## bioluminescent dinoflagellates species

(, Zirbel, M. J., Veron, F. and Latz, M. I. Pyrocystis fusiformis, the largest species tested, had the brightest flashes over the same flow range. Two species of bioluminescent dinoflagellates, similar in size and shape, and one nonluminescent diatom species were offered to A. tonsa as prey items. bio luminescent squid. Furthermore, the bioluminescence threshold shear stress for L. polyedrum is similar for steady and unsteady laminar Couette flow (von Dassow, 2003). Non-photosynthetic dinoflagellates mainly feed on the diatoms as well as other dinoflagellates while Pyrosystis Noctiluca size ranges from 200-400 micrometers and feeds on zooplankton and fish eggs. 4). Pyrocystis Fusiformis Once the required They Bioluminescence is the characteristic feature of dinoflagellates. For C. horrida, there was an inflection at a wall shear stress of 0.4 N m−2. Non-photosynthetic dinoflagellates mainly feed on the diatoms as well as other dinoflagellates while Pyrosystis Noctiluca size ranges from 200-400 micrometers and feeds on zooplankton and fish eggs. Further details of data collection are found in Latz and Rohr (Latz and Rohr, 1999). (, Vanderploeg, H. A. and Paffenhöfer, G. A. In a similar way, the response thresholds for escape behavior of copepods appear to be tuned to environmental shear levels (Fields and Yen, 1997). For 20-ms response latency, this position is again within Rcritical, indicating that flashes for these species will not occur within the feeding current. Both species are considerably larger in size than L. polyedrum and are non-spherical. Supported by the Office of Naval Research (grant N00014-95-1-001 to M.I.L.) The grow lamps should be placed about three feet away to avoid heat damage and should light for solid twelve hours. No data were collected when the flow was transitioning between laminar and turbulent flow, for a wall shear stress of 3–7 N m−2. Equivalent spherical diameter = (length × width2)1/3. \$8.50. Their bioluminescence is an unusual defense mechanism and provides a great opportunity to discuss animal adaptations. Dinoflagellate bioluminescence is a powerful model system for assessing flow sensitivity because each flash is a near-instantaneous reporter (Eckert, 1965; Widder and Case, 1981a) to suprathreshold levels of shear in the immediate fluid environment of a single cell (Hamman and Seliger, 1972; Latz et al., 1994, 2004). Donaldson, T. Q., Tucker, S. P. and Lynch, R. V. (, Eppley, R. W., Holm-Hansen, O. and Strickland, J. D. H. (, Eppley, R. W., Reid, F. M. H., Cullen, J. J. et al. But the light can be different depending on the bioluminescent species present. Suspension feeders such as copepods capture organisms by movement of the second maxillae to direct particles from the feeding current toward the mouth (Koehl and Strickler, 1981; Vanderploeg and Paffenhöfer, 1985; Price and Paffenhöfer, 1986) where they are macerated by the mandibles (Arashkevich, 1969). The pipe-flow apparatus was the identical system used by Latz and Rohr (Latz and Rohr, 1999), consisting of a 75-L acrylic tank attached through a gently constricting inlet to a 6.35-mm internal diameter clear polycarbonate pipe 1 m in length. In support of this assumption is the finding that the response threshold for copepod escape jumps is similar for steady tangential and spatially changing extensional shear stress (Kiørboe et al., 1999). Maximum intensity values based on pooled data for each species began to plateau at wall shear stress values of approximately between 2.5 and 6 N m−2 for C. fusus, 0.1 N m−2 for C. horrida, 2 N m−2 for L. polyedrum and 1 N m−2 for P. fusiformis. In turbulent flow, shear stresses vary in time and space. These aforementioned dinoflagellates are often chosen to grow at home for aesthetic purposes or out of curiosity. In turbulent flow, although flashes would often overlap, preventing accurate measures of the intensity of individual flashes, general trends in maximum flash intensity as a function of wall shear stress could still be discerned. Images of live cells of the bioluminescent dinoflagellates (A) Ceratium fusus (scale bar = 50 μm), (B) Ceratocorys horrida (scale bar = 25 μm), (C) Lingulodinium polyedrum (scale bar = 25 μm) and (D) Pyrocystis fusiformis (scale bar = 100 μm). and the IAR and ILIR programs at SSC San Diego (to J.R.). Pyrocystis fusiformises lose their flagella as they mature and are thus non-motile dinoflagellates with tapering ends. Search for other works by this author on: In fully developed laminar and turbulent pipe flow, shear stress is greatest at the pipe wall and decreases linearly to zero at the pipe centerline. Together with mathematical models coupling cell response to flow conditions (Deane and Stokes, in press), flow-stimulated dinoflagellate bioluminescence can be applied as a quantitative tool for probing complex flow fields not amenable to conventional