Calcareous dinoflagellate cysts in South and equatorial Atlantic surface sediments: diversity,distribution, ecology and potential for palaeoenvironmental reconstruction

Several marine, peridiniphycidean dinoflagellate species produce calcareous cysts during their life cycle, which are relatively resistant to chemical and physical degradation and are therefore often found in large quantities in oceanic bottom sediments. Although the use of these calcareous cysts as proxies for palaeoenvironmental and palaeoclimatic reconstructions has seen many advances over the last decade, until now only relatively patchy and regional information was available on their recent distribution patterns and ecology, especially at the species level. In this paper, comprehensive calcareous cyst diversity and distribution data have been compiled from published and unpublished work for 167 South and equatorial Atlantic Ocean surface sediments, ranging from 20°N to 50°S, and 30°E to 65°W. The main aim has been to focus on the complex, often non-linear, relationships between individual species’ distributions and the physicochemical and trophic conditions of the overlying (sub)surface waters through the use of x–y graphs of cyst abundance vs. (sub)surface water environmental parameters, and detrended correspondence analyses. Ten cyst species and the calcareous vegetative coccoid species Thoracosphaera heimii were observed in the bottom sediments, each species showing its own characteristic distribution pattern in relation to the environmental conditions of the upper water masses above them (e.g. sea surface temperature, productivity, stratification). The sensitive reactions of various species to unique combinations of environmental parameters shows that each species has its own specific ecological traits, thus rejecting the bundled use of ‘calcareous cyst accumulation’ as a general proxy for oligotrophy or stratification in future palaeoenvironmental analyses. The acquired ‘reference’ data set of this study is large and diverse enough to allow its future application in quantitative palaeoenvironmental reconstruction models, and shows that there is still an enormous reconstruction potential concealed in many fossil calcareous dinoflagellate cyst assemblages.     

Distributions of calcareous dinoflagellate cysts in surface sediments of the western equatorial Atlantic Ocean,and their potential use in palaeoceanography

Only very few studies focus on recent calcareous dinoflagellate cyst diversity, geographic distribution and ecology, so that information on the distribution patterns and environmental affinities of individual cyst species is extremely limited. This information is, however, essential if we want to use calcareous dinoflagellate cysts for palaeoenvironmental reconstruction. Surface sediment samples from the generally oligotrophic western equatorial Atlantic Ocean, offshore northeast Brazil, were therefore quantitatively analysed for their calcareous dinoflagellate cyst content, including the calcareous vegetative coccoid Thoracosphaera heimii. Seven calcareous dinoflagellate cyst species/morphotypes and T. heimii were encountered in high concentrations throughout the area. Substantial differences in the distribution patterns were observed. The highest concentrations of cysts are found in sediments of the more oligotrophic, oceanic regions, beyond the influence of Amazon River discharge waters. Dinoflagellates producing calcareous cysts thus appear to be capable of surviving low nutrient concentrations and produce large numbers of cysts in relatively stable and predictable environments affected by minimal seasonality. To test for the environmental affinities of individual species, distribution patterns in surface sediments were compared with temperature, salinity, density and stratification gradients within the upper water column (0–100 m) over different times of the year, using principal components analysis and redundancy analysis. T. heimii and four of the seven encountered cyst species (Sphaerodinella? albatrosiana, two morphotypes of Sphaerodinella? tuberosa and Scrippsiella regalis) relate to these parameters significantly and the variations in the cyst associations appear to be associated with the different surface water currents characterising the area. The results imply that calcareous dinoflagellate cyst distributions can potentially be used to distinguish between different open oceanic environments and they could, therefore, be useful in tracing water mass movements throughout the late Quaternary.     

甲藻孢囊在长江口海域表层沉积物中的分布

为了了解长江口海域赤潮爆发潜势,于2002年4月至5月用采泥器采集了位于122°～123．5°E、29°～32°N之间12个站位的表层沉积物,分析沉积物中甲藻孢囊的分布．共分析鉴定出孢囊类型29种,其中自养型11种,异养型18种．每个站位的孢囊种类在10～21之间,孢囊密度为11．7～587孢囊·g-1干泥之间．远岸海域孢囊种类较为丰富,密度也较高．在调查区域内,孢囊密度及种类自西向东、自北向南逐渐增加．亚历山大藻孢囊分布广泛,最高密度为40．4孢囊·g-1干泥,其他赤潮种类的孢囊如链状裸甲藻、多边舌甲藻、锥状斯氏藻、科夫多沟藻和无纹多沟藻等都在长江口海域有分布．     

