Observations on the diel migration of marine dinoflagellates off the Baja California coast

The vertical distribution of Gonyaulax polyedra, Ceratium furca, Gymnodinium sp., Ceratium dens, Gonyaulax digitale, Prorocentrum micans, Polykrikos kofoidii, and Peridinium depressum was followed during two 36 h time-series stations, off the Baja California coast. Water samples were taken at 6 h intervals, at 7 light penetration depths, for phytoplankton identification and for chemical and biological analysis. The results indicate that the dinoflagellates were able to perform vertical migration against the physical water movements, such as upwelling internal waves and wind mixing, that existed in the area during this study. Attempts to correlate the vertical dstribution of the dinoflagellates with the seawater density and nitrate distribution lead to the conclusion that even a weak density gradient acted as a barrier to the downward migration, and that nitrate assimilation in darkness was not required for their growth. Differences in the vertical distribution at 12.00 hrs of each species supports the hypothesis that migration is of a phototactic nature. A comparison of this study with previous ones on vertical migration in other areas suggests that a species should not be defined in general as a positive or negative phototactic organism, but that the direction of the migration is related to the light intensity at the sea surface.Contribution No. 77-004 of the Bigelow Laboratory for Ocean Sciences, West Boothbay Harbor, Matne 04575, USA.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

The ontogeny of phototaxis by larvae of the crab Rhithropanopeus harrisii

Phototaxis by each zoeal stage of the crab Rhithropanopeus harrisii (Gould) was quantitatively measured by means of a microscope closed-circuit television system. The megalopa stage is indifferent to light stimulation and was thus not tested. The action spectrum for positive phototaxis is similar at each zoeal stage, having the most pronounced maximum at about 500 nm and smaller maxima at 400 and 280 nm. Responsiveness to various intensities of 500 nm light is also similar at each stage. After dark-adaptation strongest positive phototaxis occurs at intensities between 1.0 and 3.0x10^-4 W/m², with no negative phototaxis to lower intensities. After light-adaptation, the positive response occurs to higher intensities between 20 and 0.01 W/m², with a pronounced negative response to lower intensities. Generally, mean swimming speeds during positive phototaxis do not change with stimulation intensity, although during negative phototaxis mean speeds do vary with intensity. At the highest intensities which initiate a negative response, mean values are significantly greater than values for positive phototaxis. Based on the pattern of phototaxis at each developmental stage, a prediction of events during diurnal vertical migration is possible. The negative response after light-adaptation might also function as a shadow reflex.     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Occurrence of a shadow response among brachyuran larvae

A closed-circuit television system was used to study phototaxis and behavioral responses upon sudden decreases in light intensity in light-adapted Stage I zoeae from 7 species of Brachyura (Panopeus herbstii, Menippe mercenaria, Uca pugilator, Callinectes sapidus, Sesarma cinereum, Pinnotheres maculatum, and Libinia emarginata). All species except one show positive phototaxis to high intensities of 500 nm light and negative phototaxis to low intensities. Upon a sudden light intensity decrease, larvae show a shadow response, which depends upon the initial and final intensities. If the initial intensity is sufficient to induce positive phototaxis, and the light is extinguished, the larvae of 6 species stop swimming and passively sink (sinking response). Reducing the intensity to an absolute level that causes negative phototaxis, induces a sinking response followed by negative phototaxis. For intensity reductions that occur in the intensity range above that for negative phototaxis, a sinking responseis observed. The intensity decrease that initiates the sinking response is independent of initial stimulus intensity and duration, is consistent between species, and is equivalent to a decrease by 0.5 OD (optical density) units. The sinking response is greatest at intensity decreases of at least 1.0 to 1.1 OD units. It is concluded that the shadow response is common in brachyuran larvae inhabiting coastal-estuarine areas.     

Rapid response to changing light environments of the calanoid copepod<Emphasis Type="Italic"> Calanus sinicus</Emphasis>

Calanus sinicus is a large calanoid copepod and a dominant species in the coastal waters of Japan. During a research cruise in Sagami Bay on 18 June 1996, we found C. sinicus performing an unusual diel vertical migration (DVM), a behavior that has not been reported in previous studies on this species. This study examined the DVM of C. sinicus under different light environments and revealed the copepods characteristic response to light. Field and laboratory results show that the DVM of C. sinicus is flexible and also confirmed its sensitivity and its rapid response to changing light environments. It is suggested that C. sinicus reacts to changes in absolute light intensity. This feature may be common in oceanic copepod species. The copepods quick reaction to light variation provides decreased predation risks and increased feeding opportunities, which make them a dominant survivor in coastal water habitats.Communicated by T. Ikeda, Hakodate     

