Factors affecting the feeding rate of the winkle Littorina littorea

The winkle Littorina littorea (L.) feeds mainly when moistened by the tide and is quiescent when exposed to the drying effects of the wind and sun. Upper shore winkles thus experience a reduced feeding time compared with individuals living lower in the intertidal zone, but are able to completely compensate by an increased rate of radular activity when immersed. The rate of radular activity is influenced by the body size of winkles, by experimental temperature and by acclimation temperature, but is mainly modified by the duration of exposure to air which directly controls the rate of radular activity on immersion.     

Geographic variation in the growth rates of Littorina littorea and L. saxatilis

The two intertidal grazers Littorina littorea and L. saxatilis occupy extensively overlapping distributions on both sides of the Atlantic. Morphological, electrophoretic and physiological evidence suggests that the direct developer L. saxatilis is more geographically differentiated than the planktonic developer L. littorea. This hypothesis was tested by collecting both species in 1980 and 1981 from three geographically separated beaches in Massachusetts, USA, marking them, and releasing them in two of the sites. Individuals from all three L. littorea populations, grown at a common site, tended to grow at similar rates, while L. saxatilis exhibited consistent population differences in individual growth. This study supports the view that L. saxatilis achieves its broad geographic distribution via locally adapted ecotypes, while L. littorea achieves its distribution by individual physiological plasticity.     

The nature of heat coma in Littorina littorea (Mollusca: Gastropoda)

 The nature of heat coma was examined in the edible periwinkle Littorina littorea (L.). Duration of acclimation did not influence heat-coma temperature at 12 °C, although other acclimation temperatures were important in influencing thermal tolerance, with positive shifts in coma temperature observed in response to elevated temperatures. Previous thermal history also influenced heat-coma temperatures. Individuals subjected to repeat heat-coma events on a daily basis showed significant declines (P < 0.05) in coma-temperature; in contrast individuals exposed to repeat heat-coma events on a weekly basis showed no decline in thermal tolerance. Size-effects occurred at selected sites, where decreased heat-coma temperatures were recorded in large individuals. Received: 16 August 1999 / Accepted: 13 June 2000     

Effect of shell repair on shell strength in the gastropod Littorina irrorata

Repair of damage to the outer lip of gastropod shells results in a scar whose presence could weaken the shell against crushing forces. Shell strength in Littorina irrorata (Say), measured as force required to break the shell between a moving bar and a stationary steel plate, was found to be similar for scarred and unscarred individuals. This suggests that scars do not affect crushing resistance adversely. Cracks tended to propagate along sutures and along growth lines. These seams appear to be more important than scars as lines of weakness in the shell of L. irrorata.     

Quantitative alterations in the structure of the digestive cell of Littorina littorea on exposure to cadmium

The effects of sublethal concentrations of cadmium (Cd) on the morphology of the digestive gland of the periwinkle Littorina littorea have been investigated using a planimetric procedure. Five planimetric parameters were recorded (mean epithelial thickness: MET; mean diverticular radius: MDR; mean luminal radius: MLR; MLR/MET and MET/MDR) and tested by diverse statistical methods. The results indicate that MET is depressed after long exposures at high Cd concentrations whilst MLR increases. Variations in MDR have been attributed to different conditions in the tissues surrounding digestive tubules, and are not-significant in relation to the treatment. However, certain increases might be significant due to the presence of parasites or hemocyte proliferation. The ratios MLR/MET and MET/MDR are highly sensitive, and can be expressed as statistical functions of metal concentration and exposure time. Combination of these five planimetric parameters is reliably indicative of the structural changes occurring in the digestive gland of molluscs under conditions of environmental stress.     

