Scanning electron microscopy of metamorphosis in four species of barnacles (Cirripedia Thoracica Balanomorpha)

Metamorphosis and early juvenile development was followed in the laboratory in Balanus amphitrite and B. improvisus (family Balanidae) and in Semibalanus balanoides and Elminius modestus (family Archaeobalanidae) from cyprid settlement until 24 h after ecdysis. Stages of development were studied in vivo and with scanning electron microscopy. Events in metamorphosis and early juvenile development are very similar in all four species and can be interpreted in terms of a highly modified moult. Notably, there was no indication whatsoever of the amorphous decorticated settler previously described from metamorphosis of B. amphitrite. The shape of a juvenile barnacle with cirri and incipient shell plates is apparent immediately after the shedding of the cypris carapace, and rudimentary peduncle can be distinguished below the developing wall plates. A basal row of cuticular hairs encircles the peduncle in all species except S. balanoides. These hairs seem to serve a restraining function during early development, when the juvenile barnacle is only attached by the initially secreted cyprid cement. Similarly situated hairs are also present in very young juveniles of the lepadomorph Scalpellum scalpellum. In Semibalanus balanoides nothing indicates that the rostrum originates from the fusion of two latera as previously claimed, since this plate is single as soon as it can be distinguished.     

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.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Influence of light and temperature on the photosynthetic rate of marine benthic diatoms

The influence of light and temperature on photosynthetic rate as measured by C¹⁴O₂-fixation of marine benthic diatoms was investigated, using both intact sediment samples (Marshall et al., 1973) and suspensions of diatoms harvested by a lenstissue technique (Eaton and Moss, 1966). After C¹⁴-incubation, sediment samples were filtered, burned in a sample oxidizer, and their activity determined in a liquid scintillation counter. Photosynthetic rate of mixed field populations is saturated by a light intensity of approximately 10,000 lux; at still higher light intensities no photoinhibition was found. In contrast to the mixed field populations, unialgal cultures of the benthic diatom Amphiprora alata Kütz. exhibited strong photoinhibition at higher light intensities (10,000 to 60,000 lux). Within a range of 4° to 20°C, the photosynthetic rate increased about 10%/C^o. No differences in photosynthetic pattern were observed between epipelic and epipsammic species.     

Environmental control of the breeding of three boreo-arctic cirripedes

Three species of barnacles with boreo-aretic distribution were shown to require maintenance for several weeks below a critical temperature before the breeding condition could be attained. The temperatures critical for Balanus balanoides (L.) Balanus balanus (L.) and Balanus crenatus (Bruguière) were found to be between 10° and 12°C, 10° and 14°C, and at about 17°C, respectively. Although the strong influence of continuous light and the weak influence of continued feeding in delaying the onset of breeding in B. balanoides were confirmed, there remained some outstanding anomalies between the breeding behaviour of this species under laboratory conditions and between the tide marks. It was found impossible to initiate breeding by the application of conditioning procedures significantly in advance of the time of the normal Autumn breeding season in B. balanoides. Breeding appears to be inhibited, independently of external conditions, for a set period after the preceding brood cycle. Evidence points to a similar, largely endogenous, control of breeding in B. balanus which also breeds once annually, but not in B. crenatus which breeds continuously so long as food and temperature levels permit.     

Influence of activity and salinity on the weight-dependent oxygen consumption of the rainbow trout Salmo gairdneri

Two types of secretory cell have been found in the cement glands of the cypris larva of Balanus balanoides (L.). Histochemical tests show the presence of proteins, phenols and the enzyme polyphenol oxidase in both the glands and secreted cement. It is proposed that cyprid cement is composed of tanned protein, all the components of which are derived from the cement glands.     

