Comparison of swimming speed and photophysiological responses to different external conditions among three Karenia brevis strains

Behavior, growth, and production are integral in the life history of Karenia brevis, an autotrophic, dinoflagellate HAB species, and are important variables in modeling blooms in the Gulf of Mexico. This study compares swimming speeds, growth rates, and photosynthetic responses of recent isolates of K. brevis (specifically the Apalachicola – APA, Manasota – MAN, and Jacksonville – JAX strains) over a range of light intensities and temperatures. Strain swimming speeds were similar and remained fairly constant from 17 to 30 °C, but decreased markedly at 13 °C. Photosynthetic responses of the strains to different acclimated temperatures had opposite trends with APA exhibiting higher electron transport rates (ETR) at higher temperatures and MAN exhibiting higher ETR at lower temperatures. In the light experiments, the cells’ internal physiological state (represented by photosynthetic yield, ETR, and neutral lipid reserves) and swimming capabilities were examined in the dark after 6 h incubations in the radial photosynthetron. For all strains, at initial incubation light intensities swimming speed decreased and ETR increased. As incubation light intensities increased, ETR decreased and swimming speed increased. At the highest incubation light intensities, ETR and swimming speed decreased. Neutral lipids followed a pattern similar to ETR, only lipids peaked after ETR at a light intensity that corresponded to the increase in swimming speed. The results suggest that cells may partition energy selectively depending on the needs of the cell. Information was combined to characterize a generalized species response to light and temperature ranges.     

Seasonal analysis of Daphnia pulicaria swimming behavior

Our study documents individual swimming behavior of Daphnia pulicaria over a yearly cycle in a temperate lake. We collected D. pulicaria, a common freshwater zooplankton, from Lake Mendota on 10 dates between July 1994 and June 1995 from two depths, 2 m and 10 m. The Daphnia were rushed to the laboratory and video-taped as they swam in lake water under lake-ambient temperature and light conditions. Five-second swimming tracks of individual Daphnia were filmed and digitized using a motion analysis system. We measured average turning angle, swimming speed and sinking rate for each track. D. pulicaria swimming behavior varied over the annual cycle. We found significant differences in turning angle between depths and among months. Sinking rate and swimming speed were significantly different among months but not depths. Sinking rate and swimming speed were not significantly correlated with water temperature. Our results were contrary to Stokes' Law predictions, in that D. pulicaria had the slowest sinking speed in June, not in the winter when water temperatures were lowest and viscosity was highest. Body length was significantly correlated with all three swimming variables. We also studied swimming behavior in clonal populations of D. pulicaria in different concentrations of the alga, Chlamydomonas reinhardtii. D. pulicaria did not change swimming speed, turning angle or sinking rate over a range of food concentrations. Finally, swimming behavior in a D. pulicaria clone, tested at two temperatures in the laboratory, confirmed the results from our seasonal study; Daphnia did not sink as predicted by changes in viscosity.     

ABUNDANCE OF A NOVEL DINOFLAGELLATE PHYLOTYPE IN THE ROSS SEA,ANTARCTICA1

The biodiversity of protistan assemblages present in microhabitats of the Ross Sea, Antarctica, was examined using molecular biological methods to obtain a greater understanding of the genetic diversity present. Sequencing of 18S clone libraries indicated genetically diverse collections of organisms in the water column, ice, and meltwater layer (slush), but a single small subunit ribosomal DNA (srDNA) sequence type dominated clone libraries (>30%) from seawater and slush samples taken within the ice pack of this ecosystem. The BLAST searches indicated that this dominant clone was derived from a dinoflagellate, and that it shared sequence similarity (97.6%–98.3%) with both Karenia and Karlodinium species. Phylogenetic analyses based on small subunit ribosomal gene sequences supported its placement as a sister group to these taxa, and suggested that it represented a novel genus. The dinoflagellate was successfully recovered in culture, and morphological analyses have shown that it contains chloroplasts, is gymnodinoid, appears not to have thecal plates, and has an apical groove and sulcal structure that confirm its placement as a relative of the Karenia/Karlodinium group. The abundance of this phylotype in natural samples was confirmed by quantitative PCR analyses of water and slush communities, and suggests that this dinoflagellate can be a major constituent of the protistan assemblages of some Antarctic microhabitats of the Ross Sea.     

