Salinity tolerance of benthic estuarine diatoms as tested with a rapid polarographic measurement of photosynthesis

Measurements of net photosynthesis of benthic estuarine diatoms were made by polarographic registration of oxygen saturation. A measuring cell was constructed in which media with salinities of 2.0 to 100.7 were pumped over the algae between measurements. Diatoms from unialgal cultures and mixed populations from intertidal flats appeared to be highly tolerant of extreme salinities. During short-term exposures (20 min) the net photosynthesis of the algae did not drop below 70% of the initial values, within the salinity range 4.0 to 60.0. Prolonged exposure (up to 6 h) gave essentially the same results. Populations of benthic diatoms, sampled from field stations with mean salinities of about 30, 12, and below 5, showed only gradual differences in their tolerance of salinities between 2.0 and 33.7. Two species, Navicula arenaria and Nitzschia sigma, were cultured in media ranging in salinity from 8.0 to 45.0 and from 2.0 to 45, respectively. The tolerance to changing salinity was only slightly affected by the salinity of the medium in the preculture. The role of salinity in the production and distribution of intertidal diatoms is discussed.     

Effects of salinity on respiration and nitrogen excretion in two species of echinoderms

The 30-d survival limit of Eupentacta quinquesemita and Strongylocentrotus droebachiensis is 12–13 S. The activity coefficient (1 000/righting time in seconds) of stepwise acclimated sea urchins declined from 16.3 at 30 S to 3.5 at 15 S. Oxygen consumption rates (QO₂) of both species held at 30 S and 13°C were highest in June and lowest in December. During the summer, when environmental salinity is most variable in southeastern Alaska, the QO₂ of both species held at 30, 20 and 15 S varied directly with salinity. Perivisceral fluid PO₂ varied directly with acclimation salinity in sea urchins, but not in sea cucumbers. Perivisceral fluid oxygen content of acclimated sea urchins was significantly lower at 15 and 20 S than at 30 S due to reduced PO₂ and extracellular fluid volume at the lower salinities. The QO₂ of both species varied directly with ambient salinity during a 30-10-30. semidiurnal pattern of fluctuating salinity. No change occurred in the average QO₂ of either species over a 15-30-15. semidiurnal pattern of fluctuating salinity. Sea urchin perivisceral fluid PO₂ declined as ambient salinity fluctuated away from the acclimation salinity in both cycles and increased as ambient salinity returned to the acclimation salinity. Total nitrogen excretion of stepwise acclimated sea cucumbers declined significantly from 30 to 15 S, but there was no salinity effect on total nitrogen excretion in sea urchins. Ammonia excretion varied directly with salinity in stepwise acclimated sea cucumbers (67–96% of total nitrogen excreted), but there was no salinity effect on ammonia excretion (89–95% of total nitrogen excreted) of sea urchins. Urea excretion did not vary with salinity in sea cucumbers (2–4% of total nitrogen excreted) or sea urchins (2–9% of total nitrogen excreted). Primary amines varied inversely with salinity in sea cucumbers (2–30% of total nitrogen excreted), but did not vary with salinity in sea urchins (2–4% of total nitrogen excreted). The oxygen: nitrogen ratio of both species indicated that carbohydrate and/or lipid form the primary catabolic substrate. The O:N ratio did not vary as a function of salinity. Both species are more tolerant to reduced salinity than previously reported, however, rates of oxygen consumption and/or nitrogen excretion are modified by salinity as well as season.     

Effects of salinity fluctuations on the respiration rate of the southern oyster drill Thais haemastoma and the blue crab Callinectes sapidus

