Physiological responses of phytoplankton to turbulent and stable environments in an upwelling region

The assimilation of ¹⁴C into photosynthetic products was monitoredin two phytoplankton blooms growing in turbulent environmentsduring February and March, 1981. The March community appearedto store a large proportion of carbon in sugars, polysaccharidesand lipids in the upper euphoric zone, but as the cells weremixed to deeper depths the flow of carbon was directed intoorganic acids, amino acids and protein near the bottom of theeuphotic zone. In February, the winds that induced initial deepmixing moderated during the second half of the study allowingthe water mass to stabilize. Total assimilation by the communitywas greater under mixing conditions than in the stable water,implying faster growth in a turbulent environment. When thecommunity was growing in the stable water a greater percentageof carbon was incorporated into protein, however. Since algalgrowth is linked to protein synthesis it is suggested that growthin the stable environment was faster than in the turbulent environment.     

Physiological responses of phytoplankton to natural light regimes

Physiological responses to changes in light intensity and qualityare known to occur on many time scales. The mechanisms controllingthe responses on short time scales (such as the ‘flicker’effect and state transitions), medium rime scales (did changesand light-shade adaptation), and long time scales (species succession)are poorly understood. On the basis of present information,however, it is suggested that models of the effects of verticalmising on primary production could be improved by incorporatingmore basic information about some of the shorter time-scaleresponses. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP) First International Workshop heldat the Limnological Institute, University of Konstanz, in April1982.     

Marine phytoplankton growth and survival under simulated upwelling and oligotrophic conditions

Dialysis cultures of a diatom, Chaetoceros affinis Lauder, and a dinoflagellate, Gymnodnium splendens Lebour were subjected to simulated upwelling conditions which were followed by oligotrophy. Upwelling was simulated by pumping nutrient-enriched sea water past the enclosed cultures. Oligotrophy was simulated by pumping nutrient-poor sea water past the cultures. After the period of upwelling oligotrophy was carried on for 65 days for G. splendens and 80 days for C. affinis. Both algae survived oligotrophic conditions as evidenced by the ability to initiate new cultures or by observation of motility. Nutrient analyses showed that oligotrophy developed slowly — over a period of several days — so that the cultures were subjected to slow changes similar to those which might occur in the sea when upwelling ceases or when upwelled water moves offshore. C. affinis responded to oligotrophy by developing small flagellated cells (male gametes), by losing discrete protoplasts in silicified cells, and by the breaking up of normal chains of cells. Plastids in G. splendens cells became more discrete under oligotrophic conditions and some large non-flagellated cells were observed. The use of dialysis cultures to simulate features of the ocean is discussed.     

Rapid eco-evolutionary responses in perturbed phytoplankton communities

Biodiversity currently faces unprecedented threats owing to species extinctions. Ecologically, compensatory dynamics can ensure stable community biomass following perturbation. However, whether there is a contribution of genetic diversity to community responses is an outstanding question. To date, the contribution of evolutionary processes through genotype shifts has not been assessed in naturally co-occurring multi-species communities in the field. We examined the mechanisms contributing to the response of a lake phytoplankton community exposed to either a press or pulse acidification perturbation in lake mesocosms. To assess community shifts in the ecological response of morphospecies, we identified taxa microscopically. We also assessed genotype shifts by sequencing the ITS2 region of ribosomal DNA. We observed ecological and genetic contributions to community responses. The ecological response was attributed to compensatory morphospecies dynamics and occurred primarily in the Pulse perturbation treatment. In the Press treatments, in addition to compensatory dynamics, we observed evidence for genotype selection in two species of chlorophytes, Desmodesmus cuneatus and an unidentified Chlamydomonas. Our study demonstrates that while genotype selection may be rare, it is detectable and occurs especially when new environmental conditions are maintained for long enough to force selection processes on standing variation.     

Effects of simulated upwelling and oligotrophy on chemostat-grown natural marine phytoplankton assemblages

Natural phytoplankton assemblages from the Scripps Pier were grown in two chemostats under conditions that simulated two rates of upwelling followed by oligotrophic conditions. At a moderate upwelling rate (D = 0.3·day^?1) centric diatoms were selected, while at a low rate (D = 0.1·day^?1) a mixture of species dominated. Pumping of low-nutrient water (oligotrophy) resulted in a mixture of species at both rates. Upwelling at a high rate decreased diversity of the crop as compared with the low rate or oligotrophy. These results are compared with those of others who have subjected natural assemblages to continuous culture.     