BACTERIA‐INDUCED MOTILITY REDUCTION IN LINGULODINIUM POLYEDRUM (DINOPHYCEAE)1

Biotic factors that affect phytoplankton physiology and behavior are not well characterized but probably play a crucial role in regulating their population dynamics in nature. We document evidence that some marine bacteria can decrease the swimming speed of motile phytoplankton through the release of putative protease(s). Using the dinoflagellate Lingulodinium polyedrum (F. Stein) J. D. Dodge as a model system, we showed that the motility‐reducing components of bacterial‐algal cocultures were mostly heat labile, were of high molecular weight (>50 kDa), and could be partially neutralized by incubations with protease inhibitors. We further showed that additions of the purified protease pronase E decreased dinoflagellate swimming speed in a concentration‐dependent manner. We propose that motility can be used as a marker for dinoflagellate stress or general unhealthy status due to proteolytic bacteria, among other factors.     

Fish swimming stride by stride: speed limits and endurance

Summary Steadily swimming fish show a species-specific stride length and tail tip amplitude. These are constant over the entire speed range if expressed as a fraction of the body length. The speed of a fish equals the stride length times the tail beat frequency. We describe how maximum tail beat frequencies, and hence maximum swimming speeds, are related to temperature and body length.Maximum sustained swimming speeds, endurance during swimming at higher speeds, and maximum burst velocities of 27 species are compared. The rate of decline of endurance with increasing speed is either gradual or steep, with only a few cases in between Steady swimmers show the steepest decline.The published effects of temperature on endurance are not consistent.The effect of body size on the endurance curve could be investigated for two species. The maximum sustained speed decreases with increasing length, and the slope of the endurance curves steepens with increasing length with the same factor in both species. The maximum burst speed is 10 Ls^-1 on average.     

Behavioral responses of the coastal copepod Acartia hudsonica (Pinhey) to simulated dinoflagellate bioluminescence

Light pulses were used to mimic dinoflagellate bioluminescence and test its effects on the swimming behavior of Acartia hudsonica (Pinhey). The horizontal swimming patterns of the copepod were tracked and described using a video-computer system. Single flashes of light of 60 ms duration, with a wavelength of peak emission of 475 nm and an intensity of 2 μE · m^?2 · s^?1 caused a “startle” response consisting of a short burst of high speed swimming. A series of these flashes repeated every 5 s resulted in higher average swimming speed, more swimming speed bursts, and straighter paths. These behavioral changes are similar to those previously found for A. hudsonica in the presence of bioluminescent dinoflagellates. The effects of altering the intensity, duration, and color of the simulated dinoflagellate flash were also tested. Our results support the hypothesis that dinoflagellate bioluminescence is a highly evolved adaptation for repelling nocturnal grazers.     

亚成体巨须裂腹鱼游泳能力及活动代谢研究

以野生雅鲁藏布江巨须裂腹鱼(Schizothorax macropogon)为对象,通过自制的鱼类游泳实验装置,测定了4个温度(5、10、15和18℃)梯度下亚成体巨须裂腹鱼的临界游泳速度(Ucrit)及流速变化对耗氧率的影响,并通过摄像记录分析了不同游泳速度下的游泳行为。野生亚成体巨须裂腹鱼的临界游速随着温度的变化呈近似线性的递增趋势(P<0.001),4个温度下的绝对临界游速(Ucrit-a)分别为(0.88±0.07)、(1.09±0.07)、(1.24±0.15)和(1.49±0.15)m/s;若以单位时间内游过的体长倍数(BL/s)表示,相对临界游速(Ucrit-r)分别为(3.96±0.21)、(4.4±0.16)、(4.9±0.18)和(5.35±0.14)BL/s。根据不同温度及流速下耗氧率的变化情况,采用非线性拟合得到了4个温度梯度下耗氧率与游泳速度关系的幂函数模型(P<0.05)。模型表明耗氧率随游泳速度的增大而增加,且温度越高耗氧率随游泳速度的变化越显著。4个温度下的速度指数分别为2.4、2.6,2.8及3.1,表明有氧运动的效率随温度升高有所降低。在自然水温条件下(5—9℃),摆尾频率(TBF)与流速的关系呈线性正相关(P<0.001),而运动步长(Ls)的变化与流速没有显著关系,出现由高至低再升高的三个阶段。录像分析表明在流速逐渐增加的过程中,巨须裂腹鱼采用了三种不同的游泳方式,以实现降低能量消耗的目的。研究可为鱼道等过鱼设施的设计提供参考,对数量日益减少的巨须裂腹鱼保护具有较大的意义。     