Response to light by trochophore larvae of Spirobranchus giganteus

Trochophore larvae of Spirobranchus giganteus (Pallas) respond positively to white light at levels of illumination from 1 to 2 168×10¹⁴ quanta cm^-2 s^-1. In this range the strength of the response is not correlated with irradiance level. The response is increased by dark adaptation. At low levels of irradiance (0.1-2.0×10¹⁴ quanta cm^-2 s^-1) larvae respond positively to blue (360-510 nm, max. 430 nm) and green (475–620 nm, max. 530 nm) light but not to wavelengths of 590 nm or over. The light response develops gradually during the 12 h following the appearance of the eyespot and is maintained throughout the remainder of the planktonic phase.     

Light responses of the larva of the serpulid polychaeteGaleolaria caespitosa

AdultGaleolaria caespitosa were collected at Cape Banks and Balmoral Bay, both in the vicinity of Sydney, Australia, from October 1985 to February 1986. Planktotrophic larvae of this serpulid polychaete are negatively phototactic at irradiance levels between 10 and 2 000x10¹⁴ q cm^-2 s^-1. The effective spectral range lies below 530 nm. Early metatrochophore larvae are less strongly photonegative; later they become photopositive. The photopositive phase persists into the actively swimming 3-setiger stage. Older setigerous larvae are photoneutral, demersal and positively thigmotactic. The strength of the photonegative response is not affected by the level of irradiance. The photopositive response is weaker at low levels of irradiance. The photonegative behaviour of young larvae may keep them at an optimal level for feeding. The subsequent photopositive phase may increase chances of being washed ashore and the terminal, photoneutral, thigmotactic phase may be important in the selection of a suitable site for settlement. Comparisons are made with light responses of two other serpulid species. Larval responses to light may be important infitness, especially in relation to successful niche occupation by adults.     

An energy budget for the free-swimming and metamorphosing larvae of Balanus balanoides (Crustacea: Cirripedia)

Analysis of biochemical components and measurements of oxygen consumption rates of cypris larvae of Balanus balanoides (L.) maintained in the laboratory at 10°C for up to 5 weeks after capture shows that lipid is the primary energy reserve, although later protein is utilised. Initially, the cyprids swim freely with an oxygen consumption rate of ca. 37×10^-3 l O₂ h^-1 cyprid^-1, but within a few days the rate falls to ca. 21×10^-3 l O₂ h^-1 cyprid^-1 when they cease swimming and explore the substratum. The cost of metamorphosis was calculated both from the loss of biochemical components and oxygen consumption rates during metamorphosis; the values were 2.8×10^-2 and 3.2×10^-2 cal cyprid^-1, respectively. A budget was collated from the data on respiration and biochemical composition, whereby the energy per cyprid was partitioned into that required for essential structural components (6.8×10^-2 cal), that needed for metamorphosis (3.0×10^-2 cal) and an excess available for swimming and exploring, which in the batches studied was about 5.0×10^-2 cal. This excess is mainly derived from the utilisation of lipid reserves and is used up usually 2 1/2 to 4 weeks after capture. During these measurements, samples of cyprids were taken at weekly intervals to test the rate of settlement and success of metamorphosis. The results showed that they lose their competence to metamorphose successfully approximately at the same time (3 to 4 weeks) that the energy supply for swimming and exploration is used up.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Growth and competition of the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana. II. Light limitation

The marine diatoms Phaeodactylum tricornutum (Bohlin) and Thalassiosira pseudonana (Hasle and Heimdal) were grown under both continous illumination and a 14 h light: 10 h dark cycle at light intensities ranging from 1.53×10^-4 to 2.95×10^-1 ly min^-1. Under both photoperiods, T. pseudonana exhibited higher division rates than P. tricornutum at high light intensities, but the reverse was true at all light intensities <3×10^-3 ly min^-1. Comparison of these results with available data on light-limited growth of other planktonic algae suggests that P. tricornutum may be unusually efficient at maintaining its cell division rate at low light intensity. This efficiency may contribute substantially to its success in turbid, nutrient-enriched mass algal culture systems, the only environments in which it is known to attain great numbers.Contribution No. 4086 from the Woods Hole Oceanographic Institution.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

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.     