Respiration rates of Thais haemastoma and Callinectes sapidus were determined as a function of salinity with a flow-through respirometer at 20°C. Respiration rates were measured at 10, 20 and 30 S for acclimated animals. The effects of 10-5-10, 20-10-20, 30-10-30 and 10-30-10 S semidiurnal cycles (12 h) of fluctuating salinity on the rate of respiration of the oyster drill were studied. During each cycle, salinity was changed from the acclimation salinity over a 4 h interval, held at that salinity for 2 h, returned to the acclimation salinity over 4 h and held at that salinity for 2 h. The effects of diurnal (24.8 h) salinity cycles on respiration in the oyster drill and blue crab were also studied. Salinity was changed from the acclimation salinity over a 10.4 h interval, held at that salinity for 2 h, then returned to the acclimation salinity over 10.4 h and held at that salinity for 2 h. The respiration rate of 30 S acclimated oyster drills (679 l O₂ g dry weight^–1 h^–1) was significantly higher than for individuals acclimated to 10 S (534 l O₂ g dry weight^–1 h^–1). Blue crab respiration was 170 l O₂ g dry weight^–1 h^–1 at 30 S, and was significantly higher at 10 and 20 S than at 30 S. With the exception of the 20-10-20 S semidiurnal cycle, the respiration rate of oyster drills declined as salinity fluctuated in either direction from the acclimation salinity and increased as ambient salinity returned to the acclimation salinity. Semidiurnal cycles (12 h) of fluctuating salinity produced greater changes in the respiration rate of snails than analogous diurnal cycles (24.8 h). A 10-30-10 S pattern of fluctuation caused a greater percentage reduction in the steady state respiration rate of oyster drills than the 30-10-30 S pattern. The respiration rate of blue crabs varied inversely with fluctuating salinity. Relatively minor changes occurred in blue crab respiration rate with fluctuating salinity. Blue crab respiration rate characteristically dropped during the initial phase of declining salinity at a rate directly proportional to the rate of salinity decrease, perhaps representing a metabolic adjustment period by the blue crabs. The respiratory response of T. haemastoma to salinity is consistent with its incomplete volume regulation, while the response of C. sapidus is compatible with its ability to regulate extracellular fluid osmotic and ionic composition.     

Survival and chloride ion regulation of the porcelain crab Petrolisthes armatus exposed to mercury

Acute toxicity bioassays conducted at various salinities demonstrated that mercury (as mercuric chloride) at low concentrations was lethal to Petrolisthes armatus. Ninety-six hour LC₅₀ values varied from 50 to 64 parts per billion (ppb) of mercury, depending on test salinities. Lower salinities. decreased the time to death of mercuryexposed crabs. Differences in survival after 96 h due to salinity were not statistically significant. Blood chloride concentrations were regulated hyperchloride to the medium at low salinities and hypochloride at high salinities by acclimated crabs. The salinity isochloride to blood was 20 S. Transfer of crabs from 15 S to salinities ranging from 7 to 35 S resulted in new steadystate chloride levels within 12 h. Exposure to 50 ppb mercury did not alter chloride ion regulation of either acclimated crabs or crabs adjusting to new salinities.     

Influence of salinity and temperature on the oxygen consumption in young juveniles of the Indian prawn Penaeus indicus

Routine oxygen consumption of very young juveniles (0.1 g) of Penaeus indicus H. Milne Edwards was significantly influenced by ambient temperature and weight of the animal, but not by ambient salinity, when tested at salinities (7, 21, and 35) to which they had been long-term (over 10 days) acclimated. Standard oxygen consumption of young juvenile prawns (1 to 3 g), subjected to step-wise changes in ambient salinity, from sea water to low salinity waters (2 to 6), and measured after short-term (24 h) salinity acclimation at each step, was lowest at salinities where prawns such as those tested occur naturally (10 to 15). The metabolic rates do not appear to have a direct relation to the osmotic gradient, even when the influence of interfering activity is eliminated. It appears that factors other than osmotic gradient will have to be sought in order to explain the metabolic patterns of P. indicus in relation to salinity.     

The significance of temperature,salinity and zinc as lethal factors for the mussel Mytilus edulis in a polluted estuary

Mussels, Mytilus edulis L., were subjected to high temperatures, low salinities and dissolved zinc in order to investigate possible environmental hazards of a discharge of heated effluent near Newport on the Yarra River estuary, Victoria, Australia. Exposure to zinc at 0.8 mg l^-1 for 14 d in otherwise favourable conditions significantly increased mortality resulting from subsequent exposure to temperatures between 29° to 31°C for 24 h without added zinc. Mussels collected from water of temporarily lowered salinity (8–16 S) showed significantly lower thermal resistance than controls collected from marine salinities (35 S). Mussels taken from a marine environment and exposed to 10 S died at a rate which increased with temperature. Mussels acclimated for 14 d to combinations of 10°, 16° and 22°C and 22 and 35 S, and subsequently exposed to increased zinc concentrations accumulated zinc to levels which were independent of temperature and salinity. The zinc was lethal more quickly at 22°C and 35 S than at the lower temperatures and salinities. The modes of toxic action of the salinity, zinc and temperature factors are discussed and it is argued that zinc which has been found accumulated in mussels near Newport could be reducing their resistance to raised temperatures and perhaps other stresses, probably as a result of effects on lysosomal functioning. The evidence suggests that the heated effluent will accelerate any toxic effects of zinc or low salinities which occur near Newport and so poses a hazard in winter as well as in summer.     