study. See text and Latz and Rohr (Latz and Rohr, 1999) for a description of methods used to calculate thresholds. Symbols represent results from single flow rates for data pooled from all experiments. The highest value of maximum intensity for each species was 3.41 ± 1.18 × 109 photons s−1 for C. fusus, 2.59 ± 0.67 × 109 photons s−1 for C. horrida, 5.77 ± 2.70 × 109 photons s−1 for L. polyedrum and 4.79 ± 2.75 × 1010 photons s−1 for P. fusiformis. (, Krasnow, R., Dunlap, J., Taylor, W. et al. Longer shear exposure, at much lower shear stress levels, affects dinoflagellate swimming, population growth and morphology. (, Huntley, M. E., Sykes, P., Rohan, P. et al. The accelerations within this range are from 0.31 to 25 m s−2 and are not considered to be stimulatory (Latz et al., 2004). Motile cells possess two dissimilar flagella arising from the ventral cell side = dinokont flagellation (Fig. The length scales of the turbulence ranged from the radius of the pipe (Davies, 1972) to of the order of 10 μm. Bioluminescent Dinoflagellates. Dinoflagellate bioluminescence provides a nearly instantaneous index of flow sensitivity. Both modes of stimulation involve mechanical deformation of the cell that if sufficient will activate a calcium signaling pathway (von Dassow, 2003), generation of a vacuole action potential (Eckert, 1966; Widder and Case, 1981a) with proton flux into the cytoplasm and pH activation of the luminescent chemistry (Fritz et al., 1990). Bioluminescent dinoflagellate ecosystems are rare, mostly forming in warm-water lagoon s with narrow openings to the open sea. Lunula is the topographic (, Buskey, E. J., Strom, S. and Coulter, C. (, Chen, A. K., Latz, M. I. and Frangos, J. Average intensity, expressed as photons s−1, as a function of wall shear stress for (A) Ceratium fusus, (B) Ceratocorys horrida, (C) Lingulodinium polyedrum and (D) Pyrocystis fusiformis. This species of dinoflagellate is very beautiful but is usually very tricky to grow. (, Lapota, D., Geiger, M. L., Stiffey, A. V. et al. The lights startle the predators and also attract high trophic level organisms that feed on dinoflagellate’s predators. 4. Wall shear stress increases to the first power with average flow rate in laminar flow, but as the 1.75 power in turbulent flow (Schlichting, 1979). (, Pahlow, M., Riebesell, U. and Wolf-Gladrow, D. A. For C. fusus, the slope was 1.2 ± 0.6, while it was 1.6 ± 0.5 for P. fusiformis. By this criterion, for L. polyedrum, C. horrida, C. fusus and P. fusiformis to be subjected to the small-scale spatial structure of turbulent flow, 10 LK would have to be <50, 100, 350 and 900 μm, respectively. units of photons s−1) (Latz and Rohr, 1999; Rohr et al., 2002). Thus, L. polyedrum had the lowest population response proportion at its threshold, while C. horrida had the highest. Threshold values of wall shear stress were 0.116 ± 0.02 N m−2 for C. fusus, 0.024 ± 0.009 N m−2 for C. horrida and 0.087 ± 0.02 N m−2 for P. fusiformis. In turbulent flow, the larger the organism relative to the energetic scales of the turbulence, the more effective the turbulence is at deformation (Levich, 1962). Except for C. fusus, the change in average intensity (photons m−3) associated with transition from laminar to turbulent flows for pooled data was less than or equal to that expected by extrapolating the power regression found in laminar flows to higher values of wall shear stress (Fig. firefly luciferin. Thick solid line is for a flow rate of 0.279 mL s−1 (Jakobsen, 2001). (, Rohr, J. J., Allen, J., Losee, J. et al. Although C. fusus exhibited a conspicuous increase in average intensity (presented as photons m−3) through transition to turbulent flow, P. fusiformis, which is larger than C. fusus, did not. Therefore change the culture after this period. Symbols represent individual values from the pooled set of experiments for each species for laminar (solid) and turbulent (open) flows. They need an appropriate amount of sunlight, food, temperatures, and environmental stressors. N, number of experiments for each species. The description of good for this organism is an area that remains at a moderate temperature. impart blue-green light as other bioluminescent organisms in their phylum. Non-marine bioluminescence is less widely distributed.The two best-known forms of land bioluminescence are fireflies and glow worms. The spines of C. horrida may increase shear sensitivity, perhaps by acting as levers that accentuate the effect of shear flow (Zirbel et al., 2000). However, measurements of average intensity are often expressed per unit volume [i.e. Conversely, average flow rate increases to the first power as a function of wall shear stress in laminar flow and to the 1/1.75 power with wall shear stress in turbulent flow. However, flashes occurring at or after predator contact may yet benefit the individual cell if it is released unharmed because of a ‘startle response’ and/or benefit the population if the flash disrupts grazing behavior (Buskey and Swift, 1983, 1985; Buskey et al., 1983). A stiff cell wall may decrease flow sensitivity by minimizing flow-induced deformation (Märkl et al., 1991; Namdev and Dunlop, 1995; Joshi et al., 1996) or possibly increase sensitivity by distributing external forces across the cell as observed for other cell types (Helmke and Davies, 2002). 