Effect of light on apical length growth of Fucus distichus spp. edentatus (Phaeophyta)

The length growth of Fucus distichus L. ssp. edentatus (Pyl.) Powell in continuous light is compared to growth in natural daylight. In continuous light the length growth of vegetative and fertile apices were saturated at 25 to 40 Wm^-2 and 15 to 30 Wm^-2, respectively. In natural daylight vegetative apices reached saturation at about 300 Wm^-2 (in May), while fertile apices were not saturated at 180 Wm^-2 (in March). This shows that saturation levels are dependent on the light regime and that they may be greatly underestimated when continuous light is used. At 20 Wm^-2 vegetative and fertile apices were daylength saturated at 8 and 12 to 16 h, respectively. Maximum growth in vegetative apices occurred in July, but growth rates were high also during wintertime (45% of July maximum in March at 4.0°–4.2°C). This corresponds to the experimental results which indicate that F. d. edentatus grows well at low irradiances and during short photoperiods.     

Effect of short-term variations in light intensity on photosynthesis of a marine phytoplankter: A laboratory simulation study

Parameters of photosynthesis and growth were measured for a marine diatom (Lauderia borealis) grown in axenic continuous culture under three different light regimes: constant, simulated diurnal variation, and fluctuating. The light fluctuations were systematic increases and decreases in light intensity superimposed on the diurnal regime. In the first two regimes, a morning maximum and an afternoon depression in photosynthesis were observed. In the fluctuating light regime, the afternoon depression was less pronounced and the morning maximum was enhanced. The results may be explained by postulating a time-dependent value for the light-saturated rate of photosynthesis. Light utilization [mmol O₂ cell^-1 (E m^-2)^-1] was the same for the diurnally varying and fluctuating regimes, despite the fact that the peak light intensity in the fluctuating regime was double that of the diurnally varying regime.     

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.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. I. Physiological investigations of Chaetoceros armatum and Asterionella socialis in laboratory cultures

Two species of diatoms, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which account for virtually all of the plant biomass present in the surf zone along the Olympic Peninsula (USA), have been isolated and grown in culture, and comparisons have been made with respect to their physiological behavior in response to aeration, light, temperature, salinity, and nutrients. A. socialis displayed the greater maximum growth rate under most situations, which would give this species a natural advantage over C. armatum. However, in the natural habitat, C. armalum is by far the more important of the two, since the blooms often consist of almost pure stands of this species. This may be due to its apparent ability to use low light levels more efficiently. The maximum growth rate of C. armatum was reached at light levels of 750 to 1000 lux in cultures, whereas A. socialis required a higher light intensity (4000 lux) for maximum growth. C. armatum did not produce its mucilage envelope under any culture conditions. Such a mucilage envelope, with its accompanying clay particles, is invariably present surrounding the diatom chains when they are collected from the natural habitat. The absence of the mucilage envelope (and clay particles) may explain the peculiarities of iron nutrition which were observed in culture experiments, i.e., a dependence on the presence of iron as NaFe-EDTA or ferric citrate, and failure to utilize ferric choloride as a source of Fe for growth.Contribution No. 652 from the Department of Oceanography, University of Washington.     