Physiological responses of estuarine phytoplankton to ultraviolet light-induced fluoranthene toxicity

Polycyclic aromatic hydrocarbons (PAHs) are common pollutants associated with urbanization and suburbanization in estuarine systems, but little is known about their effects on the physiological properties of microalgae. We examined the effects of ultraviolet (UV)-activated fluoranthene toxicity on (a) the growth, chlorophyll a content cell⁻¹, and pigment composition of axenic Ankistrodesmus sp. (an estuarine benthic green microalga) and (b) the phytoplankton population growth and pigment composition of natural communities from an urbanized (Murrells Inlet) vs. forested (North Inlet) salt marsh estuary. The zeaxanthin/violaxanthin ratio increased in Ankistrodesmus sp. cultures exposed to UV light in the presence of fluoranthene, supporting the hypothesis that xanthophyll cycling is an energy dissipative response to photoinduced PAH toxicity in this species. Exposure of natural communities to the combination of UV light and fluoranthene resulted in decreased chlorophyll production and increased zeaxanthin violaxanthin⁻¹ in samples from the urbanized estuary (Murrells Inlet), but not North Inlet, suggesting that phytoplankton in the former “fluoranthene-impacted” estuary were more susceptible to fluoranthene toxicity. Consideration of xanthophyll cycling as a microalgal response to UV-activated PAH toxicity has implications to understanding the influence of these contaminants on microbial food web structure and ecosystem production.     

A modelling investigation of the distribution of stratification and phytoplankton abundance in the Irish Sea

Thermal stratification and phytoplankton abundance are modelledon a 5 km grid covering the Irish Sea. The water column is approximatedby three layers. The top layer is uniformly mixed by wind stirringand the bottom by tidal energy, while linear gradients can occurin the middle layer. The model is forced with hourly meteorologicaldata and mean tidal energies. Primary production is representedby a model with a single nutrient and a single phytoplanktonpopulation. The results from the model show good agreement withdata collected on a Ministry of Agriculture, Fisheries and Food(MAFF) cruise in May 1992 and with historical data. When advectionis included, driven by depth-averaged currents, the surfacetemperature patterns are improved but bottom temperatures indeep water are raised and high concentrations of chlorophyllare carried offshore from coastal regions. This indicates alimitation of using depth-averaged currents and a need to accountfor differences in phytoplankton species composition in coastaland offshore waters. Calculations demonstrate the importanceof salinity variations to stratification and phytoplankton growth.Smoothing the wind mixing energy has the effect of delayingthe onset of the spring bloom in areas where wind mixing issignificant. Removing the diurnal cycle of solar heating alsodelays the spring bloom. The chlorophyll gradient in the middlelayer has a large impact on the response of the model to short-termvariability in the meteorological forcings.     

九寨沟两种常见藓类植物对模拟氮沉降的生理响应

为探讨氮沉降对九寨沟藓类植物的影响，该研究以当地优势藓类植物锦丝藓(Actinothuidium hookeri)和塔藓(Hylocomium splendens)为对象，以NH₄NO₃为氮源，设置对照(0 kg N·hm^-2·a^-1)、低浓度(20 kg N·hm^-2·a^-1)、高浓度(50 kg N·hm^-2·a^-1)3种处理，开展为期6个月的氮沉降模拟实验。结果表明：(1)氮沉降处理导致两种藓类植物的活性氧、丙二醛、叶绿素、脯氨酸和可溶性蛋白含量显著增加，同时锦丝藓过氧化氢酶、过氧化物酶、超氧化物歧化酶、抗坏血酸过氧化物酶活性增加。(2)对于生长旺期和生长末期的塔藓，氮沉降导致其过氧化物酶、过氧化氢酶、抗坏血酸过氧化物酶活性降低。(3)锦丝藓的综合隶属函数值随氮沉降浓度增大而增加，在生长旺期和生长末期，塔藓综合隶属函数值对氮沉降的响应存在差异。综上认为，两种藓类植物对氮沉降处理的生理响应存在差异，高浓度氮沉...     

