A comparison of the reproductive cycles of Modiolus modiolus (L.), Cerastoderma (=Cardium) edule (L.), and Mytilus edulis L. in Strangford Lough,Northern Ireland

Summary This paper examines the reproductive cycles of three ecologically important marine bivalves-Modiolus modiolus (L.), Cerastoderma edule (L.), and Mytilus edulis L. in Strangford Lough, Northern Ireland over a period of almost five years.Whilst Modiolus does not appear to become sexually mature until it is several years old, Mytilus and Cerastoderma can reproduce in their first and second years of life respectively. In Cerastoderma and Modiolus sexual maturity is preceded by a period of rapid somatic growth.The subtidal Modiolus population remained in a more or less fully ripe condition virtually throughout the period of this investigation suggesting that this particular population lacked any marked cyclical reproductive activity. We interpret this as evidence of slow but almost continuous release of gametes throughout much of the year, a suggestion which is supported by recruitment data. A small intertidal population of Modiolus in Belfast Lough monitored over a period of two years exhibited a much more seasonal cycle. Here spawning occurred mainly during the autumn and winter. These data suggest that localised environmental factors are exceedingly important in controlling the annual reproductive cycle of this species. Cerastoderma from the mid-tidal sand flats ripened rapidly during the spring and spawned over a relatively restricted period in the summer. In Strangford Lough Mytilus occurs predominantly in the low-shore and while it spawns mainly in the spring and summer the annual cycle is considerably more protracted and variable than in Cerastoderma.Variations in the duration of the spawning periods in these bivalves can perhaps be explained in terms of both environmental stability and the immediate physical conditions experienced by these particular populations. The reproductive strategies exhibited by Cerastoderma, Modiolus, and Mytilus in Strangford Lough are considered in relation to population stability and to the different patterns of mortality which characterise these species in their respective local habitats.     

Effect of salinity on germination,seedling emergence,seedling growth and ion accumulation of a euhalophyte Suaeda salsa in an intertidal zone and on saline inland

The effect of salinity on germination, seedling emergence, seedling growth and ion accumulation of a euhalophyte Suaeda salsa L. in an intertidal zone and on saline inland were investigated. Brown seeds of S. salsa were heavier and better developed than black seeds in both the intertidal zone and on saline inland. The brown seeds/black seeds ratio for S. salsa in the intertidal zone was much higher than that for S. salsa on saline inland. More germinated seeds grew as seedlings under high salinity for S. salsa from the intertidal zone than S. salsa on saline inland; high salinity decreased the shoot length more severely for S. salsa from saline inland than for S. salsa from the intertidal zone; the seedling growth at a range of NaCl, measured either as shoot length or shoot dry weight, for S. salsa from the intertidal zone was lower than that of S. salsa from saline inland. In conclusion, for S. salsa from the intertidal zone there appears to be selection for slower growth and producing more brown seeds. The establishment of populations of S. salsa in different saline environments depends on the responses of seed germination, seedling emergence and seedling growth to salinity. These characteristics may determine the natural distributions of S. salsa populations in different saline environments.     

Water loss from aerially exposed mussels

The valve activity and water loss from aerially exposed Mytilus edulis L. and Modiolus modiolus (L.) have been investigated in a series of laboratory experiments. Mytilus is capable of maintaining long periods of complete, or almost complete, valve closure when exposed in air, and this allows the retention of water in the mantle cavity and protects the tissues from evaporative water loss. Over periods of three days or more the amount of water lost from emersed Mytilus was found to be less than that retained in the mantle cavity at the beginning of exposure, suggesting that during normal periods of exposure the tissues are never directly subject to water loss. In contrast, Modiolus shows periods of gaping during which water loss is rapid due to drainage from the mantle cavity and evaporation from the tissues. The exposure of the tissues to air that results from gaping, makes the water loss susceptible to environmental influences of which wind was found to be the factor which caused the greatest increase in water loss and, as a consequence, an important factor in causing death through dehydration. The different abilities of Mytilus edulis and Modiolus modiolus to control water loss may be related to their intertidal distributions.     