Swimming performance and metabolic rate of three tropical fishes in relation to temperature

Oxygen consumption was measured for three tropical fishes,Exodon paradoxus, Leporinus fasciatus andLabeo erythrurus in relation to swimming speed and temperature. For each species the logarithm of oxygen consumption (mg 0₂ · g^–1 · h^–1) increased linearly with relative swimming speed (1 · s^–1) with the value of the regression coefficients varying inversely with temperature. Active metabolism and critical swimming speed ofE. paradoxus andL. fasciatus increased with temperature to a maximum at 30 and 35° C respectively. Basal metabolic rates ofE. paradoxus andL. fasciatus increased with temperature. Metabolic rates and critical swimming speed of the three fishes studied were consistent with values for polar, temperate and other tropical species over their respective thermal ranges of tolerance. Tropical fishes have lowered their metabolism and swimming performance from that expected for many temperate species at the same temperature.     

Dinoflagellate-targeted PCR reveals highly abundant and diverse communities of parasitic dinoflagellates in and near Zhubi Reef,South China Sea

While diversity of symbiodiniacean dinoflagellates has been a focus of coral reef ecological research, information on the diversity of planktonic dinoflagellates in reef ecosystems remains limited. We used dinoflagellate-targeted PCR to investigate dinoflagellate diversity for a coral reef ecosystem. In the summer of 2007, plankton samples were collected from a lagoon, atoll, and open sea area of Zhubi Reef in the Nansha Islands, South China Sea. Sequencing of dinoflagellate-specific SSU rDNA clone libraries from samples in each of these habitats revealed high diversity and numerous novel dinoflagellate lineages. Gymnodiniales were most abundantly represented in all three water areas. Lagoon assemblages were co-dominated by Syndiniales and Gonyaulacales, the atoll by Gonyaulacales and Peridiniales, and the open sea by Syndiniales and Prorocentrales taxa. Species in the Syndiniales (group II) genus Amoebophrya were represented by eight new sequences and 13 previously described clades and were dominated by species reported to infect Gymnodiniales, Gonyaulacales, Peridiniales, and Prorocentrales taxa. And Amoebophrya were particularly abundant and diverse in the lagoon. Our results suggest that Amoebophrya probably play an important role in regulating dynamics of dinoflagellate assemblages in the Zhubi Reef coral ecosystem. In contrast, the few symbiodiniacean taxa detected occurred only in the open sea, suggesting planktonic aposymbiotic Symbiodiniaceae rarely occur in the reef ecosystem. We demonstrate the usefulness of a dinoflagellate-specific molecular technique for profiling dinoflagellate communities, and uncover diversity and the potential importance of parasitic lineages in a coral reef ecosystem.

     

Water temperature, light intensity and zooplankton density and the feeding activity of juvenile brook charr (Salvelinus fontinalis)