Seasonal distribution of dinoflagellate resting cysts in surface sediments from Changjiang River Estuary

In order to understand the distribution of dinoflagellate cysts, surface sediments were collected from 15 stations in Changjiang River Estuary from 122°E to 123.5°E and from 29°N to 32°N in four cruises from May 2002 to February 2003. In the present study, 38 different cyst morphotypes representing 21 genera and 6 groups were identified, while 1 type was not identified into genus level. Species number and cell density of dinoflagellate cysts ranged from 10 to 25 species and from 12 to 587 per gram of dry weight, respectively. There were no obvious differences in cyst composition and density among seasons. However, the highest cyst species number and density were recorded in summer and winter, respectively. Cysts of heterotrophic dinoflagellates, which held 55.7% of the overall cyst density averagely, dominated cyst assemblages. Cyst density and species number increased from the west to the east, from the north to the south within the study area. Cysts of toxic dinoflagellates Alexandrium cat‐enella and Alexandrium tamarense complex distributed widely and were observed in almost all stations, with the maximum cell density of 81 per gram of dry weight.     

甲藻孢囊在长山群岛海域表层沉积物中的分布

2007年10月,在大连长山群岛海域采集了25个站点的表层沉积物,研究了甲藻孢囊的水平分布。在沉积物样品中共检测出甲藻孢囊46种,其中Pentaphasodinium dalei和Warowia rosea,在我国是首次报道。各站位孢囊种类数在7-20之间,平均14种;丰度在182-2960 个/g干重之间,平均为1201 个/g干重。香农-威纳指数(H')的变化范围为2.4-3.4。麻痹性贝毒藻--亚历山大藻的孢囊在调查海域分布广泛,最高密度达829个/g干重,可能是造成该海域虾夷扇贝贝毒超标的原因之一。     

Organic-walled dinoflagellate cysts in western equatorial Atlantic surface sediments: distributions and their relation to environment

In contrast to the wide range of studies carried out in temperate and high-latitude oceanic regions, only a few studies have focused on recent and Holocene organic-walled dinoflagellate cyst assemblages from the tropics. This information is, however, essential for fully understanding the ability of species to adapt to different oceanographic regimes, and ultimately their potential application to local and regional palaeoenvironmental and palaeoceanographic reconstructions. Surface sediment samples of the western equatorial Atlantic Ocean north of Brazil, an area greatly influenced by Amazon River discharge waters, were therefore analysed in detail for their organic-walled dinoflagellate cyst content. A diverse association of 43 taxa was identified, and large differences in cyst distribution were observed. The cyst thanatocoenosis in bottom sediments reflects the seasonal advection of Amazon River discharge water through the Guyana Current and the North Equatorial Countercurrent well into the North Atlantic. To establish potential links between cyst distribution and the environmental conditions of the upper water column, distribution patterns were compared with mean temperature, salinity, density and stratification gradients within the upper water column (0-100m) over different times of the year, using correspondence analysis and canonical correspondence analysis. The analyses show that differences in these parameters only play a subordinate role in determining species distribution. Instead, nutrient availability, or related factors, dominates the distribution pattern. The only possible indicators of slightly reduced salinities are Trinovantedinium applanatum and Lingulodinium machaerophorum. Four assemblage groups of cyst taxa with similar environmental affinities related to specific water masses/currents can be distinguished and have potential for palaeoenvironmental reconstruction.     

A sampling device to measure in situ germination rates of dinoflagellate cysts in surface sediments

The design and function of a new device to collect in situ germinatingcells from settled cysts of dinoflagellates in surface sedimentsare reported. Experiments to determine the effectiveness ofthe sampler indicate that it is applicable to studies on thepopulation dynamics of dinoflagellates.     

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea: distribution patterns and influence of main environmental gradients

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.     

