The dorsal light reactions of larval and metamorphosing flatfish

Before metamorphosis, plaice and turbot show dorsal light reactions generally similar to those of other species, suggesting that their balance mechanisms are also similar, and that their utriculi are functional. The reactions are unusual in one respect in that larvae of both species tilt away from light incident from below the horizontal. In turbot, during metamorphosis, the null position (that taken up when illumination is from directly above) becomes gradually inclined to the right until the larva is horizontal at the end of metamorphosis. There is evidence that the null position of the static system is reset to an inclined position during this change. In plaice, during metamorphosis, the shift in null position is smaller, in the opposite direction, and at the end of metamorphosis there is a rapid change to a horizontal position. The null position of the static system does not appear to be reset. In plaice the influence of light on balance increases during metamorphosis, possibly due to it affecting the brain or the otolith organs. This is not seen in turbot, but light has a strong influence on the balance of this species both before and during metamorphosis. At the end of metamorphosis both plaice and turbot appear to utilize only static information from the sacculi and lagenae. Light could be seen to influence balance in swimming turbot, but not in plaice.     

Copper metabolism in the plaice, Pleuronectes platessa (L.)

Copper metabolism in a teleost, the plaice, Pleuronectes platessa (L.) from a natural environment has been studied. Distribution in the various tissues of the metal and of five key copper-dependent enzymes: ceruloplasmin EC 1.16.3.1; Superoxide dismutase EC 1.15.11; tryptophan oxygenase EC 1.13.1.12; cytochrome oxidase EC 1.9.3.1 and monoamine oxidase EC 1.4.3.4 have been determined. The copper distribution was found to be similar to that in mammals with the greatest concentrations in the brain and heart. Distribution of the copper enzymes is also similar to that found in mammals. A preliminary characterization of the copper enzymes showed that plaice cytochrome oxidase has a pH maximum 2 pH units more alkaline than the mammalian enzyme and that plaice tryptophan oxygenase is more sensitive to heat denaturation than the mammalian enzyme. The present data form a base-line against which studies on factors affecting the copper metabolism in a teleost can be assessed.     

The settlement, distribution and movements of O-group plaice Pleuronectes platessa (L.) in Filey Bay, Yorkshire

The pelagic larval stages and the demersal O-group stage of plaice were sampled in the immediate vicinity of Filey Bay, Yorkshire (54°12'N, 00°16'W), using a modified Gulf V high-speed plankton sampler and beam trawls. The distribution and movements of these fish were studied from the time of first settlement in May through the period of metamorphosis and the O-group stage. It was found that the larvae settled in water deeper than 5 m and moved into shallow water following metamorphosis. This established a typical distribution of larger fish in shallower water than the smaller fish. When the fish grew over 40 mm they moved out again into deeper water which leads to a reversal of the lengthdepth distribution. Further movements during the winter months and of the I-group fish were studied by tagging. It was found that emigration from the nursery ground did not occur until the I-group stage.     

Light,predation and the lekking behaviour of the ghost swift Hepialus humuli (L.) (Lepidoptera,Hepialidae)

We examined the timing of the crepuscular lekking flight of male ghost swift moths in southern Sweden with respect to variations in: (i) the quality of the visual mating signal; and (ii) the behaviour of potential vertebrate predators (mainly bats). The moths'' display flights started ca. 57 min after sunset, and occurred during 20–30 min at incident light intensities between 10.0 and 2.0 lux. Owing to the falling and more shortwave ambient light after sunset, the brightness contrast between the moth wings and the background (grass) increased steeply at the time of display onset. The silvery white male wing colour thereby seems to maximize conspicuousness, and may be a secondary adaptation that facilitates visibility at low light intensities. The display timing itself is probably determined by other factors, possibly predation. By displaying only for a short period at dusk, the moths seem to avoid most birds, which normally do not forage at these light levels, and gleaning bats, which typically do not start to feed until the light intensity has fallen even further. Nevertheless, aerial-hawking bats were often (54% of the evenings, n = 22) seen at the leks, and one species (Eptesicus nilssonii) frequently fed on the displaying moths (22% of the moths observed, n = 83). H. humuli represents an ancient clade among the Lepidoptera. By restricting its sexual behaviour to a short time window at dusk, when predation risk may be minimized (but still high), it may to some extent compensate for the lack of sophisticated predator defence systems such as aposematic or mimetic coloration, manoeuvrable flight, and ultrasonic hearing, which predominate among the more recent Lepidopteran clades. However, the time window solution restricts the moths'' activities considerably and the lack of defence still carries a considerable cost in terms of predation.     

Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine bird: the blue tit (Parus caeruleus L.) and the blackbird (Turdus merula L.)

The spectral absorption characteristics of the retinal photoreceptors of the blue tit (Parus caeruleus) and blackbird (Turdus merula) were investigated using microspectrophotometry. The retinae of both species contained rods, double cones and four spectrally distinct types of single cone. Whilst the visual pigments and cone oil droplets in the other receptor types are very similar in both species, the wavelength of maximum sensitivity (λ_max) of long-wavelength-sensitive single and double cone visual pigment occurs at a shorter wavelength (557 nm) in the blackbird than in the blue tit (563 nm). Oil droplets located in the long-wavelength-sensitivesingle cones of both species cut off wavelengths below 570–573 nm, theoretically shifting cone peak spectral sensitivity some 40 nm towards the long-wavelength end of the spectrum. This raises the possibility that the precise λ_max of the long-wavelength-sensitive visual pigment is optimised for the visual function of the double cones. The distribution of cone photoreceptors across the retina, determined using conventional light and fluorescence microscopy, also varies between the two species and may reflect differences in their visual ecology. Accepted: 8 January 2000     

Reactivation of contraction in detergent-lysed teleost retinal cones

Teleost retinal cones contract in the light and elongate in the dark. In the green sunfish, Lepomis cyanellus, the necklike myoid region of the cone contracts from as much as 120 micrometers (midnight dark- adapted) to 6 micrometers in fully light-adapted state. When dark- adapted fish are exposed to light (1.4 lux), cone myoids contract with a linear rate of 1.5 +/- 0.1 micrometers/min. We report here that detergent-lysed motile models of teleost retinal cones exhibit calcium- and ATP-dependent reactivated contraction, with morphology and rate comparable to that observed in vivo. For reactivation studies isolated dark-adapted retinas were lysed with nonionic detergent Brij-58 (0.1- 1.0%). In reactivation medium containing 10(-5) M free calcium and 4 mM ATP, the lysed cones contracted with normal morphology at in vivo rates (1.4 +/- 1 micrometer/min). Little contraction was observed if ATP or detergent was deleted from the medium or if free calcium levels were less than 10(-8) M. Ultrastructural examination of cone models lysed with 1% Brij-58 revealed that, in spite of extensive extraction of the cytoplasmic matrix, cytoskeletal components (thin filaments, intermediate filaments, microtubules) were still present. Thus we have produced extensively extracted motile models of teleost retinal cones which undergo calcium- and ATP-dependent reactivated contraction with normal morphology at physiological rate.     

Outer retinal fine structure of the garfish Belone belone (L.) (Belonidae, Teleostei) during light and dark adaptation – photoreceptors, cone patterns and densities

In the present EM study, we investigate the retina of Belone belone , a visually-orientated marine predator living close to the water surface. In the duplex retina, four morphologically different cone types are observed: unequal and equal double cones, long single cones and triple cones. In the light-adapted state, five different cone patterns occur: row, twisted row, square, pentagonal and hexagonal patterns. High double cone densities are found ventro-nasally, ventro-temporally and dorso-temporally. Throughout the retina the double cone/single cone ratio is 2 : 1, in the ventral part, however, a 1 : 1 ratio occurs. In the vitreous body we found a curtain-like intraocular septum dividing the retina into two morphologically different regions. In most areas of the dark-adapted retina the cone patterns are absent at the ellipsoid level, with long single cones standing more vitreally in the light path than double cones. The mosaics are retained, however, in the outer nuclear layer. Typical dark adaptation, i.e. the retinomotor movements of the retinal pigment epithelium and photoreceptors in response to the dark adaptation (light change) is not present in the peripheral ventral and parts of the central ventral area. In both regions we found a twisted row pattern of cones having a vitreal position. The findings are discussed with respect to the photic habitat and feeding habits of this species.     