Cyclic AMP-dependent protein kinase: role in anoxia and freezing tolerance of the marine periwinkle Littorina littorea

A key regulatory mechanism underlying the switch between aerobic and anaerobic metabolism amongst anoxia-tolerant marine molluscs is reversible protein phosphorylation. To assess the role of cAMP-dependent protein kinase (PKA) in aerobic–anaerobic transitions, the effects of anoxia on the activity and subcellular distribution of PKA were assessed in foot and hepatopancreas of the marine periwinkle, Littorina littorea. Exposure to N₂ gas at 5 °C caused a rapid decline in the percentage of total enzyme present as the free catalytic subunit (PKAc) in both tissues; the percentage of PKAc fell from ∼30% in controls to 3% after 1 h anoxia and remained low over 72 h. Total PKA also fell by 30% after 72 h anoxia in hepatopancreas but rebounded during aerobic recovery. Freezing at −8 °C elicited parallel results for both percentage of PKAc and total PKA, suggesting that PKA responses to freezing were stimulated by the ischemia that develops when hemolymph freezes. Anoxia also led to a shift in PKA subcellular distribution in hepatopancreas (but not in foot), the percentage of total PKA activity associated with the nuclear fraction dropping from 25% in controls to 8% in 12 h anoxic snails with opposite changes in the cytosolic fraction. The catalytic subunit (PKAc) of foot PKA was purified to a final specific activity of 63.5 nmol phosphate transferred per minute per milligram protein. Enzyme properties included a molecular weight of 33 to 35 kDa, an activation energy from Arrhenius plots of 65.1 ± 4.8 kJ mol⁻¹, and substrate affinity constants of 151 ± 6 μM for the phosphate acceptor, Kemptide, and 72 ± 9 μM for Mg.ATP. Activity was strongly reduced by mammalian PKA inhibitors (H-89, PKA-I), by neutral chloride salts (I₅₀ values 165 to 210 mM) and by NaF (I₅₀ 62 mM). Reduced PKA activity under anoxic or freezing conditions would facilitate the observed suppression of the activities of numerous enzymes that are typically PKA-activated and thereby contribute to the overall anoxia-induced metabolic rate depression. Received: 19 November 1997 / Accepted: 30 September 1998     

The paradox of Rockall: why is a brooding gastropod (Littorina saxatilis) more widespread than one having a planktonic larval dispersal stage (L. littorea)?

In benthic invertebrates dispersal of planktotrophic larvae is generally considered more effective than is, for example, the rafting of adults or egg masses. It is certainly true that over short distances, viz., in the range of tens of kilometres or less, a moderately long-lived planktotrophic larva represents an effective mechanism of dispersal. However, turbulent mixing and mortality will decrease the concentration of planktotrophic larvae, and at some distance from the ancestral population the density of settlers may be too low to enable future matings between adults of low mobility. On the other hand, adults, juveniles or benthic egg masses drifted over long distances may colonize new habitats. The crucial point is the type of larval development of the organism. If the founder group belongs to a species with direct development or which produces very short-lived planktonic larvae, the low mobility of all life-stages will maintain a population within a restricted area so that mates will be likely to encounter each other even in a small population. Even if transport of benthic stages happens very rarely, this may be more influential than larval dispersal over long distances. To show that this may be true the detailed geographical distribution of two intertidal gastropod species with contrasting modes of development is presented and further support from the literature for this hypothesis is discussed.     

Intraspecific physiological variability of the gastropod Littorina saxatilis related to the vertical shore gradient in the White and North Seas

 Physiological responses to desiccation and temperature stress as well as behavioural responses to fast and abrupt environmental changes were investigated in high- and low-shore Littorina saxatilis (Olivi) from several populations from the White and North Seas. Variations in evaporation rates, resistance to air exposure and to acute and chronic temperature stress between animals from different shore levels were similar in White and North Sea periwinkles, consistent with the adaptive nature of these variations. High-shore snails were found to be able to conserve body water reserves better, to resist higher temperatures and to survive longer under conditions of combined temperature and desiccation stress than their low-shore counterparts. In a temperature range of 25 to 35 °C, the rate of evaporative water loss was positively correlated with temperature in low-shore snails while being largely temperature-independent in high-shore snails. Median lethal time during air exposure in L. saxatilis was negatively but not linearly related to the temperature of exposure. In a temperature range of 30 to 38 °C, the resistance to heat exposure in air was only slightly dependent on the temperature, with Q ₁₀ = 1.4 for the median lethal time; the heat resistance dropped drastically at temperatures above 38 °C, with Q ₁₀ = 593.8. This suggests different mechanisms of temperature resistance in different parts of the studied temperature range. In contrast, behavioural response to extreme salinity fluctuations was not uniform in the high- and low-shore periwinkles from the White and North Seas, which may reflect specific environmental conditions at different shore levels in the two areas studied. Observed physiological and behavioural variations are discussed from the viewpoint of different adaptive strategies employed by eulittoral and eulittoral-fringe animals within populations of a single species. Received: 13 December 1999 / Accepted: 11 April 2000     