Larval behavior of predacious sister-species: orientation,molting site,and survival in Chrysopa

Field and laboratory studies compared two features of larval behavior in a pair of predacious sisterspecies of green lacewings: one (Chrysopa slossonae) a specialist on a single species of colonial aphids (the woolly alder aphid) that occur on branches and trunks of alder trees, the other (C. quadripunctata) a general aphid feeder whose primary prey is dispersed on foliage of diverse types of trees. First, a few hours after hatching, larvae of the two species develop significantly different phototactic responses; the differences correspond well with the spatial distributions of their prey. Most C. slossonae exhibited negative phototaxis, a response that helps move hatchlings inward on alder trees toward the woolly alder aphid colonies, whereas most C. quadripunctata hatchlings showed positive orientation to light, a response that tends to keep them in tree canopies with their prey. Second, in greenhouse experiments, a significantly greater proportion of C. slossonae larvae (second instars) molted within woolly alder aphid colonies and remained with the aphids than did C. quadripunctata larvae. These differences indicate that the specialist larvae have evolved a high degree of behavioral fidelity to their prey. However, larvae (second instars) of the two species that were released near ant-tended woolly alder aphid colonies in the field had similar recovery (= survival) rates. Consequently, natural selection may not act on behavioral traits that influence larval fidelity to prey during the late second and early third instars.     

Vertical migration ofGonyaulax catenata andMesodinium rubrum

Diel migration patterns ofGonyaulax catenata andMesodinium rubrum in the central Baltic during spring 1986 are presented. The depth at which maximum cell concentration of these species occurred was dependent during daytime to a great degree on light intensity; it is hypothesized that the organisms migrated upwards toward higher light intensities until a certain threshold irradiance was reached, after which migration was directed downwards to avoid light intensities higher than the threshold. This threshold level probably depended on nutrient conditions and on the daily average of total irradiance of the foregoing days (light history). Furthermore, it is hypothesized that migration is an important mechanism for adapting to daily irradiance fluctuations. The dependence of migration on irradiance was more obvious forG. catenata than forM. rubrum.     

Influence of temperature on the growth rate of Skeletonema costatum in response to variations in daily light intensity

Daily light intensities (I _o) can vary 10-fold during the winter-spring and late-summer diatom blooms in New England, USA, coastal waters. Laboratory cultures and natural populations incubated in dialysis sacs were examined to determine the time course of growth rate in Skeletonema costatum (Greville) Cleve in response to variations in daily light intensity during two bloom periods in Narragansett Bay, Rhode Island, USA. Log-phase cultures of S. costatum require 2 d to attain maximum growth rates at 2°C following transfer to saturating intensities. At 20°C, only 1 d is required. As temperature increases, Detonula confervacea (Cleve) Gran, Thalassiosira nordenskiöldii Cleve and Ditylum brightwellii (West) Grunow also exhibit rapid increases in mean daily division rates (K) following transfer to saturating light intensities. Thalassiosira pseudonana Hustedt, however, did not alter the time required to achieve maximum K as temperature varied. Natural populations of S. costatum did not show a well-defined relationship between K and light. Throughout a winterspring bloom, K was limited by low temperatures and exhibited no clear response to variations in I _o. A change in K in response to variation in I _o may occur on a daily basis during the summer, when temperatures are near 20°C; this has yet to be verified for in situ populations.     

Copepod photobehavior in a simulated natural light environment and its relation to nocturnal vertical migration

To investigate the roles of light in initiating, controlling and directing nocturnal vertical migration, photoresponses of the adult, female copepod Acartia tonsa Dana were measured under simulated natural underwater light conditions using a video system. Copepods were adapted to a range of background light levels and tested with the following stimuli: absolute quantal intensity, absolute change in quantal intensity and relative (%) change in quantal intensity. The stimulus initiating vertical movements was relative change in quantal intensity, while responsiveness was controlled by the level of light adaptation. A. tonsa swam upward in response to decreases. Response with minimal stimulation occurred at an adaptation intensity close to that in the copepod's natural habitat at the time of the migratory ascent (near the bottom of the Newport River estuary, North Carolina, near sunset). Analysis of the angles of upward movement showed that light is not a directional cue. Relative increases in intensity resulted in sinking, with minimal stimulation required at an adaptation intensity close to that in the field when the migratory descent occurs near sunrise. These results offer a reasonable explanation of how light cues may shape nocturnal vertical migratory patterns.     