26 No. It is no doubt that bioluminescent dinoflagellates are of marine origin and their life cycle is bounded within the sea. The average shear stress across the pipe is two-third of wall shear stress (, $L_{\mathrm{K}}\ =\ \left(\frac{{\nu}^{3}}{{\epsilon}}\right)^{0.25}$, Biology and the Mechanics of the Wave-Swept Environment, The Response of Bioluminescent Organisms to Fully Developed Pipe Flow, Naval Oceanographic Office Technology Report, Proceedings of the First International Conference on Toxic Dinoflagellates, Journal of Plankton Research Vol. In summary, the present results suggest that bioluminescence is stimulated by predator contact but not by typical oceanic flows that might continually deplete luminescent reserves. The container has to be transparent which allows proper distribution of sunlight. This difference between threshold and oceanic background dissipation rates is consistent with the general lack of background bioluminescence observed in the ocean interior (Boden et al., 1965; Widder et al., 1989; Buskey and Swift, 1990). With few exceptions, such as breaking waves and wave-forced bottom shears in shallow nearshore areas, these threshold shear stress levels are several orders of magnitude larger than typical oceanic ambient flows. (, Boden, B. P., Kampa, E. M. and Snodgrass, J. M. (, Buskey, E. J., Reynolds, G. T., Swift, E. et al. If organisms are stimulated near the wall, then the local value of shear stress will be less than the maximum value at the wall. The only difference is that maximum intensity for C. fusus appeared to level off at slightly higher wall shear stress levels than for L. polyedrum. Thus, for any siphon flow rate and a 20-ms response latency, only Ceratocorys horrida will flash within the siphon flow field. Dinoflagellates in marine environments can multiply rapidly with the right combination of sunlight and nutrients, and when the water around them is agitated their distinctive blue or green glow becomes visible. No data were collected when the flow was transitioning between laminar to turbulent, for a wall shear stress range of 3–7 N m−2. A. As expected, the slope of the power regression of intensity (photons s−1) as a function of wall shear stress was decreased by 1 for the photons m−3 regression: 0.60 ± 0.04 for C. fusus, 3.27 ± 0.44 for L. polyedrum and 2.53 ± 0.57 for P. fusiformis. Dinoflagellates also require to be placed in a good corner at your home. While generally for the dinoflagellate species studied bioluminescence would not be stimulated by predator feeding currents, bioluminescence may still serve as an index of mechanical sensitivity to contact and handling by the predator. These dinoflagellates impart blue-green light when disturbed, whatever the disturbance may be ranging from a swimmer’s hand to breaks due to waves. Except for C. horrida, this relationship for laminar flow was modeled as a simple power regression (R2 = 0.85–0.87). They have a life cycle of about 5-7 days and they reproduce asexually during this time frame. copepods) that graze upon the dinoflagellates (Burkenroad, 1943; Morin, 1983; Mensinger and Case, 1992; Abrahams and Townsend, 1993; Fleisher and Case, 1995). Laboratory cultures of C. fusus Ehrenb, C. horrida Stein [strain 89A from the Sargasso Sea, see (Latz and Lee, 1995)] and P. fusiformis Murray were grown in seawater with f/2 additions at half strength (Guillard and Ryther, 1962) minus silicate as previously described (Latz and Rohr, 1999) on a 12:12 h light–dark cycle. They have a specialized organelle in their body called scintillations which oxidize luciferin to produce light. Dinoflagellates need a proper container to grow at home. Whether cells with these attributes have an antipredation advantage has yet to be proven but is an attractive hypothesis. It is a marine organism that can impart light. Photomultiplier measurements in units of counts s−1 were converted to photons s−1 using a previously described calibration procedure (Latz and Rohr, 1999). This study is the first to examine the relationship between cell size and flash intensity using a range of characterized laminar and turbulent flow conditions. For the pipe-flow experiments, the shear exposure time, based on average flow rates, is between 20 s at near threshold flows and 0.4 s at the highest flow rates (Latz and Rohr, 1999). The ecological significance of dinoflagellate bioluminescence extends beyond the direct interaction of dinoflagellate and predator. Bioluminescence can be