SUMMARY 1. The objective of this study was to evaluate the effects of zooplankton biomass (as a measure of density), fish biomass, light intensity and water temperature on the attack rate and swimming characteristics (i.e. swimming speed and angle of turn) of juvenile (1+) brook charr (Salvelinus fontinalis) in field enclosures. We used a portable underwater camera system in a series of pelagic enclosures to quantify the feeding behaviour of brook charr over a gradient of natural conditions. 2. In simple linear or non‐linear regression models we found (i) that attack rate and angle of turn were positively related to water temperature, (ii) that attack rate and swimming speed were positively related to zooplankton biomass and light intensity and (iii) that attack rate was positively related to swimming speed. In multiple regression models, fish biomass, light intensity and variance of the angle of turn accounted for 87% of the variation in attack rate. Light intensity and water temperature accounted for 86% of the variation in swimming speed. Fish gut fullness and attack rate accounted for 83% of the variation in the variance of the angle of turn executed by fish. 3. The increase in the number of attacks as zooplankton biomass increases conforms to the general positive functional response observed in other fish species. Our results also support the hypothesis that swimming speed increases with prey biomass. We did not observe a plateau in attack rate as zooplankton biomass increased. As our experiments were performed under natural biotic and abiotic conditions, factors other than zooplankton biomass might affect or limit this response, such as water temperature and light intensity. 4. Because zooplankton biomass was correlated with water temperature and light intensity, it was not possible to evaluate the independent contribution of these factors on the attack rate and swimming characteristics (swimming speeds and angle of turn) of brook charr. However, this study highlighted the impact of these factors on the feeding behaviour of juvenile brook charr when feeding in the pelagic habitat under natural conditions, and their importance in future models of optimal foraging and fish habitat quality.     

Maximum sustainable swimming speeds of late-stage larvae of nine species of reef fishes

We examined the maximum sustainable swimming speed of late-stage larvae of nine species of tropical reef fishes from around Lizard Island, Great Barrier Reef, Australia. Larvae were captured in light traps and were swum in flumes at different experimental swimming speeds (of 5 cm s⁻¹ intervals) continuously for 24 h. Logistic regression was used to determine the speed at which 90% of larvae were able to maintain swimming, and this was used to indicate the maximum sustainable swimming speed for each species. Maximum sustainable swimming speeds varied among the species examined, with the lethrinid maintaining the fastest sustainable swimming speed (24 cm s⁻¹), followed by the Pomacentridae (10-20 cm s⁻¹) and the Apogonidae (8-12 cm s⁻¹). U-crit (maximum speed) explained 64% of the variation in sustainable speed among species, whereas total length only explained 33% of the variation in sustained swimming. A regression fitted across species suggests that 50% U-crit is a good approximation of the speed able to be maintained by these larvae for 24 h. A model based on a cubic relationship between sustained swimming time and speed was found to be more successful than either length or U-crit as a method of estimating sustainable swimming speed for most of the species examined. Overall, we found that swimming speed is an important factor when considering the potential for active swimming behaviour to influence dispersal patterns, recruitment success and levels of self-recruitment in reef fish larvae and needs to be carefully considered in models of larval dispersal.     

Swimming behavior of Daphnia: its role in determining predation risk

Individual swimming behavior of zooplankton can play an importantrole in determining how planktivorous fishselect their prey.Although several studies have documented the effect of preysize, contrast or degree of pigmentation, escape ability, encounterrate and abundance in determining predation risk, the importanceof individual behavior has received relatively little attentionby aquatic ecologists. Recent advances in the technology ofvideo recording and computer analysis of motion have allowedus to collect digitized three-dimensional videorecords of free-swimmingzooplankton such as Daphnia. We found that Daphnia clones, includingthose within a single species, exhibit a wide range of swimmingbehaviors as measured by swimming speed. The individual behaviorof a species cannot be adequately described by looking at oneclone. We alsoshow that different behavior observed in liveDaphnia can play an important role in determining attractivenessto visual predators. Given a choice between two clones of equalsize and visibility contrast, fish selected indi viduals fromthe faster swimming clone. Our results suggest that currentmodels of prey selection would be improved by the incorporationof individual swimming behavior because it is an important factordetermining overall prey visibility.     