The combined effects of salinity and temperature on larvae of the mussel Mytilus edulis

The combined effects of salinity and temperature on survival and growth of larvae of the mussel Mytilus edulis (L.) were studied. The effects of salinity and temperature are significantly related only as the limits of tolerance of either factor are approached. Survival of larvae at salinities from 15 to 40 is uniformly good (70% or better) at temperatures from 5° to 20°C, but is reduced drastically at 25 °C, particularly at high (40) and low (20) salinities. Larval growth is rapid at a temperature of 15 °C in salinities from 25 to 35, at 20 °C in salinities from 20 to 35. Optimum growth occurs at 20 °C in salinities from 25 to 30. Growth decreases both at 25° and 10 °C; the decline is most drastic at high (40) and low (20) salinities.Part of a study completed at the Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut, USA, while on a UNESCO Fellowship.     

Effects of salinity gradients on the tolerance and bioenergetics of juvenile blue crabs (Callinectes sapidus) from waters of different environmental salinities

Juvenile Callinectes sapidus Rathbun were collected from brackish and hypersaline coastal environments in August 1986 and July 1987, respectively. The brackish collection site was a salt-marsh near Grand Isle, Louisiana (USA), and the hypersaline site was in the barrier island system on the north end of the Laguna Madre near Corpus Christi, Texas (USA). On the dates of collection, salinities fluctuated daily between 20 and 30 S and between 30 and 45 S at the brackish and hypersaline collection sites, respectively. The high-salinity 21 d LC₅₀ (50% mortality) was 56.0 for brackish-water individuals and 66.5 S for hypersaline individuals. The brackish-water individuals survived 0 S. The lowsalinity 21 d LC₅₀ was 0.5 S for the hypersaline individuals. Respiration and excretion comprised a small portion of the energy budget and did not vary with salinity for individuals from brackish water. However, both respiration and excretion increased with decreasing salinity in individuals from the hypersaline environment. Respiration accounted for more energy than excretion. As energetic expenditure (due to respiration and excretion) was relatively small, scope for growth usually paralleled energy absorption. Scope for growth responses to salinity differed significantly between crabs from the two environments. Peaks in scope for growth for both the brackish-water and hypersaline individuals corresponded to salinities normally encountered by these crabs in their natural habitats. Individuals from the brackish-water population had maximal energy absorption and scope for growth at 10 and 25 S. Individuals from the hypersaline population displayed maximal energy absorption at 35 S and maximal scope for growth at 35 and 50 S.     

Influence of salinity on embryonic development and the distribution ofSepia officinalis in the Delta Area (South Western part of The Netherlands)

The effect of salinity on embryonic development ofSepia officinalis (cuttlefish) in the Delta Area (South Western part of The Netherlands) was investigated in 1988/1989, and compared with data concerning the distribution ofS. officinalis in the three main parts of this area: Oosterschelde, Westerschelde and Grevelingen. Embryos hatched in water collected at Yerseke (Oosterschelde), Vlissingen (Western part of the Westerschelde) and Bommenede (Grevelingen), i.e., at salinity values above 28.1, but not in water sampled at Hoedekenskerke and Hansweert (Middle and Eastern part of the Westerschelde; salinities below 22.0). Under laboratory conditions, using diluted Oosterschelde water, the highest hatching percentages ofS. officinalis were found at salinities above 29.8. Some embryos hatched at a salinity value of 26.5 but no hatching occurred at salinities below 23.9. In embryos exposed to salinity changes during late embryonic development, the developmental rate decreased at salinity values of 28.7 or less. Below 22.4 embryos with morphological malformations were found. It can be concluded that salinity is an important factor limiting the distribution ofS. officinalis in most parts of the Delta Area, with the exception of the Western part of the Westerschelde and the Grevelingen.Contribution no. 489 of the Library of the Delta Institute for Hydrobiological Research     

Effects of salinity and temperature on Corophium volutator and C. arenarium (Crustacea: Amphipoda), with particular reference to distribution

The responses of the post-embryonic stages of Corophium volutator (Pallas) and C. arenarium Crawford to the combined effects of salinity and temperature show that gravid females have a wider tolerance than nongravid adult females which in turn are more tolerant than adult males. C. volutator is more tolerant of low salinity (2 to 10) than C. arenarium, but the latter is more tolerant of salinities above 45. The embryos of C. volutator develop normally and hatch at lower salinities and temperatures than those of C. arenarium, in which successful development was recorded at higher temperatures. Females undergoing a pre-copulatory moult failed to lay eggs below salinities of 3 (C. volutator) and 10 (C. arenarium), but in both species the lowest salinity at which all females moulted and laid eggs was 20. The results are discussed in relation to the distribution of both species.     