Temporal resolution in the eyes of marine decapods from coastal and deep-sea habitats

 Temporal responses of eyes from four decapod species taken from sublittoral (Pandalus montagui Leach), coastal [Nephrops norvegicus (L.)] and deep-sea [Paromola cuvieri (Risso) and Chaceon (=Geryon) affinis A. Milne Edwards and Bouvier] habitats were examined. The electroretinogram responses to a range of sinusoidal intensity modulations between 0.5 and 40 Hz were recorded. Recordings were made from individuals adapted successively to two background light intensities. The sublittoral and coastal species showed faster responses when adapted to the higher light level and they also responded best to intermediate frequencies. When adapted to the lower light level, all species responded most strongly to low-frequency stimuli. Physiological and ecological reasons for the differences in responses are suggested. Received: 2 July 1999 / Accepted: 26 October 1999     

Comparative study of behavioral-sensitivity thresholds to near-UV and blue-green light in deep-sea crustaceans

The behavioral sensitivities of five species of deep-sea crustaceans (order Decapoda: Acanthephyra curtirostris, A. smithi, Notostomus gibbosus, Janicella spinacauda and Oplophorus gracilirostris) to near-UV and blue-green light were studied during a research cruise off the coast of Hawaii in 1993. Two of the five species have electrophysiologically-measured spectral sensitivity peaks at 400 and 500 nm, while the remaining three species have a single sensitivity peak at 490 to 500 nm. In the current study, behavioral mean threshold sensitivities (defined as the lowest irradiance change to which the shrimp would give a behavioral response) were determined for tethered specimens of each species at two wavelengths, 400 and 500 nm. The mean behavioral threshold sensitivities of the two species with putative dual visual-pigment systems were approximately the same to near-UV and blue-green light, while the other three species were significantly less sensitive to near-UV vs blue-green light. Results from these experiments indicate that (1) behavioral information obtained from tethered shrimp accurately reflects their spectral sensitivity, and (2) the sensitivity of the putative dichromats to near-UV light is sufficiently low to detect calculated levels of near-UV light remaining in the down-welling field at their daytime depth of 600 m. Possible functions of this high sensitivity to short wavelength light are discussed.     

Initial observations of the zooplankton microdistribution on the fringing coral reef at Aqaba (Jordan)

Diurnal and nocturnal zooplankton was sampled simultaneously at the surface and at 35 m depth on the outer slope of the fringing reef off the Jordanian coast in the Guff of Aqaba, Red Sea, over a period of 8 d during July 1982. Complementary surface sampling was carried out above the reef and in the open sea in the morning and afternoon for a period of 5 d. Five groups of organisms were classified according to changes in their vertical distribution in the water column which is analyzed and discussed. Zooplankters of the first group appear to be positively phototactic (Calanus robustior, Mecynocera clausi, Oithona helgolandica, Corycella rostrata). Those of the second group also seem to be positively phototactic, but only as regards average and low light (moonlight) intensities (Nannocalanus minor, Acartia negligens, Sagitta enflata, S. hexaptera, Thalia democratica, Oikopleura fusiformis, O. longicauda). The zooplankton of the third group displays a negative phototaxis (Oithona nana, calyptopis and furcilia euphausiid stages, pelecypods and other mollusc veligers). Species exhibiting uncertain behaviour or caught in low numbers and exhibiting no particular distribution are relegated to the fourth and the fifth groups. The horizontal microdistribution of the zooplankton community is not easily discernible: certain neritic or oceanic affinities can be identified, however, by comparing the WP2 standard net surface samples collected above the reef with those from the open sea. The importance of zooplankton communities is emphasized, and the necessity of elucidating the temporal and spatial migrations and exchanges between reef and planktonic organisms in order to gain more insight into their trophic relationship with the reef system.     

Responses of cirripede larvae to light. I. Experiments with white light

The phototactic response of the nauplius larva of Balanus balanoides, B. crenatus and Elminius modestus shows darkadaptation; the response of the cyprid of B. balanoides shows both phototaxis and low photokinesis. The phototactic responses and the orientation of the cyprid to white light at settlement require an intensity of illumination slightly above 10^-5 lux. The ability to select a shaded position by cyprids of R. balanoides requires a higher intensity of 10^-2 to 10^-4 lux; hence a different mechanism may be involved. Barnacle larvae are sufficiently sensitive to be able to respond to light beneath the sea surface, even on cloudy, moonless nights.     