ENCYSTMENT OF THE DINOFLAGELLATE GYRODINIUM UNCATENUM: TEMPERATURE AND NUTRIENT EFFECTS1

Sexual reproduction and encystment of the marine dinoflagellate Gyrodinium uncatenum Hulburt were induced in nitrogen and phosphorus-limited batch cultures. Sexuality did not occur under nutrient-replete conditions even when growth rate was reduced by non-optimal temperatures. Growth was optimal over a broader temperature range than encystment and virtually no cysts were produced at some low and high temperatures where growth occurred. Most cells initiated sexuality as intracellular pools of each limiting nutrient reached minimum or subsistence levels as much as four days after extracellular nutrients were exhausted. High nitrogen cell quotas during the phosphorus experiment indicate that sexuality was induced by a shortage of phosphorus and not by an indirect effect on nitrogen uptake. Total cyst yield corresponded to successful encystment of 9–13% of the motile populations, yet 60–85% of the plateau-phase motile cells were planozygotes (swimming zygotes formed from fusing gametes). Batch culture studies monitoring total cyst yield may thus seriously underestimate the extent of sexuality. More importantly, the number of cysts produced in a dinoflagellate population may be significantly reduced by environmental factors acting on the cells after sexual induction and fusion.     

Evolution of fast development of planktonic embryos to early swimming

Planktonic embryos of marine animals swim at an early stage and age. Although natural selection has apparently favored rapid development of structures for swimming, taxa have not converged on the same, minimal time from first cell division to first swimming. Comparisons of 34 species with planktonic embryos in 10 phyla revealed factors that account for variation in time to swimming. Time to first swimming correlated significantly with time from first to second cleavage (first cell cycle) in analyses of all embryos sampled and separately within the Spiralia and Echinodermata. Time to first swimming also correlated significantly with egg diameter in some clades, but not in all. Correlations between egg diameter and cell cycle duration were low except for the three species of Urochordata. Development to a feeding or nonfeeding larva did not affect time to first swimming beyond effects attributable to egg size. Time to first swimming did not correlate with type of locomotion developed (uniciliated cells, multiciliated cells, or muscle). Nonetheless, differences in locomotion are associated with changes in cell cycle durations prior to swimming. The ratios of time to first swimming and time for first cell cycle suggests that allocation of time to multiplication of cells versus differentiation of cells is resolved differently in species with different types of locomotion.     

Form and Function of the Dinoflagellate Transverse Flagellum1

A reexamination of the dinoflagellate transverse flagellum in relation to swimming in more than 50 species, using a television recording system, has revealed the following new facts: the flagellar beat always proceeds counterclockwise when seen from the cell apex; the cell always rotates in the direction of the flagellar beat, and fluid is propelled in the opposite direction. These observations can be explained by the actions of flagellar mastigonemes not included in previous models. The shape of the flagellar wave is not isotropic. New explanations are offered for other morphological features of the cell as they relate to swimming.     

Patterns in early embryonic motility: effects of size and environmental temperature on vertical velocities of sinking and swimming echinoid blastulae

Early embryonic swimming is widespread among marine invertebrates, but quantitative information about swimming behaviors is scarce. Swimming may affect encounters with predators, positioning in the water column, and nutrient absorption. Measured rates and patterns of swimming and sinking for blastulae of four eastern Pacific echinoid species show that sinking speeds equal or exceed swimming speeds. Swimming speed scaled negatively with embryo size, though sinking speed did not scale with size. Analysis of swimming paths of Strongylocentrotus franciscanus revealed a temperature dependency in swimming pattern that affected speed of upward movement. Sinking speeds were significantly greater at 10 degrees C than at 14 degrees C for blastulae of all four species examined. In Dendraster excentricus, killing the blastulae annulled this temperature effect, indicating an active density regulation by these embryos. Finally, measurements of particle velocities around sinking and swimming D. excentricus blastulae show that swimming creates a more localized disturbance than sinking. Embryonic swimming may therefore decrease rather than increase encounters with pelagic predators. Results from subsequent experiments in which embryos were reared in low-oxygen environments suggest that any oxygen-absorption advantages of swimming have little, if any, effect on the development of D. excentricus embryos.     

