Photoresponses of Chaoborus larvae

The diel vertical migration of Chaoborus larvae is a well known phenomenon. In order to quantify the ability of larvae to utilize underwater light cues in their migration, we measured photoresponses of fourth-instar Chaoborus punctipennis larvae in the laboratory. The action spectrum for these larvae was characterized by a maximum in sensitivity at 400 nm, a plateau at a lower sensitivity from 480 to 560 nm, and a region of much lower sensitivity at wavelengths longer than 620 nm. Dark-adapted larvae exhibited a positive phototaxis at low light intensity which shifted to a negative phototaxix as light intensity increased. At 540 nm the threshold intensity was 1.5 × 10^?9 W/m² for positive phototaxis and about 10^?6 W/m² for negative phototaxis. Light adaptation decreased sensitivity and altered the phototactic pattern. Larvae have a clear circadian rhythm in negative phototaxis, in which greatest responsiveness occurs early in the day. We suggest that the rhythm in photoresponsiveness primarily controls the timing of the downward migration at dawn.     

Photoresponses of the copepod Mesocyclops edax

The predatory copepod Mesocyclops edax is an important componentof many zooplankton communities where it typically makes extensivedid vertical migrations. To describe the effect of light onadults we measured their photoresponses in the laboratory. Theresponse spectrum is characterized by a wide plateau of greatestsensitivity from about 480 – 580 nm. These animals areadapted to perceive light during the day since their regionof maximum sensitivity overlaps the spectral region of highestquantal intensity underwater (575 – 700 nm). The thresholdintensity for positive phototaxis by dark adapted animals wasabout 5 x 10^–1 Wm^–2 at 540 nm, and they were positivelyphototactic up to an intensity of 5 x 10^–1 Wm^–2.Above this intensity phototaxis is no longer observed. Light-adaptedanimals were less sensitive than dark-adapted, but their generalpattern of response to light intensity did not differ. Thereis no rhythm in phototaxis. Their photoresponses may providea mechanism for controlling vertical migration so as to minimizeexposure to planktivorous fish. ¹Contribution No. 1375-AEL from UM-CEES, Appalachian EnvironmentalLaboratory.     

Photoresponses of late instar <Emphasis Type="Italic">Chaoborus punctipennis</Emphasis> larvae to UVR

With a few clear exceptions (e.g., Daphnia) it is uncertain if most aquatic invertebrates can detect and respond to ultraviolet radiation (UVR). It is known that many aquatic invertebrates are vulnerable to UVR and that anthropogenically-induced increases in surface UVR have occurred in recent decades. We examined the photoresponses of late larval instars of Chaoborus punctipennis to different combinations of UVA (320–400 nm), UVB (300–320 nm) and visible light (400–700 nm) to determine whether the larvae can detect and/or avoid UVR. To accomplish this, we exposed late instar C. punctipennis larvae to a directional light source of UVR only (peak wavelength at 360 nm), visible light only or visible plus various wavebands of UVR. We examined negative phototaxis for 10 min at a quantum flux of 2.62 x 10¹³ quanta s^–1 cm^–2 (S.D. = 3.63 x 10¹² quanta s^–1 cm^–2). In the dark, larvae stayed close to the surface of the experimental vessels. Under all treatments containing visible light the larvae exhibited negative phototaxis and occupied the bottom of the vessels. Under UVR only, the larvae occupied the middle of the water column. Our results suggest that late instar C. punctipennis larvae are unable to detect and avoid UVB and short UVA wavelengths but they can detect long UVA wavelengths.     

Behavioral dissection of Drosophila larval phototaxis

A behavior generally comprises multiple processes. Analyzing these processes helps to reveal more characteristics of the behavior. In this report, light/dark choice-based Drosophila larval phototaxis is analyzed with a simplistic mathematical model to reveal a fast phase and a slow phase response that are involved. Larvae of the strain w¹¹¹⁸, which is photophobic in phototaxis tests, prefer darkness to light in an immediate light/dark boundary passing test and demonstrate a significant reduction in motility in the dark condition during phototaxis tests. For tim⁰¹ larvae, which show neutral performance in phototaxis tests, larvae unexpectedly prefer light to darkness in the immediate light/dark boundary passing test and demonstrate no significant motility alteration in the dark condition. It is proposed that Drosophila larval phototaxis is determined by a fast phase immediate light/dark choice and an independent slow phase light/dark-induced motility alteration that follows.     