Circadian regulation of teleost retinal cone movements in vitro

In the retinas of many species of lower vertebrates, retinal photoreceptors and pigment epithelium pigment granules undergo daily movements in response to both diurnal, and in the case of teleost cone photoreceptors, endogenous circadian signals. Typically, these cone movements take place at dawn and at dusk when teleosts are maintained on a cyclic light (LD) regime, and at expected dawn and expected dusk when animals are maintained in continuous darkness (DD). Because these movements are so strictly controlled, they provide an overt indicator of the stage of the underlying clock mechanism. In this study we report that both light-induced and circadian-driven cone myoid movements in the Midas cichlid (Cichlasoma citrinellum), occur normally in vitro. Many of the features of retinomotor movements found in vivo also occur in our culture conditions, including responses to light and circadian stimuli and dopamine. Circadian induced predawn contraction and maintenance of expected day position in response to circadian modulation, are also normal. Our studies suggest that circadian regulation of cone myoid movement in vitro is mediated locally by dopamine, acting via a D2 receptor. Cone myoid contraction can be induced at midnight and expected mid-day by dark culture with dopamine or the D2 receptor agonist LY171555. Further, circadian induced predawn contraction can be increased with either dopamine or LY171555, or may be reversed with the dopamine D2 antagonist, sulpiride. Sulpiride will also induce cone myoid elongation in retinal cultures at expected mid- day, but will not induce cone myoid elongation at dusk. In contrast, circadian cone myoid movements in vitro were unaffected by the D1 receptor agonist SCH23390, or the D1 receptor antagonist SKF38393. Our short-term culture experiments indicate that circadian regulation of immediate cone myoid movement does not require humoral control but is regulated locally within the retina. The inclusion of dopamine, or dopamine receptor agonists and antagonists in our cultures, has indicated that retinal circadian regulation may be mediated by endogenously produced dopamine, which acts via a D2 mechanism.     

Low levels of light pollution may block the ability of male glow-worms (Lampyris noctiluca L.) to locate females

Light pollution has been proposed as a factor in the decline of Lampyris noctiluca because it has the potential to interfere with reproductive signaling and has been shown to impact the ability of males to locate light lures in a suburban environment. To compare and test the replicability of this effect in a natural setting and population, imitation females were set out under light polluted and control conditions at varying light pollution intensities in an undisturbed British chalk grassland. Very low levels of light pollution were found to interfere with phototaxis: no males were attracted at either 0.3 or 0.18 lux background lighting versus 33 males collected at paired dark controls. These background illumination levels are much lower than that of 1.5 lux which is recommended by local city councils in Britain to light footpaths. A survey of female L. noctiluca numbers and distribution showed a trend towards female clumping that was not statistically significant. We also found no evidence of light interfering with female signaling behavior.     

Vulnerability of newly settled plaice (Pleuronectes platessa L.) to predation: effects of habitat structure and predator functional response

Plaice (Pleuronectes platessa) nursery grounds on the Swedish west coast have been subject to increasing cover of annual green macroalgae during recent years, with growth of algae starting at the time of plaice settlement in April to May. A laboratory experiment was performed to investigate how the vulnerability to predation of metamorphosing plaice was affected by the presence of filamentous algae. Predation by shrimps (Crangon crangon) on settling plaice larvae was higher on sand than among algae, whereas predation by crabs (Carcinus maenas) was unaffected by habitat type, suggesting a lower overall mortality of plaice in the vegetated habitat. When predators and prey were presented with a combination of the two habitats, predation by shrimps was as high as that in the sand treatment alone, whereas predation by crabs was lower than that in the two treatments with one habitat. Based on these results, an additional experiment was performed, investigating the functional response of shrimps to six densities of juvenile plaice in a sand habitat with alternative prey present. The proportional mortality of juvenile plaice (12-16 mm total length (TL)) was density-dependent and was best described by a type III (sigmoid) functional response of the predatory shrimps. The results suggested that the combined predation pressure from shrimps and crabs was lower among algae than on sand, but settling plaice and predatory shrimps chose the sand habitat. Plaice densities in the sigmoid part of the obtained functional response curve represented normal to high field densities of plaice on the Swedish west coast, suggesting that shrimp predation could have a stabilising effect on plaice recruitment. The formation of macroalgae mats could therefore lead to a concentration of plaice juveniles in the remaining sand habitat and increased mortality through density-dependent predation by shrimps.     