Effects of salinity and calcium concentration on arm regeneration byOphiothrix angulata (Echinodermata: Ophiuroidea)

At 33 salinity a tissue stump formed 2 to 3 d after autotomy and developing ossicles were present by the fourth day inOphiothrix angulata (Say). Regeneration proceeded rapidly from the sixth day until the thirteenth day, when the rate decreased greatly. The length of the regenerated arm and the number of ossicles formed did not vary over a salinity range of 28 to 38 S, but were significantly less at 23 S. The number of ossicles regenerated increased linearly (y=1.9 x-7.7;r=0.9089) with the calcium concentrations ranging from 3.8 to 9.5 mM. No ossicle formation occurred at 3.8 mM calcium concentration. Rate of net uptake of calcium-45 into the ossicles of intact individuals in salinities of 28 and 33 was significantly greater than that in 23 and 38 S. However, net uptake rate of calcium into the soft tissues of the arms was significantly higher at 18S than at the lower two salinities.     

Osmoregulation of the grass shrimp Palaemonetes pugio exposed to polychlorinated biphenyls (PCBs). I. Effecton chloride and osmotic concentrations and chloride-and water-exchange kinetics

Grass shrimp, Palaemonetes pugio, were capable of hypo- and hyper-osmotic regulation of body fluids. Hemolymph chloride and osmotic concentrations were maintained at relatively stable levels over a wide salinity range. Following an abrupt transfer from intermediate (14 and 17) to high (31 and 35) or low (1 and 2) salinities, hemolymph chloride levels exhibited initial overshoot and undershoot, respectively, of new steady-state levels. Osmotic concentrations exhibited an initial undershoot at low, but not overshoot at high salinity. Chloride space in salinity-acclimated shrimp was relatively stable at salinities from 1 to 35. Changes in chloride space following salinity transfer paralleled those of hemolymph chloride levels, and are discussed in the light of alterations in intracellular sodium concentrations reported earlier. Rate constants for chloride turnover indicated independent exchanges of sodium and chloride ions. Water-turnover measurements showed that permeability of P. pugio was greatest at the isosmotic salinity (17) and reduced at salinities which were associated with active osmoregulation. Exposure to sublethal and 96-h LC₅₀ levels of Aroclor^® 1254 did not seriously alter hemolymph chloride and osmotic concentrations, chloride space or chloride-exchange kinetics in adult shrimp. Disruption of hemolymph chloride regulation in juvenile shrimp was associated with large mortalities not observed in adults. Shrimp exposed to Aroclor 1254 at 17 S exhibited reduced water permeability similar to levels previously observed in controls at high and low salinities in response to osmotic or ionic gradients. Exposure to PCBs did not result in further reduction in permeability at the latter salinities.     

Effects of salinity and bromine on zygotes and embryos of Fucus vesiculosus from the Baltic Sea

Zygotes and young embryos derived from Fucus vesiculosus collected in the archipelgo of Stockholm in 1990, growing at a salinity of 6 to 7 S, were cultured under different salinity conditions and in media of different bromine concentrations. Optimum salinity was 10 to 12 S for germination (rhizoid initiation) while apical hair formation showed a broader tolerance curve with an optimum at 8 to 14 S. Bromine caused inhibition of early development of F. vesiculosus. At 6 salinity a 50% reduction in germination took place at 10.0 mM Br and at 1.25 mM Br only 4.7% of the embryos developed apical hairs, as compared to 32.7% in the control. Bromine toxicity decreased at higher salinities. The results indicate that F. vesiculosus in the Baltic Sea has diverged from its Atlantic progenitors and to some extent acclimated to low salinity. Still, the salinity in the normal environment of the tested population is lower than optimum, leading to a lower degree of germination of zygotes, a lower growth rate of young embryos and probably also a higher sensitivity to additional stress factors such as chemical pollution.     