彩叶草对模拟干旱胁迫的生理响应

采用盆栽试验,对彩叶草进行PEG-6000浓度为0(对照)、5%、10%、15%、20%(W/V)模拟干旱胁迫,研究在干旱胁迫下彩叶草的生长、渗透调节能力及抗氧化酶活性的变化。结果表明:与对照相比,随着PEG-6000浓度的增加,鲜质量、干质量、含水量、水势、根系脱氢酶活性、无机离子含量包括K+、Na+、Ca2+、Mg2+等均呈下降趋势;NO-3含量呈先下降后上升趋势;硝酸还原酶活性、可溶性蛋白含量、可溶性糖含量、超氧化物歧化酶活性均呈先上升后下降趋势;脯氨酸含量、游离氨基酸含量、过氧化物酶活性、过氧化氢酶活性、超氧阴离子(O-2·)产生速率、质膜透性则呈上升趋势。因此,模拟干旱胁迫对彩叶草生长有抑制作用,且随着PEG-6000浓度增加,其生长受抑制和水分胁迫程度加重;模拟干旱胁迫下,彩叶草不积累K+、Na+、Ca2+、Mg2+和NO-3等无机离子进行渗透调节,而积累脯氨酸、可溶性蛋白、可溶性糖、游离氨基酸等有机小分子物质进行渗透调节,但这4种小分子物质增加幅度不尽相同;轻度模拟干旱胁迫虽增强彩叶草抗氧化酶活性,但仍表现轻度的氧化伤害;重度模拟干旱胁迫加重彩叶草氧化伤害。研究结果可为彩叶草耐旱生理机制的研究积累资料,也为其节水型栽植和养护提供依据。     

Copepod abundance in the western Irish Sea: relationship to physical regime, phytoplankton production and standing stock

The distinct patterns of stratification in the North Channeland stratified region of the western Irish Sea influence theseasonal abundance of phytoplankton. The 3–4 month productionseason in the stratified region was characterized by productionand biomass peaks in the spring (up to 2378 mg C m² day^–1and 178.4 mg chlorophyll m^–2) and autumn (up to 1280 mgC m^–2 day^–1 and 101.9 mg chlorophyll m^–2).Phytoplankton in the North Channel exhibited a short, late productionseason with a single summer (June/July) peak in production (4483mg Cm^–2 day^–1) and biomass (–160.6 mg chlorophyllm^–2). These differences have little influence on copepoddynamics. Both regions supported recurrent annual cycles ofcopepod abundance with similar seasonal maxima (182.8–241.810³ind. m^–2) and dominant species (Pseudocalanus elongatusand Acartia clausi). Specific rates of population increase inthe spring were 0.071 and 0.048 day¹ for the North Channel andstratified region, respectively. Increased copepod abundancein the stratified region coincided with the spring bloom, andwas significantly correlated with chlorophyll standing stock.Increased copepod abundance preceded the summer production peakin the North Channel. This increase was not correlated withchlorophyll standing crop, suggesting that a food resource otherthan phytoplankton may be responsible for the onset of copepodproduction prior to the spring bloom. Hetero-trophic microplanktonas an alternative food source, and advection of copepods fromthe stratified region, are proposed as possible explanationsfor copepod abundance increasing in advance of the summer peakin primary production.     

Chlorophyll standing crop and phytoplankton production in the western Irish Sea during 1992 and 1993

Chlorophyll standing crop and phytoplankton production werestudied in the western Irish Sea over a 21 month period during1992 and 1993. For both years, the start of the production seasonwas first observed in Dundalk Bay and occurred progressivelylater in more northerly coastal and offshore waters. Standingcrop and production exhibited marked spatial heterogeneity with12.5- to 19-fold differences in crop observed over distancesof 20–30 km. Distinct regional differences in the lengthof the production season were apparent. The longest season,6–7 months with a production of 194 g C m^–2, occurredin Dundalk Bay. The season lasted 3–4 months in the summerstratified region with a production of 140 g C m^–2. Northerly,offshore mixed waters and coastal waters of Northern Irelandsupported a short (2–3 months) season and production of194 and 140 g C m^–2, respectively. The similarity in seasonalproduction between Dundalk Bay and coastal waters of NorthernIreland, and between the summer stratified and northern mixedregions, is attributed to the intensity of production duringthe summer. Between 59 and 79% of seasonal production in thenorthern mixed region and coastal waters of Northern Irelandtook place during June and July, compared to 29–40% inDundalk Bay and the summer stratified region. Lower summer productionin the latter two may be due to nutrient limitation and thishas implications for the sensitivity of these two regions toanthropogenic nutrient enrichment.     