Using KCl to determine size at competence for larvae of the marine gastropod Crepidula fornicata (L.)

Competent larvae of the marine gastropod Crepidula fornicata (L.) were induced to metamorphose (i.e., lose the velum) by elevating sea-water [KCl] by 5–50 mM. The response was optimal at 15–20-mM elevations, at which 50% metamorphosis was obtained in <4 h. Larvae that did not metamorphose during brief exposures (1–5 h) to elevated [KCl] generally maintained the larval form following transfer to control sea water, suggesting that competent larvae must be continuously immersed in the test solutions for metamorphosis to occur. The smallest larvae to respond to elevated [KCl] had shell lengths of ≈700–800 μm, the range of shell lengths within which larvae of this species become responsive to natural inducers. All larvae >≈1125 μm shell length metamorphosed in response to increased [KCl]. Rearing temperature may affect the size at which larvae of this species become responsive to K⁺. CaCl₂ (20-mM concentration elevations), GABA (4×10⁻⁷, 4×10⁻⁶ M), and NaCl (10–20-mM concentration elevations) generally failed to trigger metamorphosis. Twenty-mM elevations of [RbCl] and [CsCl] induced 100% metamorphosis but the juveniles were immobile and died after several days. Elevating [KCl] appears to be a reliable way to assess competence and trigger metamorphosis in larvae of C. fornicata.     

Activity and respiration in the anemone, Metridium senile (L.) exposed to salinity fluctuations

Activity and respiration in the anemone, Metridium senile (L.), were monitored under both constant and fluctuating salinity conditions. During constant exposure to 50% sea water it was found that the animals retracted the tentacles and that the rate of oxygen consumption decreased by ≈50%. The same response was elicited from animals in 100% sea water in a contracted state. Animals exposed to continually fluctuating salinities were found to retract the tentacles, contract the body wall, and produce amounts of mucus during periods of decreasing salinities. These reactions were reversed during exposure to increasing salinity. Oxygen consumption never ceased entirely in animals exposed to dilute sea water and it was found that during declining oxygen tension M. senile regulated its oxygen consumption until the environmental oxygen tension fell to ≈30% saturation.     

An endogenous activity rhythm and the rhythmicity of catches of Nephrops norvegicus (L).

Field and laboratory studies were carried out with Nephrops norvegicus (L.) collected from depths ranging from 10 to 184 m off the west coast of Britain. An endogenous circadian rhythm of locomotor activity was exhibited by prawns from all depths when kept under constant conditions in the laboratory. The rhythm was expressed as nocturnal activity mainly within the burrow, though the prawns repeatedly vacate the experimental burrow during the same period. If, as seems likely, prawns in the field also show burrow-orientated locomotor activity at night this would explain why they are caught in smaller numbers when on commercial fishing grounds attempts are made to trawl them during darkness.Peak catches in trawls occur at differing times of day according to depth, and other workers have suggested that this reflects emergence from burrows at optimal light intensities. Present laboratory experiments show that there is no endogenous basis for a distinct ‘emergence’ rhythm. They suggest that high catches at different times of day at different depths are determined by exogenous factors but not solely by some optimal light intensity. It was not possible to induce emergence from artificial burrows by exposing prawns to light intensities within the presumed optimum range for their depth of capture. Dusk and dawn changes of light intensity may have direct effects upon activity, and light may also have an indirect effect by controlling the availability of suitable food organisms and predators. Light may also have a direct effect by suppressing emergence around midday in intermediate depths, particularly in view of the poor or non-existent light-adapting responses of the retinal shielding pigments.     