Inorganic nitrogen metabolism inUlva rigida illuminated with blue light

Inorganic nitrogen metabolism in blue light was studied for the green algaUlva rigida C. Agardh collected in the south of Spain (Punta Carnero, Algeciras) in the winter of 1987. NH₄ ⁺ has been reported to inhibit NO₃ ^- uptake; however,U. rigida showed a net NO₃ ^- uptake even when the NH₄ ⁺ concentration of the external medium was three or four times greater than the concentration of NO₃ ^-. NO₃ ^- uptake rates were similar in both darkness and in blue light of various photon fluence rates (PFR) ranging from 17 to 160 mol m^-2 s^-1. Since NO₃ ^- uptake is an active mechanism involving the consumption of ATP, respiratory metabolism can provide enough ATP to maintain the energetic requirement of NO₃ ^- transport even in darkness. In contrast, NO₃ ^- reduction inU. rigida was highly dependent on the net photosynthetic rate. After 7 h in blue light, intracellular NO₃ ^- concentrations ([NO₃ ^-] _i ) were higher in specimens exposed to intensities below the light compensation point (LCP) than in those incubated at a PFR above the LCP. When PFR is below the light compensation point, NO₃ ^- reduction is low, probably because all the NADH produced by the cells is oxidized in the respiratory chain in order to produce ATP to maintain a steady NO₃ ^- transport rate. The total nitrogen (TN) and carbon (TC) contents decreased from darkness to 33 mol m^-2 s^-1 in blue light. In this range, catabolic processes prevailed over anabolic ones. In contrast, increases in TN and TC contents were observed above the light compensation point. The C : N ratio increased with light intensity, reaching a stable value of 17 at 78 mol m^-2 s^-1 in blue light. Intracellular NO₃ ^- concentration and NO₃ ^- reduction appear to be directly controlled by light intensity. This external control of [NO₃ ^-]_i and the small capacity ofU. rigida to retain incorporated NO₃ ^-, NO₂ ^- and NH₄ ⁺ ions may explain its nitrophilic character.     

Balanus amphitrite (Cirripedia: Thoracica) — A potential indicator of fluoride,copper, lead,chromium and mercury in North Adriatic lagoons

The capacity of the barnacle Balanus amphitrite Darwin to accumulate pollutants above ambient levels was examined in two North Adriatic lagoons with respect to fluoride, copper, lead, chromium and mercury. Levels in soft tissues ranged from 138 to 312 ppm (dry weight) for fluoride, 41 to 109 ppm for copper, 7.1 to 11.7 ppm for lead, 2.10 to 3.89 ppm for chromium, and from 0.96 to 1.35 ppm for mercury. The concentration factor was of the order of 10² for fluoride, 10³ for copper, and possibly higher than 10³ for lead, chromium and mercury. The differences of about 2:1 in the fluoride and copper concentrations in the waters of the two lagoons were clearly reflected by the levels in the barnacles. B. amphitrite that had set on experimental panels had levels similar to those found in specimens collected from long-term natural populations as early as 42 days after immersion of the panels. Compared with literature data, the accumulation levels found in B. amphitrite for fluoride were among the highest known for invertebrates; the values for copper and lead were considerable, but exceeded by others published for B. balanoides; the results for chromium and mercury were one order of magnitude lower than values reported for other suspension-feeders or indicator organisms. B. amphitrite appears to possess most of the properties considered essential for a biological indicator. Eventual determination of the response time of the barnacle to changes in environmental level could profitably be carried out utilizing experimental panels.     

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae) in culture

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae), cultured from a cyst in sediments of Suo-nada, Japan in 1986, was investigated by culture experiments. Nitrogen limitation was effective in inducing cyst formation. After incubation in N-limited medium at 25°C with 10000 lux on a 14 h light, 10 h dark photo-cycle, cultures were subjected to six different light intensities (0 10000 lux) in order to determine effects of light intensity on cyst formation. Cysts were formed under light intensities of 1000 lux or below, but not at 10000 lux. Cysts formed in culture displayed morphological characteristics quite similar to those natural cysts observed in sediments collected from the Inland Sea of Japan. Observations on nucleus, using diaminidinophenylindole (DAPI)-staining and epifluorescence microscopy, indicated that the majority of cysts formed in culture were uninucleate, a few were binucleate. Germination of cysts was observed under adequate conditions for germination (22°C, 3500 lux with a 14 h light, 10 h dark photo-cycle) even after storage at 11°C in the dark for more than 4 mo.