深圳湾表层沉积物中甲藻孢囊的垂直分布

采集了深圳湾3个采样点约20cm的表层沉积物样品,研究了甲藻孢囊的垂直分布。共分析鉴定出孢囊类型37种,其中5种为我国沿海未报道的新记录类型,原多甲藻类孢囊是该海域种类最丰富的孢囊类型。除上表层沉积物外,深圳湾甲藻孢囊密度较低,孢囊密度大多在500cysts／gDWt以下,总孢囊密度变化范围为83～4036cysts／gDWt之间。锥状斯氏藻是深圳湾甲藻孢囊的优势种类,上表层沉积物中该孢囊密度的急剧上升证实了2000年附近海域所发生的该藻赤潮。而裸甲藻类孢囊含量明显比南海其他海域高。孢囊的香农-威弗种类多样性指数为1．50～3．96之间,并且在非养殖区、离岸海域及较深层次沉积物中较高,而孢囊密度则呈相反的变化趋势。种类多样性的下降及孢囊密度的上升在一定程度上说明深圳湾污染的变化趋势,富营养化程度非养殖区大于养殖区,近岸海域大于离岸海域,并且近年来污染逐渐加剧。     

九龙江西陂库区沉积物甲藻孢囊的分布

孢囊在甲藻的生活史中发挥重要作用,福建省九龙江从2009年起,暴发多次拟多甲藻水华事件。采用显微镜观察和单细胞PCR技术,对九龙江西陂库区2012—2013年不同月份沉积物中的甲藻孢囊进行种属判定,并对甲藻孢囊的分布及其影响因素进行分析。结果表明,西陂库区沉积物中的甲藻孢囊主要为拟多甲藻属,约占80%,其次为裸甲藻属,发现了2009年水华的优势种佩式拟多甲藻(Peridiniopsis penardii)孢囊。库区沉积物中甲藻孢囊的丰度在(13.7±1.2)—(105.2±8.3)个/g干重之间。多元相关分析结果显示甲藻孢囊的丰度与含水率呈现显著正相关性(P0.05),反映了甲藻孢囊沿水流方向逐渐积累。本研究结果填补了国内水库甲藻孢囊鉴定和萌发的研究空白,为九龙江甲藻水华的防治提供科学参考。     

Effect of salinity on the skeletal chemistry of cultured scleractinian zooxanthellate corals: Cd/Ca ratio as a potential proxy for salinity reconstruction

The effect of salinity on the elemental and isotopic skeletal composition of modern zooxanthellate scleractinian corals (Acropora sp., Montipora verrucosa and Stylophora pistillata) was investigated in order to evaluate potential salinity proxies. Corals were cultured in the laboratory at three salinities (36, 38 and 40). The other environmental parameters were kept constant. For all species analyzed, Sr/Ca, Mg/Ca, U/Ca and Li/Ca ratios were not influenced by salinity changes. The Ba/Ca ratio also lacks a systematic relationship with salinity and exhibits high inter-generic variations, up to one order of magnitude. On the contrary, the Cd/Ca ratio decreases as a function of increasing salinity, and δ¹⁸O and δ¹³C also presented a significant response, but with opposite trends to salinity variations. Since Cd/Ca is usually considered as an upwelling proxy, its salinity dependence could compromise the upwelling signal, unless some corrections can be carried out. Regardless, if the dependence found in the present dataset proved to be widespread and systematic, the Cd/Ca ratio could represent a promising salinometer awaiting further investigation. This study also confirmed the reliability of the well-established temperature proxies Sr/Ca, Mg/Ca and U/Ca, as these ratios were insensitive to salinity variations. Moreover, our results showed that δ¹⁸O or δ¹³C can be considered as reliable temperature recorders as far as the salinity effect is removed from the parameter reconstructed (e.g., temperature). Investigating the influence of salinity on the skeletal chemistry of scleractinian corals grown under controlled environmental conditions confirmed previous results, validated isotopic corrections, and identified a promising proxy of salinity.     

Spatial distribution of calcareous dinoflagellate cysts in surface sediments of the Atlantic Ocean between 13 degrees N and 36 degrees S

To enhance the limited information available about the palaeo-ecological significance of calcareous dinoflagellates, we have studied their lateral distribution in surface sediments of the equatorial and south Atlantic between 13 degrees N and 36 degrees S. Calcareous dinoflagellate cysts appear to be widely distributed throughout the studied area. In the surface sediments, concentrations (cyst per gram dry sediment) of the vegetative stage Thoracosphaera heimii are generally higher than that of the (presumably) calcareous resting cysts. Distribution patterns in surface sediments of Orthopithonella granifera (Fütterer) Keupp and Versteegh, Rhabdothorax spp. Kamptner., Sphaerodinella albatrosiana (Kamptner) Keupp and Versteegh S. albatrosiana praratabulated, Sphaerodinella tuberosa var. 1 (Kamptner) Keupp and Versteegh and S. tuberosa var. 2 and the ratios between these species have been compared with temperature, salinity, density and stratification gradients in the upper water column. Rhabdothorax spp. is characteristically present in sediments of more temperate regions characterized by high seasonality. Dinoflagellates producing these cysts are able to tolerate high nutrient concentrations, and mixing of the water column. S. albatrosiana is abundant in regions characterized by high sea surface temperatures and oligotrophic surface water conditions. In contrast, the distribution of S. tuberosa var. 2 is negatively related to temperature. The other cyst species did not show a characteristic pattern in relation to the studied environmental gradients.The ratio of Sphaerodinella tuberosa var. 2 to Orthopithonella granifera can be used for reconstructing the presence of stratification in the upper 50m of the water column, whereas the ratios of S. tuberosa var. 2 to Sphaerodinella albatrosiana and of O. granifera to Rhabdothorax spp. might be used for palaeotemperature reconstructions. Calcareous dinoflagellate cysts are abundant in oligotrophic areas and may be useful for the reconstruction of palaeoenvironmental conditions.     