Osmotic changes in an estuarine bivalve,Modiolus fluviatilis

• 1.1.Responses to different salinities monitored by opening and closing of the shell valves were observed in Modiolus fluviatilis.
• 2.2.The osmotic pressure, sodium and chloride ion concentrations were measured in the haemocoelic fluid of Modiolus fluviatilis under similar conditions.
• 3.3.Free amino acids (measured as ninhydrin-positive substances) were determined in the muscle tissue of Modiolus.
• 4.4.It appears that these free amino acids are involved in the ability of the estuarine bivalve Modiolus fluviatilis to osmoregulate in a wide range of salinities.

     

Root morphology is related to the phenotypic variation in waterlogging tolerance of two populations of Suaeda salsa under salinity

The effects of waterlogging and salinity on seedling emergence, seedling growth and ion accumulation in a euhalophyte Suaeda salsa in an intertidal zone and on saline inland soil were investigated. Seedlings of S. salsa from the intertidal zone emerged more rapidly than those of the inland population under both waterlogged and drained conditions. Waterlogging and salinity had no adverse effects on seedling emergence of S. salsa from the intertidal zone, but markedly inhibited this parameter in the inland population. Waterlogging did not affect the seedling survival, shoot dry mass, and shoot height in high salinity in S. salsa from the intertidal zone, while the opposite trend was shown in the inland population. The root dry mass was higher in S. salsa from the intertidal zone as compared to the inland population, in waterlogged treatments by 1.9, 1.3, and 1.5 times in 1, 200, and 600 mM NaCl, respectively, and in drained treatments by 1.8, 2.3, and 3.0 times in 1, 200, and 600 mM NaCl, respectively. Waterlogging increased Na⁺ and K⁺ concentrations in high salinity, but waterlogging had no effect on Cl^- concentration in shoots of S. salsa from the intertidal zone. In all NaCl treatments, waterlogging had no effect on concentrations of these ions in shoots of S. salsa from the saline inland site. In a field investigation, the fresh mass of shoots and roots were lower, whereas the root/shoot ratio was 1.5 times higher in S. salsa from the intertidal zone, compared with the inland population. These findings indicate that S. salsa population from the intertidal zone is more waterlogging tolerant than the inland population. S. salsa from the intertidal zone produced relatively more root biomass and this might help anchor plants against tidal action in the intertidal zone. The physiological and morphological characteristics may determine the natural distributions of the two S. salsa populations in their different saline environments.     

Induction of Increased Salt Tolerance in Sorghum bicolor by NaCl Pretreatment

Following 20 d of exposure to 75 or 150 mol m^–3 NaCl Sorghumbicolor (L.) Moench plants become capable of growing in mediumcontaining 300 mol m^–3 NaCl. Control plants, which havenot been pretreated, or plants pretreated for less than 20 ddie within 2 weeks when exposed to 300 mol m^–3 NaCl. Weconsider this induction of a capacity to survive in and toleratea high NaCl concentration as an adaptation to salinity. We suggestthat adaptation to salinity is more than osmotic adjustmentand that it takes longer to develop than osmotic adjustment.Concomitantly with the appearance of the ability to grow inhigh salinity, adaptation also comprises the development ofa capacity to regulate internal Na⁺ and Cl^– concentrations,even when external salinity is high. Shoot mean relative growthrates are similar for both control plants and for adapted plantsgrowing in 300 mol m^–3 NaCl, although their shoot Na⁺and Cl^– concentrations are quite different. Based on thesedata, we propose that adaptation of Sorghum to high salinityresults from a modulation of genome expression occurring duringextended exposure to non-lethal NaCl concentrations. Key words: Sorghum bicolor (L.) Moench, NaCl, salt tolerance, adaptation to salinity     

Flexible Life History Strategies of Ninespine Sticklebacks,Genus <Emphasis Type="BoldItalic">Pungitius</Emphasis>