Behavior of red king crab larvae: Phototaxis,geotaxis and rheotaxis

Zoeae of Paralithodes camtschatica were positively phototactic to white light intensities above 1 × 10¹³ q cm^?2 s^?1. Negative phototaxis occurred at low (1 × 10¹² q cm^?2 s^?1), but not high intensities (2.2 × 10¹⁶q cm^?2 s^?1). Phototactic response was directly related to light intensity. Zoeae also responded to red, green and blue light. Zoeae were negatively geotactic, but geotaxis was dominated by phototaxis. Horizontal swimming speed of stage 1 zoeae <4 d old was 2.4 ± 0.1 (SE) cms^?1 and decreased to 1.7 ± 0.1 cm s^?1 in older zoeae (P <0.01). Horizontal swimming speed of stage 2 zoeae was not significantly different from ≥4 d old stage 1 zoeae. Vertical swimming speed, 1.6 ± 0.1 cm s^?1, and sinking rate, 0.7 ± 0.1 cm s^?1, did not change with ontogeny. King crab zoeae were positively rheotactic and maintained position in horizontal currents less than 1.4 cm s^?1. Starvation reduced swimming and sinking rates and phototactic response.     

Phototaxis by the Dinoflagellate Gymnodinium splendens Lebour*

A microscope-television system was used to monitor quantitatively the behavior of Gymnodinium splendens Lebour in response to light. The predominant behavioral sequence upon stimulation is (a) an initial 2–5 sec cessation of movement (stop-response) followed by (b) positive phototaxis. The action spectra for each response are identical, having maxima at 450 and 280 nm. Upon measuring the percent response to a range of stimulus intensities, it is apparent that a stop-response is not a behavioral prerequisite for phototaxis. An identical circadian rhythm in photoresponsiveness is observed for phototaxis and for the stop-response with greatest light sensitivity occurring during the first 4 hr of the entrained light period. The implication of phototactic sensitivity and the phototactic circadian rhythm in diurnal vertical migration is discussed.     

PHOTOMOVEMENT RESPONSES OF PORPHYRIDIUM PURPUREUM1

The influence of light quality on positive phototopotaxis by the gliding, unicellular red alga Porphyridium purpureum was obtained using interference filters. Cells exposed to 3 × 10^?7 mol · m²· s^?1 of various wavelengths for 72 h showed maximum topotaxis at 420 and 440 nm. The lower threshold for positive, movement was approximately 5 × 10^?8 mol · m^?2· s^?1. Random movement occurred at nonactinic wavelengths, and no movement occurred in the dark. Cell motility appeared to be unaffected by light polarity, suggesting that the photoreceptor(s) for topotaxis and photokinesis are randomly oriented.     

Larval photoresponses of the polyclad flatworm Maritigrella crozieri (Platyhelminthes,Polycladida) (Hyman)

Phototaxis by larvae of the flatworm Maritigrella crozieri was used to determine spectral sensitivity, the ontogeny of the phototactic pattern, and the lowest light intensity to induce a directional response (intensity threshold). Adult M. crozieri live in shallow water with the tunicate Ecteinascidia turbinata and have a planktonic larval phase lasting longer than 3 weeks. The primary spectral sensitivity maximum was at 500 nm, which is probably an adaptation to the spectrum available underwater at twilight. The phototactic threshold changed with age, as dark-adapted, 1-week-old larvae had a threshold (1.84×10¹⁷ photons m⁻² s⁻¹) an order of magnitude higher than that of 3-week-old larvae. Flatworm larvae are relatively insensitive to light as compared to other invertebrate larvae. Young larvae were positively phototactic at high light intensities and negative at low, a pattern typical of a predator avoidance shadow response. In contrast, older larvae were only positively phototactic, which would be useful for transport to shallow-water adult habitats.     

The responses of Philophthalmus gralli and P. megalurus miracidia to light, gravity and magnetic fields

Miracida of an eyefluke of birds, Philophthalmus gralli, which are positively geotactic, exhibited a positive north-seeking magnetotaxis when subjected to magnetic field strengths from 3 × 10^?4 to 2 × 10^?2 T. A closely-related species, P. megalurus, which is positively geotactic only in complete darkness, exhibited no magnetotaxis under similar conditions. A positive phototactic response overrode the north-seeking magnetotaxis when P. gralli miracidia were subjected to both stimuli in a competing system. No detector mechanisms for magnetic fields are known in miracidia of digenetic trematodes. The order of responsiveness for P. gralli miracidia then is: geotaxis > phototaxis > magnetotaxis.     