Behavioral responses of larvae of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) during diel vertical migration

Larval Rhithropanopeus harrisii (Gould) show nocturnal vertical migration. Larval behavioral responses to different rates of increase and decrease in light intensity were measured in an apparatus with a natural angular light distribution. A central objective was to establish whether phototaxis actually participates in vertical migration. At sunset the level of light adaptation controlled the readiness of the larvae to migrate, while an ascent was initiated by a preductable relative decrease in intensity (e.g. 4.0x10^-3s^-1). Rates of relative decrease around sunset would evoke continuous upward swimming. Gravity was the orienting cue and there was no change in swimming speed during the ascent. At sunrise, the larval descent was initiated by exposure to an absolute light intensity of about 0.23 log unit above the lower visual threshold. Light served as an orienting cue, as larvae descended by a negative phototaxis. Thus, phototaxis is not a laboratory artifact and does participate in vertical migration. A consideration of behavioral responses of other crustacean zooplankton indicates there is considerable variation in the initiating and directing cues for vertical movements. The variety of behavioral responses of R. harrisii suggests that a synthesis of hypotheses about migration may provide the proper basis for explaining the mechanisms underlying diel vertical migration.     

Laboratory experiments on the diurnal vertical migration of marine dinoflagellates through temperature gradients

The diurnal vertical migrations of 4 marine dinoflagellates (Cachonina niei, Ceratium furca, Gymnodinium splendens, and Prorocentrum micans) were studied in a thermally-stratified Plexiglas column (1.70 m deep, 0.61 m diameter). All species migrated through temperature gradients exceeding 7 C°. Some species exhibited altered migration patterns at different salinities either between experiments or compared to observations by other investigators. P. micans exhibited complex changes in swimming speed that depended both on temperature and on the light cycle. These changes provide insight into the possible time course of a field organism's diurnal exposure to physical variability in the water column. P. micans exhibited compositional changes in response to the light cycle, but not to the cross-thermocline temperature differential. A multi-parameter response matrix is required to model coupling between organisms and biologically-active physical mechanisms realistically.     

Defenses and refuges: alternative mechanisms of coexistence between a predatory gastropod and its ascidian prey

Only five of the 14 species of solitary ascidians in the San Juan Islands, Washington, USA commonly co-occur with an abundant predator of the rocky subtidal, the gastropod Fusitriton oregonensis. None of the common subtidal species is defended by vanadium or sulfuric acid; concentrations of these substances are highest in species that are eaten readily by F. oregonensis and are uncommon in the rocky subtidal. Experimental manipulations indicate that Halocynthia igaboja are protected by stiff spines of the tunic. The defense mechanism of Pyura haustor consists of a tough, leathery tunic with inorganic inclusions, and microscopic spines around the siphons. Boltenia villosa and Styela gibbsii are eaten by snails in the laboratory, but live in areas where the predator is abundant. Both find refuge as epizooites on P. haustor and H. igaboja. The mechanisms by which Cnemidocarpa finmarkiensis persists in the presence of F. oregonensis is unknown.     

A laboratory investigation into the effects of hydrostatic pressure on the vertical migration of planktonic Crustacea

A laboratory-scale vertical migration, using adult Daphnia magna, was observed within an observational pressure vessel. The animals exhibit a clearly defined movement along the water column in response to an overhead light regime. The hydrostatic pressure was increased to 41.5 bar (600 psi) during the course of a migration. a short lived increase in swimming rate was observed, but the pattern of the migration was quickly restored; the migration continued at the high ambient pressure. The response of D. magna to light does not appear to be affected by high pressure.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

Phototaxis in Fungiidae corals (Scleractinia)

Corals within the scleractinian family Fungiidae were observed to move toward light (positive phototaxis). Negative phototactic movement was not observed in any of the specimens tested. On sandy substratum, Diaseris distorta moved faster (max. speed of 3 cm h^-1) than other species. Although D. distorta has symbiotic algae, phototactic movement also was observed both in bleached corals and in those treated with a specific inhibitor (dichlorophenyl dimethyl urea) of photosynthesis. D. distorta was phototactic even on a glass plate, and climbed up a steep slope (up to 30°). Based on experiments with Fungia fungites and D. distorta, soft tissues at the peripheral region of the dise seem to be responsible for movement via peristalsis. It is suggested that positive phototaxis in symbiontbearing fungiid corals is an important trait for selection of favorable habitats.