Synopsis We studied the life histories of the ninespine sticklebacks, Pungitius pungitius and Pungitius tymensis, collected from Japanese freshwater and brackish (sea) water habitats by examining the strontium (Sr) and calcium (Ca) concentrations in their otoliths. The Sr:Ca ratios in the otoliths changed with the salinity of the habitat regardless of identification as freshwater or brackish water type based on morphological characteristics. The ninespine sticklebacks living in a freshwater environment showed consistently low Sr:Ca ratios throughout the otolith. These samples were identified as a standard freshwater type. In contrast, all freshwater-type fishes collected from the intertidal zone showed higher otolith Sr:Ca ratios than those in the standard freshwater type, and the ratios fluctuated with the growth phase. All brackish water-type fishes collected in the intertidal zone showed the highest otolith Sr:Ca ratio throughout the otolith. In the present study, besides the two representative life history types of P. pungitius, i.e., freshwater and brackish water life history types, other sticklebacks had an anadromous life history type. These findings clearly indicate that the ninespine stickleback has a flexible migration strategy with a high degree of behavioral plasticity and an ability to utilize the full range of salinity in its life history.     

Effect of salinity change on cardiac activity in Hiatella arctica and Modiolus modiolus, in the White Sea

A great majority of salinity studies have dealt with intertidal species. Little is known about the way subtidal animals respond to salinity fluctuations. Even less details are available on invertebrates from the White Sea, which salinity is ca. 25. The heart rate of two subtidal Bivalvia—Hiatella arctica and Modiolus modiolus—exposed to different salinities was recorded. Changes in cardiac activity were monitored for 9 days of the animals’ acclimation to salinities of 15, 20, 30 and 35, and for 4 days of reacclimation (return to the initial salinity of 25). The initial response to salinity change was a significant heart rate reduction. On the other hand, cardiac activity in M. modiolus intensified at salinities of 30 and 35. Reacclimation induced different HR responses: from a decrease to a rise, depending on the species and the salinity applied in the experiment. The differences in responses to salinity are discussed with respect to the morphological and ecological characteristics of the species.     

Exposure of the Cyanobacterium Synechocystis PCC6803 to Salt Stress Induces Concerted Changes in Respiration and Photosynthesis

In order to survive and to grow in the presence of a high salinity(550 mM NaCl) Synechocystis PCC6803 increases its energeticcapacity. The salt-induced increase of electron transport ratesinvolves both cytochrome c oxidase and photosystem I. In contrast,electron transport rates measured through complexes I plus IIIof the respiratory chain, or through the photosystem II pluscytochrome b₆f complexes of the photosynthetic chain, do notshow appreciable changes. The time at which changes in electrontransport rates occur in the photosystem I and cytochrome coxidase complexes after the onset of salt stress indicates similaritiesin the adaptation of dark respiration and (cyclic) photosyntheticelectron flow. Given an increase of whole cell respiration andof PSI cyclic electron flow larger than the neosynthesis ofcytochrome aa3 and PSI reaction centers would predict, it appearsthat both adaptations require more than just synthesis of thesetwo complexes. (Received April 12, 1993; Accepted August 10, 1993)     

Effect of salinity on growth, ion accumulation and the roles of ions in osmotic adjustment of two populations of Suaeda salsa

The effect of salinity on growth, ion accumulation and the roles of ions in osmotic adjustment of two populations of Suaeda salsa were investigated. Seeds were collected from an intertidal zone or a saline inland zone in the Yellow River Delta in Shandong province, China. Seedlings were exposed to 10, 100, 200, 400 or 600 mM NaCl for 18 days in a greenhouse. NO₃ ^? concentration in the soil where S. salsa grows in an intertidal zone was much lower than that for the second population, but leaf NO₃ ^? concentration was the same in the two populations under field conditions. When plants were cultured in a greenhouse under natural light conditions, S. salsa from the intertidal zone showed fewer main stem branches and lower relative shoot growth compared to S. salsa from saline inland. Leaf Cl^? concentration of saline inland S. salsa was significantly higher than that of S. salsa from the intertidal zone, while the opposite was true for the concentration of NO₃ ^? in leaves of plants. For S. salsa from the intertidal zone NO₃ ^? contributed more than Cl^? to the osmotic potential, whereas S. salsa from the saline inland exhibited a reverse relationship under saline conditions, indicating that NO₃ ^? plays an important osmotic role in S. salsa from the intertidal zone in high salinity. In conclusion, S. salsa from the intertidal zone may employ superior control of ion uptake and content than S. salsa from the saline inland zone. The two populations of Suaeda salsa presented different ability in chloride exclusion and nitrate accumulation. These characteristics may affect the distributions of S. salsa in natural highly saline environments.     