Physiological responses of Sargasso Sea picoplankton to nanomolar nitrate perturbations

A study was conducted in July 1989 at three stations in thenorthern Sargasso Sea, where picoplankton (<1 µm)provided approximately half of the standing crop of chlorophyll.Temporal changes in the position of the nitracline at a singlelocation indicated that the vertical supply of nitrate was notat ‘steady-state’ and phytoplankton distributionstracked the nitracline. Our main experimental objective wasto examine the short-term effects of ecologically significantnitrate perturbations (+20 and +100 nM) on the physiologyof <1 µm communities growing at low (nanomolar)ambient nitrate concentrations. A chemiluminescent nitrate methodwas used to measure the time course (up to 4 h) of nitratedisappearance at in situ irradiance, in parallel with measurementsof photosynthetic ¹⁴CO₂ assimilation. Picoplankton growing at<60 nM nitrate rapidly responded to nanomolar nitratesupplements with luxury consumption and enhanced photosynthesisin proportion to their ambient nitrate environment. Light-saturatedSynechococcus populations from the most nitrate-depleted waters(13 nM) had doubled their cellular rate of photosynthesisafter 4 h, in response to a 20 nM nitrate pulse.     

Peculiarities of the copper toxic effect on the phytoplankton communities of the Baltic Sea

The ecotoxicological investigations on the resistance of various phytoplankton communities were performed in the Southern Baltic Sea during summer and winter periods. The primary production rate was studied for different Cu concentrations. The enhancement of the toxic effect as well as total biomass and productivity increase were observed for phytocenosis. Total algae biomass on most of the stations increased accordingly to the dinoflagellate growth. Both dinoflagellate impact and relative copper toxic effect declined in the Gdansk Bay opposite to the high total algae biomass observed. We assume the dinoflagellate abundance is linked with the sensitivity of the phytoplankton cells to copper.     

台湾海峡上升流区浮游植物对营养盐添加的响应

2006年6月在台湾海峡近岸上升流区通过表层水体营养盐添加的现场培养实验,研究该海区营养盐限制情况及其浮游植物水华产生的主要影响因素.对营养盐,叶绿素a浓度和浮游植物细胞丰度进行了测定,结果表明,实验中不存在明显的硅限制;氮磷营养盐均存在明显的限制,且氮限制情况更为严重.营养盐添加后,冰河拟星杆藻(Asterionellopsis glacialis)等硅藻迅速生长成为优势藻种,其对氮磷的利用机制有所不同.对氮营养盐采取吸收后迅速同化利用,相较于硝酸盐的补充,氨氮补充条件下优势硅藻更易迅速生长并迅速死亡;对磷营养盐的利用则由于体内磷库的存在,采用迅速吸收后贮存在体内慢慢消耗的利用机制.氮营养盐的补充是上升流期间浮游植物水华产生的主要因素.     

4种阔叶幼苗对PEG模拟干旱的生理响应

研究了PEG模拟干旱胁迫环境下的火力楠(Michelia macclurel)、尾叶桉(Eucalyptus urophylla)、枫香(Liquidambar formosana)、荷木(Schima superba)幼苗的生理变化。结果表明,干旱胁迫下,4种幼苗叶片的相对含水量小于对照,其中,尾叶桉和枫香下降明显;不同干旱胁迫条件下,4种树种幼苗叶片的相对电导率均显著大于对照,其中尾叶桉和枫香上升幅度大;干旱胁迫下的火力楠和荷木幼苗叶片的脯氨酸含量呈现波动,尾叶桉和枫香幼苗则显著大于对照;不同干旱胁迫时间下的幼苗叶片的叶绿素含量小幅波动;4个树种幼苗的过氧岐化酶(SOD)活性随胁迫时间增加而呈现先升后降的趋势,其中火力楠和荷木的幼苗的SOD活性持续维持在较高水平;荷木叶片的丙二醛(MDA)含量先升后降,最后和对照水平相近,其余幼苗的MDA含量均大于对照;干旱胁迫下4种幼苗叶片的可溶性糖含量增加幅度较大。主成分分析表明,4种幼苗的抗旱能力排序为荷木>火力楠>尾叶桉>枫香。     