Alteration of photoresponses involved in diel vertical migration of a crab larva by fish mucus and degradation products of mucopolysaccharides

Photoresponses involved in the descent phase of nocturnal diel vertical migration (DVM) of larvae of the crab Rhithropanopeus harrisii were measured in a laboratory system that mimicked the underwater angular light distribution. The test hypothesis was that kairomones from fish that activate zooplankton photoresponses involved in DVM are derived from polysaccharides from the external mucus of fishes. Studies considered fish mucus from the mummichog (Fundulus heteroclitus) and disaccharides (originating from chondroitin sulfate A and heparin polysaccharides) that are likely constituents of fish mucus. R. harrisii larvae descend at sunrise with an isolume and remain near the isolume during the day. Since depth maintenance near the isolume depends upon a negative phototaxis, the lowest light intensity (threshold) that induces this response was used to quantify the effects of the test chemicals. It was predicted that exposure to fish kairomones would lower the photoresponse threshold, thereby resulting in larvae remaining deeper in the water column where light for visual predation was reduced. The photoresponse threshold declined as the concentration of fish mucus increased. Disaccharides originating from chondroitin sulfate A and heparin also decreased the photoresponse threshold as compared to responses in aged, filtered seawater. Collectively, the results support the hypothesis and indicate that disaccharide degradation products of predator mucus containing sulfated and acetylated amines can serve as kairomones.     

Automated Recording of Vertical Negative Phototactic Behaviour in Daphnia magna Straus (Crustacea)

Diurnal vertical migration is a well-known phenomenon in the circadian activity rhythms of zooplankton. Our goal was to test whether negative phototaxis in Daphnia magna clone BEAK (provoked by artificially induced light stress, alternating light and dark phases in 2 h intervals), and its interference with the endogenous rhythm of diurnal vertical migration, can be automatically registered with a biomonitor. For the first time the vertical swimming behaviour of D. magna was recorded quantitatively based on non-optical data recording in a fully automated biotest system, the Multispecies Freshwater Biomonitor in a new experimental setup consisting of a column of three recording units (3-level chambers). Circadian vertical migration was clearly recorded in the 3-level chambers and the rhythm was more clear with 5 than with 1 organism per chamber. The organisms clearly responded to induced light stress with negative phototaxis, however best in larger chambers. The artificially induced rhythm was influenced by the endogenous rhythm. This approach may facilitate long-term observations of vertical swimming activity of zooplankton in the future.     

Effect of turbulence on feeding intensity and survival of Japanese flounder Paralichthys olivaceus pelagic larvae

Three‐day rearing experiments were conducted to study the effect of turbulence on the feeding intensity and survival of pelagic larvae of Japanese flounder Paralichthys olivaceus. Four levels of turbulence as control (10^?7·2 m² s^?3), low (10^?6·2 m² s^?3), mid (10^?5·6 m² s^?3) and high (10^?5·0 m² s^?3) were set by changing the flow rate of water pumped through pipes set on the bottom of the tanks. In B‐stage larvae, defined as having buds of elongated dorsal fin rays, the feeding intensity and growth were higher in the low and mid turbulence levels, while survival was highest in the control level. Most of the larvae surviving in the control level, however, were judged to be in a seriously starved condition leading to subsequent high mortality. Because the three‐day span of the rearing experiments was thought to be a little shorter than the periods before starvation‐induced, high mortality occurs. In contrast, for D‐stage larvae, their feeding and growth were optimal in the control and low levels. Feeding was more adversely affected in the high level for D‐stage larvae compared with B‐stage larvae. This is probably due to the compressed body shape and elongated dorsal fin rays of D‐stage larvae, which may be more strongly affected by turbulence and, as a consequence, the larval feeding behaviour such as pursuit and capture of prey organisms becomes less efficient than in lower turbulence. Considering the vertical distribution of B and D‐stage larvae in the oceanic water column, the optimum turbulence level range found in the present study corresponded to a wind speed of 7–10 m s^?1. Therefore, moderate weather conditions of this wind speed range are considered to potentially enhance survival of early larval stages of P. olivaceus.     

Bi‐modal sensitivity to monochromatic light by the mud snail Ilyanassa obsoleta †

Under laboratory conditions the mud snail Ilyanassa obsoleta exhibits a pronounced positive behavioral response to monochromatic light. The response spectrum for‐this positive phototaxis has maxima at 480 nm and 560–580 nm. The threshold intensities for this response are 2.06 x 10^‐7 /μW/cm² for 480 nm and 2.18 × 10^‐7 μW/cm² for 580 nm. These results are suggestive of Ilyanassa's possessing two visual pigments with different absorption maxima.     