Accumulation of ions during seed development under controlled saline conditions of two Suaeda salsa populations is related to their adaptation to saline environments

Controlled conditions were used to investigate the relationship between ion distribution in developing seeds of two Suaeda salsa populations and seed germination and seedling emergence. Seeds were harvested from S. salsa plants that had been treated with 1 or 400 mM NaCl for 122 (saline inland population) or 135 days (intertidal zone population) in a glasshouse. Germination and seedling emergence were evaluated under salinity. In both populations, more ions were accumulated in the pericarps of plants cultured in 400 mM NaCl than in 1 mM NaCl. Pericarps accumulated much higher ion concentrations in the intertidal zone population than in the saline inland population, while the opposite trend occurred for ion accumulation in the embryos. Seeds of plants from the intertidal zone population germinated more rapidly than those from plants of the saline inland population, regardless of the NaCl concentration during seed germination. However, seedling emergence under high salinity was lower with seeds from the intertidal zone population than with seeds from the saline inland population. In conclusion, S. salsa in the intertidal zone employs superior control of ion compartmentalization in the pericarps to tolerate salinity but requires a minimal level of ions in embryos to ensure seedling establishment in highly saline environments. This indicates that euhalophytes require salts during the mature seed stage to maintain seed viability and to ensure seedling emergence and population establishment.     

Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus petrolisthes

Vertical zonation of intertidal organisms, from the shallowsubtidal to the supralittoral zones, is a ubiquitous featureof temperate and tropical rocky shores. Organisms that livehigher on the shore experience larger daily and seasonal fluctuationsin microhabitat conditions, due to their greater exposure toterrestrial conditions during emersion. Comparative analysesof the adaptive linkage between physiological tolerance limitsand vertical distribution are the most powerful when the studyspecies are closely related and occur in discrete vertical zonesthroughout the intertidal range. Here, I summarize work on thephysiological tolerance limits of rocky intertidal zone porcelaincrab species of the genus Petrolisthes to emersion-related heatstress. In the eastern Pacific, Petrolisthes species live throughouttemperate and tropical regions, and are found in discrete verticalintertidal zones in each region. Whole organism thermal tolerancelimits of Petrolisthes species, and thermal limits of heartand nerve function reflect microhabitat conditions. Speciesliving higher in the intertidal zone are more eurythermal thanlow-intertidal congeners, tropical species have the highestthermal limits, and the differences in thermal tolerance betweenlow- and high-intertidal species is greatest for temperate crabs.Acclimation of thermal limits of high-intertidal species isrestricted as compared to low-intertidal species. Thus, becausethermal limits of high-intertidal species are near current habitattemperature maxima, global warming could most strongly impactintertidal species.     

Directional orientation in blue crabs, Callinectes sapidusRathbun: Escape responses and influence of wave direction

Blue crabs invade the upper intertidal at high tide and exhibit rapid, directed locomotory movements (i.e., escape response) which are significantly oriented offshore. The direction of wave surge and beach slope are conservative features of the intertidal, indicating offshore direction and can be used by blue crabs as orientation cues. Temporarily blinded crabs in the intertidal zone oriented down-slope (2–3°) and into the wave approach (up-wave) in experimental field test arenas. Up-wave orientation acuity increased in the presence of larger waves. Experiments in a wave test tank showed that orientation occurred over a wide range of surge velocities dependent upon threshold levels of surge displacement. We believe that the oriented response to wave surge serves as an important adaptation for movements by blue crabs in the nearshore zone.     