低温胁迫后苔藓植物对模拟氮沉降条件的生理响应

2008年初受强寒潮的影响, 中国华南大部分地区出现持续的异常低温。该文研究了华南地区常见的3种苔藓植物 ——刺边小金发藓拟刺亚种(Pogonatum cirratum subsp. fuscatum)、大灰藓(Hypnum plumaeforme)和石地钱(Reboulia hemisphaerica), 在人工模拟氮(N)沉降两年并经历2008年初异常低温气候后的生理响应变化, 并与2007年正常气候情况下人工加N一年后的结果进行比较, 分析苔藓植物的生长与N沉降之间的关系, 并探讨N沉降对低温胁迫后苔藓植物的补偿生长的影响。结果显示: 2008年3种苔藓植物的净光合速率和淀粉含量在加N量为0-60 kg N·hm ^-2·a^-1的范围内均随着N浓度的上升而下降, 总N含量在加N量处于0-40 kg N·hm^-2·a^-1的范围内随着N浓度的上升而上升, 至60 kg N·hm^-2·a^-1时不再上升, 甚至有所下降。2008年, 3种苔藓植物大多数碳氮代谢指标在对照及低N条件下与2007年加N 1年且在正常气候时同种N处理时相比均有不同程度的上升, 但上升幅度与加N浓度成反比, 至中高N条件时两者常较接近, 显示苔藓植物在经历低温胁迫后会出现超补偿效应, 但是在N沉降升高的条件下, 补偿生长能力下降。     

Regional differences in stratification and its effect on phytoplankton production and biomass in the northwestern Irish Sea

Differences in tidal mixing result in the formation of offshoremixed and stratified regions in the NW Irish Sea during springand summer. Stratification resulted from vertical gradientsin temperature, although vertical gradients in salinity wereimportant during the early stages of stratification. The northerncoastal and offshore mixed regions were characterized and distinguishedfrom the southern coastal and summer stratified regions by thepresence of more saline, cool near-surface water and incompletedepletion of dissolved inorganic nutrients. Distinct regionaldifferences in the production season of phytoplankton were observed.This lasted 6 months with a seasonal production of 155 g C m^–2in the southern coastal region. A shorter season, 4 and 2 months,and lower production of 101 and 96 g C m^–2 occurred inthe summer stratified and northern mixed regions, respectively.The southern mixed region supported the shortest season (<2months) and lowest production (66 g C m^–2). It is estimatedthat a daily light exposure of {small tilde}200 Wh m^–2is required for the onset of the production season and it isconduded that the subsurface light climate as a function ofsolar radiation and surface mixed layer depth, rather than nutrientavailability, controls its duration. The existence of a shortlate production season in the offshore regions is confirmedand this contrast with the season in the North Sea is attributedto differences in the subsurface light climate. Sustained productionin the southern coastal region may play an important role infish recruitment and offset any effect of the short late offshoreproduction season.     

Physiological responses of free-swimming adult coho salmon to simulated predator and fisheries encounters

The responses of free-swimming adult coho salmon (Oncorhynchus kisutch) to simulated predator and fisheries encounters were assessed by monitoring heart rate (f(H)) with implanted data loggers and periodically taking caudal blood samples. A 10- or 30-min corralling treatment was conducted to simulate conspecifics being cornered by a predator or corralled by fisheries gear without physical contact. Corralling rapidly doubled f(H) from ～31 beats min(-1) to a maximum of ～60 beats min(-1), regardless of the duration of the corralling. However, recovery of f(H) to precorralling levels was significantly faster after the 10-min corralling (7.6 h) than after the 30-min corralling (11.5 h). An exhaustive-exercise treatment (chasing for 3 min, with physical contact) to simulate a predator chasing a fish to exhaustion or a fish becoming exhausted after encountering fisheries gear resulted in increased f(H) (to 60 beats min(-1)), plasma lactate, glucose, sodium, osmolality, and cortisol (males only) and a significant decrease in mean corpuscular hemoglobin concentration. Recovery of f(H) and most blood variables was complete about 16 h after exhaustive exercise and handling. The results illustrate a clear relationship between the intensity of exercise and the duration required for recovery of f(H). Changes in f(H) were significantly correlated with those in plasma lactate, chloride, and sodium at 1 h after the exercise treatment protocols. Thus, measurements of f(H) may provide an accurate indication of the general physiological response of salmonids to exhaustive exercise in the natural environment.     