不同程度的尾损伤对镇海林蛙(Rana zhenhaiensis)蝌蚪游泳速度的影响

无尾两栖类蝌蚪的尾巴通过产生强大的游泳速度在反捕食中起到了重要的作用。以镇海林蛙(Rana zhenhaiensis)蝌蚪为实验动物来评估断尾的运动代价。以74尾具有完整尾蝌蚪作为实验组,通过截去不同尾长片段,人为分成轻微尾损伤组(30%)和严重尾损伤组(30%)并测定两组蝌蚪在断尾前后的游泳速度。以16尾完整尾蝌蚪作为对照组在实验组断尾前后同时进行游泳速度的测定。实验结果显示断尾影响蝌蚪的游泳速度,但仅在尾损伤程度达到尾长的30%以上时才产生不利的影响。这表明轻微尾损伤并不对镇海林蛙蝌蚪的游泳速度产生严重影响。在断尾前后实验组蝌蚪的游泳速度均与尾长呈正相关。在相同尾长状态下,尾损伤蝌蚪的相对游泳速度明显快于完整尾蝌蚪。因此,尾损伤的镇海林蛙蝌蚪有可能通过改变尾和身体的摆动频次等方式在断尾后对游泳速度进行了一定的补偿。尾损伤在野外频繁发生于蝌蚪的尾远端,据此推测镇海林蛙蝌蚪在自然条件下的尾损伤并不会产生严重运动代价。     

Swimming metabolism and temperature in juvenile walleye,Stizostedion vitreum vitreum

Synopsis Oxygen consumption of juvenile walleye increased between 5 and 15°C at each swimming speed between 20 and 45 cm s^–1. With further increase in temperature to 23.5°C, oxygen consumption declined. Basal oxygen consumption was estimated by extrapolation of the relationship between swimming speed and the logarithm of oxygen consumption to 0 cm s^–1. The metabolic cost of swimming, represented by the difference between total and basal oxygen consumption was independent of temperature at each swimming speed. Energy required to swim 1 km increased from 2.14 to 5.68 J g^–1 between 20 and 45 cm s^–1.     

Benthic flattened cells of the phylogenetically related marine dinoflagellates Protoceratium reticulatum and Ceratocorys mariaovidiorum (Gonyaulacales): a new type of cyst?

A planktonic‐benthic relationship has been described for many dinoflagellate species as part of their ecological strategy to overcome highly variable aquatic environments. Here, the phylogenetically and morphologically related marine dinoflagellates Protoceratium reticulatum and Ceratocorys mariaovidiorum were studied in relation to an unknown benthic life form. In vivo and fixed samples from cultures were analyzed in detail by light and scanning electron microscopy. In both species, a cell type with a morphology different from that of vegetative cells was observed in cultures grown until stationary phase. This cell type was always benthic, swimming sporadically only when it was disturbed. Its main feature included a strong dorsoventral compression. These cells originated from vegetative cells whose protoplasm underwent a progressive flattening, resulting in a gradual detachment of the reticulate and thick thecal plates and the formation of very thin non‐reticulated new plates with pores. When returned to fresh full‐strength medium, the cells recovered their spherical vegetative‐like morphology, including new reticulated thick plates and subsequent cell divisions. The kinetics of flattened cell formation showed that in both species, this cell type increased exponentially until the onset of the culture stationary phase and then decreased. The results of this study are discussed in the context of the planktonic–benthic coupling in dinoflagellate life cycles, including those newly appreciated to be well adapted to the benthic environment.     

THECATE HETEROPHIC DINOFLAGELLATES: FEEDING BEHAVIOR AND MECHANISMS1

The feeding of 18 species of thrcale hetrophi dinoflagellates from three genera (Protoperidininm, Oblea, Zygabikodinium) can all be described within one general framework. These species engulf diatoms and other prey with a pseudopod (herein terned a “Pallium”)which originates at the flagellar pore in the sulcus. The pallium is a highly plastic, membranous organ which rasily strethes to accommodate spines and many as 58 diatom cells in a chain. The contents of the phytoplanklon prey are liquified and transporued throughthe pallioum typically within 7 to 30 minutes of capture (although feeding may last 2 h) teaving an intact but empty cell wall or frustule. Thus far, with few exceptions, Protoperidinium specises have been observed feeding inly on diatoms, whereas two diplopsaloid species feed on dinoflagellates and prasinophytes as well. In four species from the three genera studied. a capture filament has been observed that connects the food to the dinoflagellate prior to extension of the pallium, sometimes allowing the cell to pull the food while swimming. A distinctive precapture swimming behavior is also deseribed foe six species, suggesting that the dinoflagellates are selective grazers.