Acclimation of gametes to reduced salinity prior to spawning inLuidia clathrata (Echinodermata: Asteroidea)

AdultLuidia clathrata (Say) were held at salinities lower and higher (25 to 35) than environmental salinity (29) during gametogenesis and after sperm and primary oocytes were produced. The sperm and primary oocytes acclimated to the experimental salinities in both cases. Fertilization and development through gastrulation were not affected at the salinities at which the parents were held, but were affected at altered salinities. This indicates that intracellular osmoregulation can occur both during gametogenesis and after the stage of arrested activity has been reached. This is an important capacity forL. clathrata, which lives in estuaries with variable salinities.     

Effects of salinity fluctuation on the hemolymph composition of the blue crab Callinectes sapidus

The hemolymph of the blue crab Callinectes sapidus was hyperosmotic during 20-10-20 S and 30-10-30 S diurnal cycles. The hemolymph became isosmotic at 26 S and hyposmotic at 28 S in the 10-30-10 S diurnal cycle. Hemolymph Na⁺ was hyperionic to seawater throughout all cycles. Hemolymph Cl^- was hyperionic below 24 S and either isionic or hypoionic from 24 to 30 S. Hemolymph K⁺ concentrations were hyperionic below 26 S and either isionic or hypoionic from 26 to 30 S. Hemolymph Mg⁺⁺ values were hypoionic over the experimental salinity range (10 to 30). Hemolymph ninhydrin-positive substances (NPS) levels were directly related to ambient salinity.     

Changes of plasma osmolality,chloride concentration and gill Na−K-ATPase activity in tilapia Oreochromis mossambicus during seawater acclimation

Changes of plasma osmolality, chloride concentration and gill Na–K-ATPase activity in tilapia Oreochromis mossambicus (obtained from Tainan Fish Culture Station of Taiwan Fisheries Research Institute, 1987) during seawater acclimation were examined. Three experiments were performed. (1) Freshwater (FW) to 30 salinity seawater (SW): plasma osmolality and chloride rose violently immediately post-transfer. At 6 h, gill Na–K-ATPase activity began to increase but most fish died from excessive plasma osmolality and Cl. (2) FW to 20 salinity SW: plasma osmolality and chloride increased immediately post-transfer, but more slowly than in (1), and began to decrease at 24 h. However it was not until 12 h post-transfer that gill Na–K-ATPase activity rose slowly. (3) FW to 20 salinity SW for 24 h, then to 30 salinity SW: after transfer to 30 salinity, plasma osmolality and chloride showed only a small increase initially then declined, while gill Na–K-ATPase activity started to rise rapidly within 3 h. The present results coincided with our previous morphological data concerning the ultrastructural responses of gill chloride cells. These are discussed to elucidate the osmoregulation mechanisms in tilapia during seawater acclimation.     

Osmotic and ionic regulation in the fiddler crab Uca rapax acclimated to dilute and hypersaline seawater

Adult male Uca rapax, collected from the central coast of Venezuela in early 1994 were gradually acclimated to salinities ranging from 1.7 to 139S. The hemolymph osmolality (791±15 mOsmol in crabs from 26S) changed less than three-fold over the entire range of concentrations tested. The urine was isosmotic with the hemolymph in crabs exposed to dilutions <26S, and significantly hyperosmotic in those exposed to media >34.8S. The hemolymph levels of Na⁺, Cl^–, K⁺, Ca²⁺ and Mg²⁺ (320±13, 405±17, 7.8±0.7, 7.2±0.1 and 31±2.2 mmol l^–1, respectively, in crabs acclimated to 26S) were maintained fairly constant over the range from 8.7 to 99S, decreasing by 15% in the more dilute media or increasing sharply to about twice those values in crabs from 139S. The excretory organs contributed to the osmoionic regulation of the hemolymph in crabs exposed to <3.5 or to >34.8S, by means of the partial reabsorption or excretion, respectively, of salts from or into the urine. The results described place U. rapax among the most powerful hypo/hyper-regulating crustaceans known.     