Two heavy metal-binding proteins in the midgut gland of the crab Carcinus maenas

Two heavy metal-binding proteins occur naturally in the midgut glands of Carcinus maenas (L.) collected from the Firth of Clyde, Scotland. These proteins, of approximately 27,000 and 11,500 MW (molecular weight) have previously been described as Cd-binding proteins after their induction by high concentrations of cadmium in the laboratory. The 27,000 MW heavy metal-binding protein is bound to about 0.10 g-at of Cd, 0.70 g-at of Zn and 0.31 g-at of Cu per mole of protein; 7.7, 7.9 and 1.1%, respectively, of the soluble Cd, Zn and Cu in the midgut gland are associated with this 27,000 MW protein (6.7, 1.6 and 0.9% of the total midgut gland Cd, Zn and Cu). The 11,500 MW protein is bound to about 0.04 g-at of Cd, 0.37 g-at of Zn and 1.54 g-at of Cu per mole of protein; 29.3, 31.3 and 41.7%, respectively, of the soluble Cd, Zn and Cu in the midgut gland are associated with this protein (25.7, 6.3 and 34.4% of the total midgut gland Cd, Zn and Cu). Neither heavy metal-binding protein is bound to measurable amounts of lead.     

Predator-induced alarm responses in the common periwinkle, Littorina littorea: dependence on season, light conditions, and chemical labelling of predators

Chemically mediated alarm reactions of the common periwinkle, Littorina littorea (L.), were studied in laboratory experiments during two consecutive summers, and one intermediate autumn season. Responses to chemical stimuli were detected as crawl-out responses, i.e. movements of snails out of the water. Snails were exposed to extracts of injured conspecifics, extracts of the mussel Modiolus modiolus (L.), and water conditioned by the predatory crab Carcinus maenas (L.), which had been maintained on different diets. In experiments carried out during the summer, a significantly larger number of snails moved out of the water when exposed to chemical stimuli from injured conspecifics, compared to chemical stimuli from injured mussels or filtered seawater. These results suggest that chemical alarm substances are present in L. littorea. Water conditioned by crabs that had been fed L. littorea released significantly more crawl-out responses compared to water conditioned by crabs that had been kept on a fish diet. When tested in autumn, no significant differences were found in responses to the above-mentioned water samples. Crawl-out responses under different light regimes were also investigated. All series of experiments carried out in the dark evoked a higher number of responses compared to series that took place in light. These findings may indicate an adaptation of snails to night-active predators. In total, the current results suggest that a L. littorea diet may chemically “label” the predator crab with snail alarm substances, and that predator-induced responses of L. littorea are actually responses to conspecific alarm substances released from crabs that have been maintained on a L. littorea diet. The response to the alarm signal, however, appears to be dependent on season and light conditions; some ecological implications of these findings are also discussed. Received: 8 January 1999 / Accepted: 29 March 1999     

The introduction of Littorina littorea to British Columbia, Canada: potential impacts and the importance of biotic resistance by native predators

Although the establishment and spread of non-indigenous species depends upon survival in the face of novel environmental conditions and novel biological interactions, relatively little attention has been focused on the specific role of native predators in limiting invasion success. The European common periwinkle, Littorina littorea, was recently introduced to the Pacific coast of Canada and provides a case study of an introduction into an area with an important predator guild (sea stars) that is functionally minor in the invader’s native habitat. Here, we assess the likelihood of establishment, spread, and negative ecological impact of this introduced gastropod, with an emphasis on the role of native sea stars as agents of biotic resistance. Size frequency distributions and local market availability suggest that L. littorea was most likely introduced via the live seafood trade. Non-native hitchhikers (e.g., the trematode Cryptocotyle lingua) were found on/in both market and field specimens. Laboratory studies and field observations confirmed that L. littorea can survive seasonal low salinity in Vancouver, British Columbia. Periwinkles also readily consumed native Ulva, suggesting that periwinkles could impact native communities via herbivory or resource competition. Unlike native gastropods, however, L. littorea lacked behavioural avoidance responses to Northeast Pacific predatory sea stars (Pisaster ochraceus and Pycnopodia helianthoides), and sea star predation rates on L. littorea were much higher than predation rates on native turban snails (Chlorostoma funebralis) in common garden experiments. We therefore expect periwinkle establishment in British Columbia to be limited to areas with low predator density, as is seen in its field distribution to date. We caution that this conclusion may understate the importance of the L. littorea introduction if it also serves as a vector for additional non-indigenous species such as C. lingua.     