The effect of lectins on the attachment and invasion of Enteromyxum scophthalmi (Myxozoa) in turbot (Psetta maxima L.) intestinal epithelium in vitro

The involvement of the lectin/carbohydrate interaction in the invasion of the turbot intestinal epithelium by Enteromyxum scophthalmi was studied in vitro using explants of turbot intestine and pre-treatment of parasite stages with the plant lectins of Canavalia ensiformis (Con A) and Glycine max (SBA). Both lectins inhibited the attachment and invasion of E. scophthalmi stages to the intestinal epithelium, though the inhibitory effect was higher for SBA than for Con A. Such results point to the involvement of N-acetyl-galactosamine (GalNAc) and galactose (Gal) residues and also of mannose/glucose residues in the E. scophthalmi-intestinal epithelium interaction. The inhibitory effect of both lectins on the parasite adhesion and penetration points to the interest of further studies to confirm the presence of putative lectins recognising GalNAc-Gal and mannose/glucose residues in turbot intestine. The obtained results demonstrated also the adequacy of turbot intestinal explants as an in vitro model to study the interaction with E. scophthalmi.     

Light intensity and schooling behaviour in larval gulf menhaden

Schooling behaviour was examined in larval gulf menhaden Brevoortia patronus under decreasing and increasing light intensities and related to movements of retinal photoreceptors and pigment. Nearest neighbour angles and nearest neighbour distances between fish were inversely related to light intensity and were independent of the direction of intensity change. Swimming speeds of individual fish were directly related to light intensity when intensity was decreased but did not increase when intensity was raised. The changes in nearest neighbour angles and distances more closely paralleled movements of retinal pigment cells than cone cell migration, suggesting that rod photoreceptors play a greater role than cones in determining the threshold light intensity for schooling. A general model is developed relating the threshold light intensity for schooling to eye diameter for a number of different species and the implications of this relationship are discussed.     

Le contrôle de l'émergence et des nages nocturnes chez les péracarides des plages de méditerranée. Eurydice affinis Hansen (Isopoda), Gastrosaccus mediterraneus Bacescu, Gastrosaccus spinifer (goës) (Mysidacea)

Along the Mediterranean sandy shores, Eurydice affinis Hansen, Gastrosaccus mediterraneus Bacescu and Gastrosaccus spinifer (Goës) are present during the day in the sand of the beaches, either near the limit of the wave wash or in the superficial fringe of the infralittoral. In the evening there is a general emergence and the animals rejoin the planktonic populations of the open sea.All the three species when kept in aktographs under constant conditions and continuous darkness exhibit a circadian rhythm of emergence and swimming activity of nocturnal frequency. The rhythm is clear and in the two Gastrosaccinae persists for ten days at least; it has been observed over about fifty days in G. spinifer. In E. affinis, the rhythm is only on some occasions clear; one frequently observes a permanent burying in the sediment, in which case the swimming rhythm can be brought to light by a mechanical disturbance or the withdrawal of the sediment.A study of the changes in the photokinetic reactions in a horizontal beam of white light, with intensities varying from 4000 to 10^?6 lux, allow the synchronization of the rhythms to be ascribed to a slight photonegative tendency of the animals at intensities > 1000 lux and a photopositive tendency, often marked, at lower intensities. The variations in the intensity of the reactions in the three species allows an explanation of the autonomous rhythms.A comparison between the photokinetic reactions of these species and those of the infralittoral Peracarida, which migrate in the nocturnal plankton, shows great differences with respect to light intensity. In the latter, the phototaxis is strongly negative and frequently accompanied by a strong photo-inhibition while in the former the phototaxis is still often positive and the kinesis is high. The photopositive reaction allows the animals to rejoin their daytime habitat soon after dawn when the swimming activity decreases.     

The intertidal movements and distribution of young fish on a sandy beach with special reference to the plaice (Pleuronectes platessa L.)

The intertidal movements of young fish were studied on a sandy beach over several tidal cycles during the day and the night. Young plaice (Pleuronectes platessa L.), by virtue of their abundance, received the most attention, and it was found that they migrate up and down the beach with each tide in company with several other less abundant species. The dab (Limanda limanda (L.)), does not make this migration.The vertical distribution of the commoner fish species was charted and each was found to occupy a specific zone in relation to the water, but with large overlaps in many cases. A close relationship between the length of plaice and dabs and the depth in which they were caught was evident during the summer months, but this relationship broke down in the winter due mainly to an offshore movement of larger fish.The food intake of plaice was studied throughout the day and night; feeding occurred mostly during the day. No effect of the tide on feeding could be detected.The possible mechanisms controlling the tide-related movements and the size-depth relationship are discussed and a general hypothesis is put forward in which it is proposed that the fish are able to select and maintain their position at a particular depth throughout a tidal cycle thereby being guided up and down the beach. Attributes of depth which the fish might use as a reference point in their movements are discussed.     