Differential predation on mangrove propagules in open and closed canopy forest habitats

Amounts of seed predation by grapsid crabs (Brachyura: Grapsidae) on two species of mangroves (Aegiceras corniculatum and Avicennia marina) were compared among different habitats in an Australian mangrove forest. For Avicennia, comparisons were between canopy gaps and the adjacent forest understory for six, mid intertidal, gaps of different sizes. For Aegiceras the comparisons were among canopy gaps in the high intertidal; open, accreting mud/sand banks where mangroves were colonizing in the low intertidal; and in the forest understory in both the high and low intertidal zones. These were repeated in the high salinity (35\%) downstream portion and the low salinity (0–5\%) upstream portion of a tidal river.Predation on Avicennia was significantly higher in the understory than in adjacent canopy gaps. Within a canopy opening, predation was greatest in the smallest gaps and lowest in the largest gaps. Predation on Aegiceras was greater in the high intertidal compared to the low intertidal, but no differences were found between river mouth and upstream locations. In the high intertidal zone of the forest, there were no differences in predation between canopy gap or forest understory sites for Aegiceras. In the low intertidal zone, however, significant differences in amount of predation were found between habitats. More Aegiceras propagules were consumed in the understory than on adjacent accreting sandbanks.Frequency of tidal inundation, which in turn affects the amount of time available to forage, is hypothesized to account for differences in predation between low and high intertidal forests and between small and large canopy gaps. Our results also suggest that shade intolerance in these two species may actually reflect an escape from predators, successful when the seeds are dispersed into open areas such as canopy gaps or mud banks.     

The Physiological Ecology of the Zebra Mussel, Dreissena polymorpha, in North America and Europe

SYNOPSIS. The zebra mussel, Dreissena polymorpha (Pallas), wasintroduced into North America in 1986. Initial North American(N.A.) studies suggested that physiological responses variedbetween N.A. and European populations. However, literature reviewindicates agreement on most aspects of physiological adaptationincluding: respiratory responses; hypoxia/anoxia tolerance;salinity limits; emersion tolerance; freezing resistance; environmentalpH limits; calcium limits; starvation responses; and bioenergeticpartitioning. The main differences among N.A. and European musselsappear to be elevated upper thermal limits and temperaturesfor optimum growth among N.A. populations. N.A. zebra musselsprobably originated from the northern shore of the Black Seain the warmest portion of the mussel's European range. However,most European physiological data come from northern Europe wherepopulations may be adapted to colder temperatures. Alternatively,N.A. research suggests that mussels may have a capacity forseasonal temperature acclimatization such that responses recordedin warmer N.A. waters may be different from those recorded innorthern Europe even after short-term laboratory acclimation.Studies of genetic variation and physiological response amongEuropean and N.A. D. polymorpha populations are required toelucidate the basis for physiological differentiation. Recentlyevolved D. polymorpha has poor resistance adaptations comparedto unionacean and sphaeriid bivalves with longer freshwaterfossil histories. Poor resistance adaptations make it less suitedfor stable habitats, instead, its high fecundities, early maturity,and rapid growth are adaptations to unstable habitats whereextensive resistance adaptations are of little value.     

Induced variability during the process of adaptation in Sorghum bicolor

The coefficient of variation of some anatomical and physiologicalparameters has been measured in plants of Sorghum bicolor (L.)Moench exposed to conditions inducing adaptation to salt. Thekinetic analysis of the variability of shoot DW and shoot Na⁺concentration revealed a specific phase of increase in variabilitybetween days 6 and 16 following exposure to salinity. The increasein variability observed in a population of adapted plants, ascompared to a population of control plants, appeared as a wholeplant process since variability for most of the parameters werepositively correlated. Variability within plant populationsof 11 Sorghum genotypes, each presenting a different capacityfor adaptation, was determined during the middle of the adaptationprocess. A strong linkage between the level of adaptation andthe increase in variability was observed. The relatively lowlevel of variability observed in populations of Sorghum varietieswhich did not adapt to salinity suggested that the increasein variability was not a direct consequence of exposure to stressconditions, but that it was specifically related to the adaptationprocess. The data presented suggest the occurrence of modificationsat the, level of genome expression during the adaptation process. Key words: Adaptation, resistance, variability, salinity, Sorghum     