Trail following,speed and fractal dimension of movement in a marine prosobranch,Littorina littorea,during a mating and a non-mating season

We quantitatively studied movement behaviour in the gastropod Littorina littorea in laboratory experiments during periods of their non-mating season (November 1992) and their mating season (April–May 1993). Snails were collected in 1992 and 1993 from one boulder shore on the north west coast of Sweden. In a comparison between the two seasons (one non-mating and one mating) we measured trail complexity of unsexed snails using fractal dimesion, the degree of mucus trail following (coincidence index of marker and tracker trails) and average movement speed of marker and tracker snails. We found no differences in fractal dimension and coincidence index of trails between the two seasons. Tracker snails moved, however, significantly faster than marker snails during both seasons. This could not be explained by trackers, when locomoting, using the mucus trail deposited by the marker to increase their speed, since there was no correlation between coincidence index and tracker speed. During the mating season we also conducted trail complexity, trail following and speed experiments comparing the behaviour of males and females. There was no difference between males and females in the fractal dimension of movement, nor was there any difference between the mean speed of male and female snails, although male marker snails tended to move faster than female marker snails. Males tracking other males, females tracking other females and females tracking males followed trails about equally long distances (i.e., coincidence indices did not differ). In contrast, males following female mucus trails showed a significantly higher degree of trail following than the other sex combinations. This new finding may suggest that females of L. littorea release pheromones in their mucus trails and that males are able to identify them.     

Biochemical composition and energy reserves in the larvae and adults of the four british periwinkles Littorina littorea,L. littoralis,L. saxatilis and L. neritoides

The biochemical composition of the adult body tissue is similar in Littorina littorea, L. littoralis, L. saxatilis and L. neritoides. In the newly crawling metamorphosed young of L. littoralis and L. saxatilis, the biochemical composition is similar to that of the mature adults. The newly released planktonic veliger larvae of L. littorea and L. neritoides, which represent and earlier stage of development than the newly crawling young, have a neutral lipid level (mean, 16.5% of ash-free dry flesh weight) approximately three times the level (5.4%) in the newly crawling young of L. littoralis and L. saxatilis. Otherwise the biochemical composition of free-swimming larvae is similar to that of newly crawling young and adults. Neutral lipid is apparently utilised by L. littoralis and L. saxatilis larvae during larval development and metamorphosis. It is suggested that neutral lipid is the major energy reserve of Littorina veliger larvae whereas in the adults, as exemplified by L. littorea, both lipid and carbohydrate are important as energy reserves.     

Genetic variation in the mangrove periwinkle Littorina angulifera

Twenty populatios of Littorina angulifera, inhabiting islands composed of the mangrove tree Rhizophora mangle, were assayed at an esterase locus to determine whether genetic differentiation was associated with distance between populations. It was predicted, on the basis of larval dispersal in this species, that genetic differentiation between populations on islands separated by long distances should be greater than those on islands located near each other. A chi0square test of homogeneity revealed significant differences in esterase gene frequencies among the 20 island populations. However, there was no association of distance between islands and genetic heterogeneity. In addition, a cline in gene frequency was found to be assiciated with latitude. Factors responsible for the observed pattern of heterogeneity at the esterase locus are discussed.     