Responses of Leptasterias hexactis (Echinodermata: Asteroidea) to low salinity

The six-rayed starfish Leptasterias hexactis (Stimpson, 1862) is seasonally exposed to low salinities in southeastern Alaska. Individuals that were gradually exposed to reduced salinities in the laboratory had a 28-d TLm of 12.9 S. The activity of L. hexactis, as measured by its activity coefficient, varied directly with salinity. Individual feeding rates of the starfish on similarly exposed Mytilus edulis, measured daily for 21 d at salinities of 30, 20 and 15 S, also varied directly with salinity. The dry weight of mussel tissue consumed was 8.84, 8.49 and 0.58 mg ·starfish^-1·d^-1 at 20, 20 and 15 S. Expressed as percent of dry starfish weight, the daily feeding rate was 1.35, 0.76 and 0.10% at 30, 20 and 15 S. Absorption efficiency decreased from 64% at 30 S to 49% at 20 S, further reducing the energy available for metabolism. Growth, measured in terms of changes in total dry weight or dry weight of soft tissues, also varied directly with salinity. Although exposure to hyposmotic conditions did induce stress responses, as indicated by reductions in activity, feeding, absorption efficiency and growth rates, L. hexactis maintained positive growth for at least a 3-wk period in the laboratory at 20 S and 13°CC. The population of L. hexactis investigated must be considered euryhaline and brief periods of exposure to hyposmotic conditions should not limit its distribution.     

Salinity dependence of sexual and asexual reproduction in the rotifer Brachionus plicatilis

A salinity dependent mictic response was observed in a clone of Brachionus plicatilis cultured in the 2 to 4 salinity range. This response was related to asexual exponential reproduction rates (G) and could be divided into three categories: (a) no mixis occurred at a salinity of 35 S and above, where G values were lower than 0.30 d^-1, (b) low mictic levels in rotifers cultured at 2 and 30 S, where G values ranged between 0.40 to 0.50 d^-1, and (c) high mictic levels in rotifers cultured at salinities ranging between 4 and 20 S, where G values ranged between 0.50 to 0.85 d^-1. Fluctuations in mictic levels varied with time during the course of the experiments. Results suggest that salinity conditions leading to optimal parthenogenic reproduction also support mixis.     

Na<Superscript>+</Superscript>/K<Superscript>+</Superscript>–ATPase activity and gill ultrastructure in the hyper-hypo-regulating crab<Emphasis Type="Italic"> Chasmagnathus granulatus</Emphasis> acclimated to dilute,normal, and concentrated seawater

We studied Na⁺/K⁺ –ATPase activity and ultrastructure in gills of the hyper-hypo-regulating crab Chasmagnathus granulatus Dana, 1851 acclimated to different salinities: 10, 30 and 45, known to be hypo-, iso-, and hyper-osmotic to the hemolymph, respectively. After centrifugation of homogenates at 11,000 g, Na⁺/K⁺–ATPase activity was almost entirely found in the pellets from the posterior (6–8) and anterior (3–5) gills, whereas very little was detected in the supernatant liquid. Specific activity of gill 6 was 41.3, 30.2, and 28.2 µmol Pi h^–1 mg prot^–1 for crabs acclimated to 10, 30, and 45, respectively, the result for 10 being significantly higher than those at 30 and 45. Although the concentration of sodium at which the reaction rate is half-maximal (K _M) was similar in the three acclimation salinities, only the enzyme from crabs acclimated to 10 was inhibited by high sodium concentration. Specific activity of gill 5 increased with the increment in external salinity (10.1, 15, and 18.1 µmol Pi h^–1 mg prot^–1 for 10, 30, and 45, respectively), the only significant difference being that between the extreme salinities. The epithelium thickness of the dorsal portion of gill 6 showed a variation among salinities: 21.7, 15.8 and 17.2 µm for 10, 30 and 45, respectively. There were significant differences in epithelium thickness between the 10 and the other salinities. In all three salinities, the ultrastructure of gill 6 epithelium showed a high density of mitochondria, estimated by their volume fraction (Vv _m=0.307–0.355). These mitochondria were packed between extensive basolateral membrane interdigitations in ionocytes and pillar cells. Gill 5 showed three cell types: pillars which possess mitochondria packed between membrane folds only in their interdigitations with neighbouring cells; type-I cells 8.0 µm thick with low density of mitochondria (Vv _m=0.088), and type-II cells, 9.9 µm thick and rich in mitochondria (Vv _m=0.423), but lacking basolateral interdigitations. Vv _m of type-I cells of gill 5 was significantly lower than those of type-II cells of the same gill and the ionocytes of gill 6. No significant difference in Vv _m was detected between the latter cell types.Communicated by P.W. Sammarco, Chauvin