observed in this mixture of marine dinoflagellates. Stiff cell components such as thecae or a cell wall could decrease flow sensitivity by minimizing local flow-induced cell deformation. Although Rcritical changed with siphon flow rate, the corresponding shear stress remained constant (Fig. Statistical tests were performed using Statview software (SAS Institute, Inc.). Pyrocustis noctiluca is marine plankton that has the ability to produce its light. Dinoflagellate produces this light through metabolizing luciferin and light up water bodies as a defense mechanism. A target cell concentration of 15 cells mL−1, as previously used with L. polyedrum (Latz and Rohr, 1999), was used for all species because at that concentration individual flashes could be resolved essentially throughout all laminar flow conditions. Regardless of the specific strategy, the range of shear sensitivity must be limited for dinoflagellate bioluminescence to have a beneficial antipredation affect. When they sense a predator approaching, they quickly let out light. The bioluminescence response thresholds for C. fusus and C. horrida have not been previously measured. Using siphon flow as a mimic of a predator feeding current, the associated velocity, shear stress and acceleration fields can be calculated as a function of distance from the mouth of the siphon based on the volume flow rate through the siphon (Kiørboe et al., 1999). The dinoflagellates (Greek δῖνος dinos "whirling" and Latin flagellum "whip, scourge") are single-celled eukaryotes constituting the phylum Dinoflagellata. materials like the species, proper solutions, lamps, and the container are The dinoflagellate P. bahamense var. These measurements involved the identical apparatus and methods of Latz and Lee (Latz and Lee, 1995). Copepods can reject unsuitable particles (Huntley et al., 1986) apparently after handling and possibly tasting the items (Vanderploeg and Paffenhöfer, 1985). bahamense has a bioluminescent capacity of 3.35 × 10 8 photons 42 and is normally present in the Indian River Lagoon, FL, during most of the summer and early fall. marker of their distal part which has the ability of nail production. The critical distance Rcritical within which stimulated organisms would be advected into the mouth of the siphon before flashing was calculated based on the 20-ms response latency of dinoflagellate bioluminescence (Widder and Case, 1981a). The slope of the power regression for values of wall shear stress <0.4 N m−2 was 2.29 ± 0.45, while the slope was 0.14 ± 0.36 for the range of higher laminar flows. (, Märkl, H., Bronnenmeier, R. and Wittek, B. Previous work using nozzle flow (Latz et al., 2004) corroborates the present quantitative finding that the response threshold for C. horrida was less than that for L. polyedrum. They are better referred to as algae and there are nearly 2000 known living species. (, Clift, R., Grace, J. R. and Weber, M. E. (. Relative cell size is the ratio of cell length, the largest dimension of size, to 10 LK at the pipe wall. Jellyfish typically feed on dinoflagellates and other microscopic algae, fish eggs, and even other jellyfish. In the present study, C. fusus and P. fusiformis were the only species whose size was on the order of the energetic turbulent length scales (Fig. Thus, mechanical stimulation of dinoflagellate bioluminescence is ecologically important not only in the context of predator interactions with dinoflagellate cells but also as a ‘mine field’ that potentially increases the risk of predation to moving animals. They noted that while the single-celled, bioluminescent dinoflagellates are usually poor competitors compared to other plankton because of their slow growth rates, the copepods reject them in favor of grazing on more poorly defended but otherwise faster-growing plankton species. In laminar flows, maximum intensity varied with flow stimulus. It is assumed, for comparison sake, that the threshold bioluminescence levels determined in steady, tangential shear in pipe flow is similar to that in unsteady, extensional shear in siphon flow. It is well known that surface breaking waves stimulate bioluminescence (Staples, 1966; Latz et al., 1994) as do bottom shears created by passing waves in a laboratory wave tank (M. I. Latz and J. Rohr, personal observations). An adjustable valve manually controlled flow through the pipe, and pressure drop was measured by two pressure ports connected to a variable reluctance differential transducer. The relatively high intraspecific variation in threshold values from different experiments for C. fusus and P. fusiformis resulted in a lack of statistically significant difference between their thresholds. Moreover, individual flash intensity was either at or near its peak at 1 N m−2. The maximum intensity of C. horrida, P. fusiformis and L. polyedrum was not noticeably affected by the transition to turbulence. Relative sensitivity was