Organic-walled dinoflagellate cysts as paleoenvironmental indicators in the Paleogene; a synopsis of concepts

The organic-walled, hypnozygotic cysts of dinoflagellates (“dinocysts”) provide a rich, albeit incomplete, history of this eukaryotic plankton group in ancient sediments. Building on pioneering studies of the late 1970s and 1980s, recent ocean drilling and more detailed, integrated studies of surface sections have provided a wealth of dino-cyst data spanning the entire Paleogene. Based on multidisciplinary approaches, these studies have been instrumental in refining existing and furnishing new concepts of Paleogene paleoenvironmental and paleoclimatic reconstructions by means of dinocysts. Since dinocysts typically exhibit high abundances in neritic settings, the dinoeyst-based environmental and paleoclimatic information is complementary to that derived from typically more offshore groups such as planktonic foraminifera, coccolithophorids, diatoms, and radiolaria. While in a recent paper we gave a broad overview of case studies from around the globe (Sluijs et al. 2005), here we focus on a summary of these analyses and present a synopsis of applied paleoecological concepts in Paleogene (65–24 ma) dinocyst studies. Representing Earth’s greenhouse-icehouse transition, this episode holds the key to the understanding of extreme transient climatic change. The present paper offers guidelines for the application of dinocyst palaeoecology to the reconstruction of Paleogene sea-surface productivity, temperature, salinity, stratification, and paleo-oxygenation as well as their application in sequence stratigraphy, oceanic circulation and general watermass reconstructions.      

Taxonomy and palaeoecology of dinoflagellate cysts from Upper Oligocene freshwater sediments of Lake Enspel, Westerwald area, Germany

Freshwater dinoflagellates play an important role as primary producers in the lacustrine environment. A new species of dinoflagellates, Cleistosphaeridium lacustre, is described from Upper Oligocene sediments of palaeo-lake Enspel. They are associated with other phytoplankton, such as diatoms, chrysophytes, green algae and benthic cyanobacteria. Mass occurrences of this species are interpreted as algal blooms and may partly reflect seasonal successions. This phenomenon was controlled by volcanic activities in the depositional area, which led to an increase in nutrient supply.     

Dinoflagellate cysts from surface sediments of Saldanha Bay, South Africa: an indication of the potential risk of harmful algal blooms

The distribution and abundance of dinoflagellate cysts from recent coastal sediments in Saldanha Bay, was investigated, and compared to the cyst assemblages of the adjacent coastal upwelling system as reflected in the sediments off Lambert's Bay on the southern Namaqua shelf. Twenty-two cyst types were identified from three sample sites off Lambert's Bay with recorded abundances between 1726 and 1863 cysts ml⁻¹ wet sediment. At least 21 distinctive cyst types were identified from 32 sample sites within Saldanha Bay. Cyst abundance in Saldanha Bay was relatively low, averaging 116 cysts ml⁻¹ wet sediment. The region off Lambert's Bay is especially susceptible to the formation of harmful algal blooms attributed to high biomass dinoflagellate blooms. Owing to these blooms and the retentive circulation characteristics of this area, cyst formation and deposition is high. Blooms can be advected into Saldanha Bay, but their development and duration in the Bay is restricted by the system of exchange that operates between the Bay and the coastal upwelling system, in that there is a net export of surface waters from the Bay. Consequently, fewer cysts are formed and deposited within the Bay thereby reducing the likelihood of in situ bloom development initiated from the excystment of cysts.     