The physiological ecology of the supratidal amphipod Talorchestia longicornis

Physiology, behavior, habitat, and morphology are used to determine the degree of adaptation to life on land for amphipod species and systemization within the four functional groups of the family talitridae. Talorchestia longicornis is a semi-terrestrial amphipod found in the supratidal zone of estuaries. The present study investigates the physiological adaptations of this species to life on land through measurements of osmoregulation and respiration. Over the salinity range of 1-40‰, T. longicornis regulated its hemolymph hyperosmotically at low salinities and hypoosmotically at high salinities. The isosmotic point was about 27‰. Analogously, hemolymph chloride levels were well regulated being hyperionic at low salinities and hypoionic at high salinities. This species is capable of respiration in both air and water. Slopes (b values) of the relationship between weight and oxygen uptake rates ranged from 0.316 to 0.590. Oxygen uptake rates were higher in air than water and at night versus day. Q₁₀ values were slightly below 2.0 for respiration in air for amphipods, irrespective of weight. These physiological adaptations, along with its behaviors, habitat, and morphology, place T. longicornis within the Group III sandhoppers of the Talitridae.     

A Nutritional Role for Amino Acid Transport in Filter-Feeding Marine Invertebrates

SYNOPSIS. Filter-feeding invertebrates are exposed to a smallbut continuous source of potential foodstuffs in the form ofdissolved free amino acids (DFAA). Studies with two genera ofbivalve molluscs, Mytilus and Modiolus, indicate the existenceof carrier-mediated transport processes for DFAA, and that theseprocesses are capable of a net accumulation of substrate fromexternal solution. The kinetics of DFAA transport in bivalvegill, the primary organ for uptake, reveal these processes tobe well-adapted for the efficient utilization of substrate atthe extremely low concentrations (approximately 1 µM)characteristic of the water column: Michaelis constants foruptake are in the micromolar range, resulting in transport ratesthat are a significant fraction of the maximum capacity of thesystem. Comparison of potential rates of DFAA accumulation torates of oxidative metabolism suggest that DFAA can serve asan important nutritional supplement to these animals. Futurework should emphasize new analytical procedures, including theuse of high performance liquid chromatography, to identify levelsof DFAA in specific environments and rates of net DFAA uptakeinto organisms from these habitats.     

Exogenous ABA as a Modulator of the Response of Sorghum to High Salinity

Treatment of Sorghum bicolor (L.) Moench, cv. 610, with abscisicacid (ABA) during the first week of sahnization with 150 molm^–3 NaCl induced enhancement of growth and acceleratedadaptation to high salinity (300 mol m^–3 NaCl) Adaptationis defined as the development of the ability of the plant tosurvive, grow, and set seeds upon exposure to a NaCl concentrationwhich is lethal for the unadapted plant In the absence of ABAthe saline pretreatment requires 20 d for the development ofadaptation (Amzallag et al., 1990), with ABA treatment the sameresult is achieved within approximately one week The exposureof the plants to non-lethal salinity (150 mol m^–3 NaCl)apparently triggers a transient sensitivity to ABA lasting forabout 8 to 10 d following the beginning of sahnization Thisperiod coincides with an increase in leaf PEP carboxylase activitywhich seems to occur faster if the plants are treated with ABA.Exogenous ABA-induced enhancement of growth and control of shootNa⁺ concentration, occur at a lower ABA concentration (10 mmolm^–3) than the induction of adaptation to salinity whichoc curs at 40 mmol m^–3 or above. The lowered shoot Na⁺concentration which is induced by a low ABA concentration isnot sufficient to induce survival of the plants in high salinity(300 mol m^–3 NaCl). Key words: Growth, adaptation to salinity, ABA