Pigment migration in the compound eye of Manduca sexta: Effects of light,nitrogen and carbon dioxide

Migration of screening pigment granules was studied in the secondary pigment cells of the compound eye of the tobacco hornworm moth Manduca sexta. The granules aggregate at the distal ends of these elongate cells during dark-adaptation, and disperse proximally during light-adaptation, to provide a longitudinal pupil regulating the entrance of light into the eye. Pigment position was measured directly during the couse of migration in sectioned quick-frozen eyes, and the pupillary response was measured in the intact eyes of living moths by reflectance microscopy. The influences of nitrogen and carbon dioxide anaesthesia on pigment migration were investigated in the light of earlier studies on other speicies showing that hypoxia results in dispersal. In accordance with these previous studies, rapid dispersal results from nitrogen hypoxia in Manduca, the pigment spreading farther than it does in light-adaptation. By contrast, the pigment disperses only slightly in response to carbon dioxide hypoxia. Carbon dioxide also inhibits the rapid, extensive dispersal caused by light and nitrogen. Thus the pseudopupil of the eye remains dilated in carbon dioxide anaesthetized moths even under bright illumination. Light-induced dispersal is restored with the addition of oxygen to the carbon dioxide atmosphere. These results suggest, contrary to the conclusions of earlier studies, that pigment dispersal in light-adaptation requires metabolic energy. The inhibition of pigment migration by carbon dioxide is unlikely to be the result of hypoxia; we suggest that low cellular pH affects the mechanism of pigment-granule motility.     

A quantitative description of the behaviour of wild juvenile plaice (Pleuronectes platessa L.)

The behaviour of young plaice (Pleuronectes platessa L.) as they migrate up and down sandy beaches with the tide is described. Their behaviour during this migration consists mainly of swimming and feeding interspersed with rarer behavioural acts. Over short periods swimming behaviour can be described by a random model in which the probability of a swim occurring remains constant. This probability varies markedly, however, from hour to hour. Two types of swimming movement are recognized: one of very short duration represents searching for food and the other, longer, category serves to transport the fish up and down the shore. Variations in the feeding rate and in the frequency, duration, and direction of swimming movements over the tidal cycle are described and related to the changing physical and biological conditions that the fish experience during their intertidal movements.     

Post-larval development of compound eyes and stemmata of Chaoborus crystallinus (De Geer, 1776) (Diptera : Chaoboridae): Stage-specific reconstructions within individual organs of vision

During metamorphosis, the dioptric apparatus of the larval compound eye of Chaoborus crystallinus (Diptera : Nematocera) is radically reconstructed. The thin larval cornea of the ommatidia is replaced by strongly curved corneal lenses, and the eucone larval cone is replaced by an imaginal cone of the acone type. Curvature of the future lens is already apparent in very young pupae, in which the cornea consists only of a thin epicuticle with corneal nipples. Fibrillary cuticle is secreted by cone and primary pigment cells throughout pupal development. Lens formation is accompanied by movement of the nuclei of the accessory pigment cells. The larval cone disintegrates unexpectedly late in young, images. During late pupal development, 7 cone cell projections emerge. In contrast to the dioptric apparatus, the retinula cells and rhabdom remain almost unchanged during metamorphosis. The main refractive element of the larval ommatidium appears to be the cone, while that of the imaginal ommatidium is the corneal lens. In addition to the compound eyes, the pairs of stemmata are retained during the whole post-larval development. Pupal stemmata show no structural differences from the larval stemmata. The stemmata are still present in 2-day-old images (“retained stemmata”), but the primary stemma loses its dioptric apparatus and is proximally relocated to the basal region of the compound eye. The reconstructions in the visual system of Chaoborus, which occur during ontogeny, are probably connected with the change from aquatic living larvae to aerial adults, and appear to fulfill stage-specific needs of vision.