Formation of kynurenic and xanthurenic acids from kynurenine and 3-hydroxykynurenine in the dinoflagellate Lingulodinium polyedrum: role of a novel,oxidative pathway

The dinoflagellate Lingulodinium polyedrum (syn. Gonyaulax polyedra) was used as a model organism for studying the effects of high and low physiological oxidative stress on the formation of kynurenic and xanthurenic acids from kynurenine and 3-hydroxykynurenine. Cell were incubated with the precursors and exposed to light (high physiological stress due to photosynthetically formed oxidants) or kept in darkness (low stress). In cultures of less than 0.5 ml cell volume/l of medium, cells took up approximately one half of 0.1 mM extracellular kynurenine within 18 h. The amino acid was partially converted to kynurenic acid, most of which was released to the medium; however, intracellular concentrations of the product were by approximately 10-fold higher than extracellular levels. Rates of kynurenic acid release exceeded by far those explained by kynurenine and tryptophan aminotransferase activities, the latter representing an additional source of kynurenic acid formation via indole-3-pyruvic acid. Light enhanced the release of kynurenic acid by approximately 4-fold; these rates were further increased by exposure to continuous light. Diurnal rhythmicity of kynurenic acid release was clearly exogenous and did not match with the circadian pattern of kynurenine or tryptophan aminotransferase activities; no rhythm was detected in constant darkness. Similar findings were obtained on turnover of 3-hydroxykynurenine to xanthurenic acid and release of the product to the medium. However, light/dark differences were relatively smaller, and additional products were formed, according to HPLC data obtained with electrochemical detection. Results are most easily explained on the basis of a recently discovered pathway of kynurenic acid formation from kynurenine, involving either non-enzymatic oxidation by H(2)O(2) or, at higher rates, enzymatic catalysis by hemoperoxidase. A corresponding mechanism may exist for the hydroxylated analogue.     

Circadian rhythm of a TCA cycle enzyme is apparently regulated at the translational level in the dinoflagellate Lingulodinium polyedrum

Previously, the authors have reported that intracellular amounts of several metabolic-related enzymes from the photosynthetic dinoflagellate Lingulodinium polyedrum(formerly Gonyaulax polyedra) showed a daily rhythm under a 12:12 h LD cycle. This led the authors to hypothesize that a circadian clock controls metabolism, including the tricarboxylic acid (TCA) cycle. In this study, the authors investigated daily changes in the levels of mRNA, protein, and enzyme activity of several metabolic enzymes during 12:12 h LD, 8:16 h LD, and constant light conditions. The NADP-dependent isocitrate dehydrogenase (NADPICDH) in the TCA cycle exhibited circadian changes of protein abundance and enzyme activity under all conditions, whereas its mRNA level remained constant throughout the cycle. These results indicate that the rhythm of NADPICDH is regulated by a circadian control of protein synthesis or modification rather than by message levels and suggest that the TCA cycle may be controlled by the circadian clock system.     

The uropod as a proxy for total length distribution in Antarctic krill: an assessment of different models

The properties of the regression of standard length on the length of the exopodit of the uropod of Euphausia superba as a method to estimate the distribution of the standard length from penguin stomach contents samples are explored. The distance between the estimated and observed distributions was measured with the Kolmogorov–Smirnov statistic. Sample variability was determined using resampling techniques. The ability of linear functions (Models I and II), allometric and non-parametric methods to recover the observed distributions was evaluated within samples and exchanging parameters between samples. Linear and allometric models proved inadequate to recover the distribution, while the differences obtained with non-parametric methods fall within the bounds of sample variability, suggesting that a linear equation does not capture the relation between total and exopodit length. The use of a non-parametric regression is recommended to increase the sample size when estimating the distributions of prey lengths in the stomach contents of Antarctic penguins.     

BET proteins: Investigating BRDT as a potential target for male contraception

While many contraception options are available for women, birth control methods for men are limited to condoms and vasectomy. Past research into male contraceptives has focused on hormonal options but the associated side effects have thus far precluded this method from reaching the market. Non-hormonal male contraceptives and vas occlusion have also been explored, but to date no method has progressed past clinical testing. Recent interest in epigenetic research has unveiled a new potential non-hormonal male contraceptive target: the testis-specific bromodomain BRDT. Potent inhibitors for bromodomain-containing proteins are described in the literature, but a BRDT-specific compound has yet to be designed, prepared and tested. The high similarity between bromodomain proteins of the BET family makes development of selective and specific inhibitors both difficult and necessary. Selective inhibition of BRDT by a small molecule is an exciting new target in the search for a new non-hormonal male contraceptive.