Yolk utilization in the gastropod Crepidula fornicata

The increases in constituents per unit weight of eggs during embryonic development of the gastropod Crepidula fornicata amounted to 14.6% for ash, 1.0% for protein, and 0.3% for non-protein nitrogen. During the same stages, fat content decreased from 33.7 to 20.3%, carbohydrate from 10.2 to 7.7% and energy content from 6209 to 5298 cal/g dry organic substance. The cumulative efficiencies for yolk utilization were 83.8% for dry weight, 61.0% for energy, 85.1% for protein, 50.7% for fat, and 63.6% for carbohydrate. A single egg contained 0.0269 cal, a single veliger 0.0164 cal. Of the 0.0105 cal expended on metabolic processes of the embryo, oxidation of fat contributed as much as 65.3%, while that of protein and carbohydrate amounted only to 18.8 and 6.3%, respectively. On the basis of ecophysiological considerations, a new classification of eggs is proposed.This paper is based on a lecture presented during the first Convention of the Indian Association of Biological Sciences in Madras, December, 1968.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.     

Age-and sex-dependent patterns of aggregation in the tropical gastropod Strombus luhuanus

The spatial dispersion of the tropical coral sand-dwelling mesogastropod Strombus luhuanus Linné, 1758 was studied at several sites in two localities (Australian Great Barrier Reef and southern Papua New Guinea), from September 1980 to February 1983. S. luhuanus usually occurred in local aggregations, many of which were relatively dense with discrete boundaries: Four types of aggregation are described: mixed age-class, juvenile, mating, and clusters. The former two are termed colonies, because they persisted over time and exhibited some coordinated movements. Mixed age-class colonies contain individuals of all ages, but within them the different ageclasses are frequently spatially segregated from each other. This was sometimes due to the younger age-classes' preference for shallower areas, but also occurred in areas of constant depth. Juvenile colonies are relatively smaller in area but higher in density, and are dominated by a single size-class. Mating aggregations are concentrations of copulating individuals, in which competition between males for access to females occurs. Clusters are concentrations of inactive individuals (juvenile and adult) piled together in close contact. All types of aggregation were concentrated in only a part of what appeared to be the suitable habitat. Most aggregations' boundaries shifted over time within the habitat. We conclude that intraspecific attraction must be an important factor which maintains the spatial structure of all aggregations, although habitat specificity sets the larger area in which they move. Processes which may underlie this pattern, and some of its implications, are discussed.     

Encapsulated development of the marine prosobranch gastropod Nucella lapillus

Nucella (=Thais) lapillus (L.) develops to the juvenile stage within an egg capsule. This study was undertaken to investigate nutritive and bacteriostatic properties of the intracapsular fluid for this species. Capsules were collected intertidally at Nahant, Massachusetts, USA, in the summers of 1982 and 1983. The average capsule contained only 1.1 l of intracapsular fluid per embryo. A statistically significant correlation was observed between capsule height and number of developing embryos contained therein, although the number of embryos per capsule was more closely related to capsule volume. Total dry weight of encapsulated, shelled embryos increased exponentially as a function of shell length. However, weight increases may be entirely due to shell calcification; the average dry tissue weight of shelled embryos was significantly below that for pre-shelled individuals, suggesting a net loss of biomass during development. Aliquots of intracapsular fluid failed to inhibit the growth of the 13 bacterial strains tested. Despite the above results, pre-shelled embryos generally survived only several days following their premature removal from egg capsules at 12°C. Shelled veligers were reared outside of the capsules with no apparent ill effects. An explanation for these survivorship results is elusive.     

Life history of the temperate subtidal gastropod Umbonium costatum

The life history and reproductive biology of the trochid sand snail Umbonium costatum (Kiener) were investigated on a subtidal sandy shore in Hakodate Bay, Japan between 1988 and 1991. Female U. costatum mature at 1 yr of age (shell diameter-11 mm), reproduce twice (in June-July and September-October) in successive years, grow to a maximum size (shell diameter=22 mm) at age 8 yr, increase annual fecundity with age from 2000 (age 1 yr) to 37000 (age 8 yr), and show a maximum monthly gonad somatic index of 8% which is constant among ages. In comparison to a previously studied life history of a tropical Umbonium vestiatium, temperate U. costatum shows more sustained growth and a longer life span after maturation. This could be explained by: (1) the optimal size model concerned with resource investment, in gametes (Sebens 1987); and by (2) bet hedging to compensate large variability in larval success at high latitudes. These two hypotheses are not mutually exclusive, but both are based on season-related extremes of environment at high latitudes where the period suitable for reproduction is short.