Phototaxis in the flagellates,Euglena gracilis and Ochromonas danica

Oriented movement with respect to laterally impinging white light of the flagellates Euglena gracilis and Ochromonas danica has been analyzed in an individual cell study with a microvideographic technique. Using the deviation of track segments (in given time intervals of 1 s) from the light direction as raw data allowed a computer based analysis of the direction distribution. A number of statistical methods employed to test the significance of the obtained results demonstrated an obvious phototactic orientation in Ochromonas which was positive (toward the light source) in low illuminance (1.25 lx=5.3×10^-3 Wm^-2) and negative in higher illuminance (>12.5 lx=5.3×10^-2 Wm^-2). Since in this flagellate the threshold for negative phototaxis is much lower than that for the step-up photophobic response, the hypothesis that negative phototaxis may be brought about by repetitive step-up phobic responses can be rejected for at least this organism. In Euglena positive phototaxis was observed in 50 lx (=0.21 Wm^-2), while an illuminance of 500 lx (=2.1 Wm^-2) caused a negative phototaxis.The experiments were carried out in this laboratory     

LIGHT‐INDUCED MOTILE RESPONSES OF THE ESTUARINE BENTHIC DIATOMS NAVICULA PERMINUTA AND CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Motility of estuarine epipelic (mud‐inhabiting) diatoms is an important adaptation to living in biofilms present within fine sediments. Motility allows cells to migrate within the photic zone in response to a wide range of environmental stimuli. The motile responses of two species of benthic diatoms to photon fluence rates and spectral quality were investigated. Cultures of Navicula perminuta (Grunow) in van Heurck and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reimann both exhibited photoaccumulation at ～200 μmol · m^?2 · s^?1 and photodispersal from photon flux densities (PFDs) of ～15 μmol · m^?2 · s^?1. Photokinesis (changing cell speed) contributed toward photodispersal for both species, and red light (λ = 681–691 nm) was most effective at inducing this process. N. perminuta showed a phototactic (directional) response, with active movement in response to a light gradient. Although this response was exhibited in white light, these directional responses were only elicited by wavelengths from 430 to 510 nm. In contrast, C. closterium did not exhibit phototaxis under any light conditions used in this study. Motile benthic diatoms thus exhibit complex and sophisticated responses to light quantity and quality, involving combinations of photokinesis and phototaxis, which can contribute toward explaining the patterns of large‐scale cell movements observed in natural estuarine biofilms.     

Baculovirus infection triggers a positive phototactic response in caterpillars to induce ‘tree-top’ disease

Many parasites manipulate host behaviour to enhance parasite transmission and survival. A fascinating example is baculoviruses, which often induce death in caterpillar hosts at elevated positions (‘tree-top’ disease). To date, little is known about the underlying processes leading to this adaptive host manipulation. Here, we show that the baculovirus Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV) triggers a positive phototactic response in S. exigua larvae prior to death and causes the caterpillars to die at elevated positions. This light-dependent climbing behaviour is specific for infected larvae, as movement of uninfected caterpillars during larval development was light-independent. We hypothesize that upon infection, SeMNPV captures a host pathway involved in phototaxis and/or light perception to induce this remarkable behavioural change.     

A change in the pattern of vertical migration of Chaoborus flavicans after the introduction of trout

The pattern of diel vertical migration exhibited by Chaoborusflavicans in Lake Lenore, Washington has changed over the pastdecade. Formerly Chaoborus larvae and pupae were present inthe water column during the day. Since 1982, a strong patternof diel vertical migration has been evident in this population.Third and fourth instar larvae and pupae reside in the sedimentsduring the day and then ascend to the surface waters at night.The change in migratory activity of C. flavicans was coincidentwith the introduction of cutthroat trout (Salmo clarki henshawi)into Lake Lenore in 1979. Predation by trout on fourth instarlarvae and pupae of C. flavicans was probably responsible forthe observed changes in migratory behavior.     

