sea for 6 consecutive years. In 2012, shipboard surveying and satellite remote sensing were used to monitor the whole blooming process. The blooms originated in Rudong sea area of the South Yellow Sea where bloom patches were of dark green and filamentous thalli were the dominant morphology. The scale of the blooms reached its peak size in Rizhao sea area of the North Yellow Sea, and decreased promptly and became insignificant in Qingdao coast where the blooms turned yellow, mostly with air sac blades. Meanwhile, vegetative cells of the green tide algae changed into cytocysts gradually from which germ cells were released as the blooms drifted northward. Additionally, chlorophyll contents and fluorescence activity of free-floating thalli in the North Yellow Sea were both significantly lower than that in the South Yellow Sea. Those studies presented here contributed to increasing our understanding about how the green tide declined gradually in the North Yellow Sea.

The world's largest macroalgal bloom in the Yellow Sea, China: Formation and implications

The world's largest trans-regional macroalgal blooms during 2008–2012 occurred in the Yellow Sea, China. This review addresses the causes, development and future challenges in this unique case. Satellite imagery and field observations showed that the macroalgal blooms in the Yellow Sea originated from the coast of Jiangsu province and that favorable geographic and oceanographic conditions brought the green macroalgae from the coast offshore. Optimal temperature, light, nutrients and wind contributed to the formation and transport of the massive bloom north into the Yellow Sea and its deposition onshore along the coast of Shandong province. Morphological and genetic evidence demonstrated that the species involved was Ulva prolifera, a fouling green commonly found growing on structures provided by facilities of Porphyra aquaculture. Large scale Porphyra aquaculture (covering >20,000 ha) along the Jiangsu coast thus hypothetically provided a nursery bed for the original biomass of U. prolifera. Porphyra growers remove U. prolifera from the mariculture rafts, and the cleaning releases about 5000 wet weight tonnes of green algae into the water column along the coast of Jiangsu province; the biomass then is dispersed by hydrographic forcing, and takes advantage of rather high nutrient supply and suitable temperatures to grow to impressive levels. Certain biological traits of U. prolifera —efficient photosynthesis, rapid growth rates, high capacity for nutrient uptake, and diverse reproductive systems— allowed growth of the original 5000 tonnes of U. prolifera biomass into more than one million tonnes of biomass in just two months. The proliferation of U. prolifera in the Yellow Sea resulted from a complex contingency of circumstances, including human activity (eutrophication by release of nutrients from wastewater, agriculture, and aquaculture), natural geographic and hydrodynamic conditions (current, wind) and the key organism's biological attributes. Better understanding of the complex biological–chemical–physical interactions in coastal ecosystems and the development of an effective integrated coastal zone management with consideration of scientific, social and political implications are critical to solving the conflicts between human activity and nature.

Genetic and Marine Cyclonic Eddy Analyses on the Largest Macroalgal Bloom in the World

In 2008, a massive Ulva prolifera bloom, with a 3-million-ton biomass covering an area of 1.29 × 10(4) km(2) at its largest, suddenly appeared from May to July in South Yellow Sea. The mechanism behind the rapid growth of these seaweeds was investigated. Molecular phylogenetic analysis of free-floating algal samples from the Yellow Sea suggested that U. prolifera belong to one population, and that temporary cyclonic eddies in the Yellow Sea in late spring and early summer may help promote the proliferation of this bloom by providing seaweeds with sufficient growth time, abundant nutrition, and favorable habitats. The initial investigation on the relationship between marine cyclonic eddies and the route of free-floating algae extends our knowledge on how the emergence of free-floating macroalgal blooms in coastal areas could yield a large biomass.

Settlement of vegetative fragments of Ulva prolifera confirmed as an important seed source for succession of a large‐scale green tide bloom

Ulva prolifera green tides in the Yellow Sea of China in the past 3 yr are among the largest macroalgal blooms ever recorded. Satellite images, as well as wind and current patterns, provide a clear picture of the formation of these green tides; however, their place of origin remains unclear. Terrestrial investigation along the coastline of the Jiangsu Province and two shipboard surveys in the Yellow Sea in 2009 showed that filamentous Ulvaceae algae were prevalent in various environments in Jiangsu Province including floating rafts for Porphyra cultivation, land‐based bivalve culturing ponds, estuaries, embankments, intertidal stones, wharfs, and hard muddy coasts. Vegetative fragments in sediment samples were found in both shipboard surveys during the bloom and postbloom periods, and all of the samples were U. prolifera. Phylogenetic analysis of 5S spacer sequences showed that the green tide in the Yellow Sea was not of terrestrial origin. However, settled vegetative fragments of U. prolifera were an important seed source for the successive and more serious green tide blooms.

World’s largest macroalgal bloom caused by expansion of seaweed aquaculture in China

In late June 2008, just weeks before the opening of the Beijing Olympics, a massive green-tide occurred covering about 600km2 along the coast of Qingdao, host city for Olympic sailing regatta. Coastal eutrophication was quickly attributed with the blame by the international media and some scientists. However, we explored an alternative hypothesis that the cause of the green-tide was due to the rapid expansion of Porphyra yezoensis aquaculture along the coastline over 180km away from Qingdao, and oceanographic conditions which favoured rapid growth of the bloom and contributed to transport of the bloom north into the Yellow Sea and then onshore northwest to Qingdao. At its peak offshore, the bloom covered 1200km2 and affected 40,000km2. This is the largest green-tide ever reported, the most extensive translocation of a green-tide and the first case of expansive seaweed aquaculture leading to a green-tide. Given similar oceanographic conditions to those that occurred in 2008, these green-tides may re-occur unless mitigation measures such as those proposed here are taken.