Microzooplankton grazing of phytoplankton in a tropical upwelling region

We measured grazing by herbivorous zooplankton (<200 μm fraction) in coastal and slope regions of the South Brazil Bight. Using the dilution technique, we performed nine experiments during the austral summer, when nutrient-rich South Atlantic Central Water is present on the shelf, and five during winter. These experiments provide the first estimates of microzooplankton grazing in the western South Atlantic Ocean. Model II regression showed a strong relationship between phytoplankton intrinsic growth rates and grazing, with a slope of 0.64 (±0.28; 95% confidence interval) indicating that microzooplankton grazing could account for the majority of phytoplankton mortality. Both phytoplankton growth and microzooplankton grazing were higher during the summer upwelling season, compared to winter. For the two experiments that were conducted in oligotrophic slope water, grazing accounted for >80% of phytoplankton production. A comparison of incubations with and without added inorganic nutrients showed no consistent stimulation of phytoplankton growth (slope of enriched versus unenriched treatments not significantly different from 1). Estimates from microscopic counts of heterotrophic organisms >10 μm indicated that copepod nauplii comprised the largest share of the microzooplankton biomass (mean 62.4 ± 5.8% SE). Grazing estimates were not correlated with microzooplankton biomass, whether or not nauplii were included, suggesting that most of the grazing was done by nano-sized zooplankton. Electronic Supplementary Material Electronic supplementary material is available in the online version of this article at and is accessible for authorized users. Handling editor: S. Wellekens     

Phytogeography of upwelling areas in the Arabian Sea

Aim The identification of the marine plant communities of two islands from different upwelling areas of the Arabian Sea, with a similar diversity in biotopes. A comparison of the species composition of these macroalgal communities and their biogeographical affinities within the Indian Ocean should give insight into the biogeographical position of the Arabian Sea within the larger Indian Ocean. The incorporation of environmental parameters in the analysis is instructive in understanding their importance in shaping the diverse marine assemblages of the Arabian Sea. Location Arabian Sea: (1) the Socotra Archipelago (Yemen; 12.47° N, 53.87° E) in the Somali upwelling area, (2) Masirah Island (Oman; 20.42° N, 58.79° E) in the upwelling area of the southern Arabian Peninsula. Methods The marine flora of different biotopes around both islands were examined by means of qualitative assessments. Ordination analysis [detrended correspondence analysis (DCA)] was used to identify the different plant communities and to correlate these with environmental parameters. The species composition of the identified communities were compared (tripartite similarity index) and their biogeographical affinity with nations bordering the Indian Ocean was determined. Indicator species analyses were performed to identify the characteristic species of the different plant communities and their biotopes. Results The DCA analysis shows a clustering of sites (plant communities) corresponding with their geographical position, linked in turn to the prevailing environmental conditions of the different coastal areas. The combined interpretation of the ordination, similarity and biogeographical analyses results in the aggregation of similar plant communities of both upwelling areas into four biotopes. Main conclusions The north coast communities of Socotra and the west coast communities of Masirah can be grouped into three biotopes related to the degree of exposure (to upwelling) and sedimentation. These biotopes are typified by indicator species, characteristic for specific substrata, and have a high biogeographical affinity with the East African coast. The plant communities of Socotra's south coast and Masirah's east coast constitute a fourth biotope, being diverse and species rich, typified by a large proportion of red macroalgae including the characteristic species of the unique Arabian Sea flora. This biotope has a pronounced biogeographical affinity with distant regions (disjunctly distributed taxa) as South Africa's East Coast and Western Australia. Within the different biotopes, the communities of Masirah are more divergent from an East African flora in comparison to Socotra, the latter being a stepping stone between the East African and Arabian Sea flora.