Larval Behaviours and Their Contribution to the Distribution of the Intertidal Coral Reef Sponge Carteriospongia foliascens

Sponges (Phylum Porifera) are an evolutionary and ecologically significant group; however information on processes influencing sponge population distributions is surprisingly limited. Carteriospongia foliascens is a common Indo-Pacific sponge, which has been reported from the intertidal to the mesophotic. Interestingly, the distribution of C. foliascens at inshore reefs of the Great Barrier Reef is restricted to the intertidal with no individuals evident in adjacent subtidal habitats. The abundance of C. foliascens and substrate availability was first quantified to investigate the influence of substrate limitation on adult distribution. Pre-settlement processes of larval spawning, swimming speeds, phototaxis, vertical migration, and settlement to intertidal and subtidal substrate cues were also quantified. Notably, suitable settlement substrate (coral rubble) was not limiting in subtidal habitats. C. foliascens released up to 765 brooded larvae sponge⁻¹ day⁻¹ during the day, with larvae (80%±5.77) being negatively phototactic and migrating to the bottom within 40 minutes from release. Subsequently, larvae (up to 58.67%±2.91) migrated to the surface after the loss of the daylight cue (nightfall), and after 34 h post-release >98.67% (±0.67) of larvae had adopted a benthic habit regardless of light conditions. Intertidal and subtidal biofilms initiated similar settlement responses, inducing faster (as early 6 h post-release) and more successful metamorphosis (>60%) than unconditioned surfaces. C. foliascens has a high larval supply and larval behaviours that support recruitment to the subtidal. The absence of C. foliascens in subtidal habitats at inshore reefs is therefore proposed to be a potential consequence of post-settlement mortalities.     

Proximate control of diel vertical migration in Phyllosoma larvae of the Caribbean spiny lobster Panulirus argus

Phyllosoma larvae of the spiny lobster Panulirus argus undergo diel vertical migration (DVM), in which they are at depth during the day and nearer the surface at night. This study determined the visual spectral sensitivity of Stage I larvae and investigated whether light plays a proximate role in DVM as an exogenous cue and as an entrainment cue for an endogenous rhythm in vertical migration. Under constant conditions, larvae have a circadian rhythm (24.5-h period) in vertical swimming that resulted in a twilight DVM pattern. The behavioral response spectrum and electroretinogram recording indicated two photoreceptor spectral classes with maxima at 360 and 486 nm. When stimulated in an apparatus that simulated the underwater angular light distribution, dark-adapted larvae showed only positive phototaxis, with a threshold intensity of 1.8 × 10(13) photons m(-2) s(-1) (3.0 × 10(-5) μmoles photons m(-2) s(-1)). They have an avoidance response to predator shadows in which they descend upon sudden decreases in light intensity of more than 69%. When stimulated with relative rates of decrease in light intensity as occur at sunset they ascended, whereas they descended upon relative rates of light intensity increase as occur at sunrise. Thus, the DVM pattern is controlled by both an endogenous circadian rhythm in swimming and behavioral responses to light at sunrise and sunset.     

Behavioural response of larval Atlantic menhaden to different rates of temperature change

Larval Atlantic menhaden Brevoortia tyrannus, spawned off North Carolina (U.S.A.) during the winter, undergo cross-shelf transport from the western Gulf Stream edge to coastal bays and estuaries. Variation in water flow direction with depth provides larvae the opportunity to enhance shoreward transport, if they can regulate their depth behaviourally. Temperature, which normally decreases with depth on the continental shelf, is one possible cue for depth regulation. Laboratory-reared larval menhaden of two different ages were exposed to varying relative rates of temperature increases and decreases, which were presented from both above and below the larvae. Temperature decreases from below caused an ascent response in both young and old larvae, but neither responded to this cue from above. The minimum (threshold) relative rates of decrease for initiating ascents were similar (7.9 × 10^?2, 10.7 × 10^?2° C min^?1) for both age larvae as were the minimum absolute amounts of decrease that must occur before a response (0.1, 0.05°C). Young larvae did not respond to a temperature increase, while old larvae ascended regardless of whether the increase was presented from above or below. Threshold relative rates of increase were 8.59 × 10^?2°C min^?1 from below and 14.79 × 10^?2° C min^?1 from above. The threshold rates and range of larval speeds during vertical movements were used to calculate vertical temperature gradients that could be perceived. These values were compared to measured gradients in areas inhabited by menhaden larvae. On the continental shelf, detectable temperature gradients appear common for temperature decreases that would occur upon descending and temperature increases upon ascending. However, it is uncommon for larvae to encounter temperature increases upon descending that would initiate an ascent response. These results support the hypothesis that menhaden larvae are capable of using temperature gradients for depth regulation.