assessed on the basis of various bioluminescence parameters of organism and population response. Only for C. fusus was there a highly significant difference in maximum intensity between high laminar and turbulent flows (t-test, t = 5.3, df = 25, P < 0.001). Bioluminescence, which means "living light," occurs in fireflies, certain fungi and fish, and microorganisms like Dinoflagellates, a species of algae. Although dinoflagellate bioluminescence occurs in the vicinity of grazing copepods (Buskey et al., 1985), it is unknown whether the bioluminescence of the studied dinoflagellates is stimulated by fluid shear in the feeding current of the predator or by contact with the predator. The time scales associated with the smallest eddies in the present turbulent pipe flow study were on the order of 0.1–1 ms (Rohr et al., 2002). There was a significant difference in thresholds among species (ANOVA, F = 19.8, df = 2,9, P < 0.0005), with the threshold for C. horrida being significantly different from those of the other two species (Fisher’s PLSD post-hoc test, P ≤ 0.002 for each), which were not significantly different from each other (Fisher’s PLSD, P = 0.09). 2). 8) for a given response latency. This organism was the smallest species studied, is thecate and has a roughly spherical shape. Further analysis used threshold values based on the regression method because it is less sensitive to differences in the flow rate increment (Latz and Rohr, 1999). However, this change in maximum intensity in turbulent flow was no higher than that extrapolated from the trend in laminar flow, suggesting that the cause was the increase in shear stress and not the turbulent nature of the flow. They require nitrate, phosphate, trace metals, and vitamins which are prepared in sterile conditions. 3) indicated how the response proportion changed as a function of wall shear stress between threshold and the highest laminar flows measured. Flow-stimulated bioluminescence has been used to visualize boundary layer shear associated with a moving dolphin (Rohr et al., 1998), examine shear stress in bioreactors (Chen et al., 2003) and quantify shear stress within breaking surface waves (Stokes et al., 2004). PyroFarms DinoNutrient Spout Pouch (10oz) 4.3 out of 5 stars 17. Morphological and bioluminescence characteristics of the dinoflagellate species studied. The threshold value of wall shear stress of 0.3 N m−2 for L. polyedrum was also slightly greater than for Couette flow, in which the threshold occurred at a shear stress of 0.1 N m−2 (Latz et al., 1994). Unless otherwise stated, values represent means with standard deviations of the mean from this data set. (, Karp-Boss, L., Boss, E. and Jumars, P. A. It may be also useful for mapping highly dissipative oceanic flows (Rohr et al., 2002). The advantage of pipe flow is that organisms are continually replenished in the flow field because of advection. Most marine light-emission is in the blue and green light spectrum, the wavelengths that pass furthest through seawater.The examples are Dinoflagellates, angler fish, and the other. The response threshold was calculated as the wall shear stress value at which the value of the fitted regression model was equal to the average background intensity for that experiment (Latz and Rohr, 1999). is usually grown at home for experiments or aesthetic purposes. This ranking, though not conclusive, is consistent with increased flow sensitivity due to increasing size and the presence of spines. 3). Otherwise, paired or unpaired t-tests were used for pair-wise comparisons. In the context of the present study, morphological features thought to affect flow sensitivity of bioluminescent dinoflagellates include cell size, the presence of spines and the possession of thecae or cell wall. For example, in the deep ocean, ϵ is on the order of 10−10 m2 s−3 (Kunze and Sanford, 1996). Nevertheless, considering the different flow characteristics of fully developed pipe and Couette flow, the similarity in response thresholds for these completely independent flow fields indicates that organisms are responding to specific, quantitative aspects of the flow, regardless of the flow field. (, Swift, E., Sullivan, J. M., Batchelder, H. P. et al. Average bioluminescence in fully developed, turbulent pipe-flow scales with cell concentration (Rohr and Latz, unpublished data), consistent with a negligible bioluminescence contribution from cell-to-cell collisions. Average intensity of bioluminescence was calculated by time averaging each time series record collected at a constant flow rate (Latz and Rohr, 1999). Without knowledge of the mechanotransduction mechanism responsible for flow-induced bioluminescence, and because morphological features co-vary among species, it is impossible to determine conclusively which morphological features are most important to flow sensitivity. 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