Effects of direct transfer from freshwater to seawater on respiratory and circulatory variables and acid-base status in rainbow trout

Summary The effects of increased ambient salinity (35 mg · ml^-1) were studied at 1, 6, and 24 h after direct transfer of rainbow trout from freshwater to seawater. Two series of experiments were carried out successively. The first series was designed to simultaneously study all the respiratory (except Hb affinity for O₂), circulatory, and acid-base variables in each fish. In this series, fish were fitted with catheters chronically inserted into the cardiac bulbus, the dorsal aorta, and the opercular and buccal cavities. In the second series, designed to study haemoglobin O₂ affinity, fish were fitted with only a dorsal aorta catheter. The ventilatory flow ( ) was markedly increased just after transfer (by 55% at 1 h), then more moderately (by 20% at 6 h and 32% at 24 h). The initial hyperventilation peak was associated with frequent couphing motions. These ventilatory changes resulted essentially from increase in ventilatory amplitude. Initially, standard oxygen consumption (MM}O₂) decreased slightly, the moderately increased (by 12% at 24 h), so that the oxygen convection requirement ( ) increased substantially. In spite of an increased ventilation, the partial pressure of oxygen in arterial blood (P _aO₂) decreased slightly at 1 h, prior to returning to control levels, while partial pressure of carbon dioxide in arterial blood (P _aCO₂) was not significantly decreased. Gill oxygen transfer factor decreased substantially at 1 h (by 35%) then more moderately (by 7% at 1 h and 12% at 24 h). These results suggest a decrease in gas diffusing capacity of the gills. As P _aCO₂ remained approximatively unchanged, the gradual decrease in arterial pH (pH_a) from 7.94 to 7.67 at 24 h must therefore be regarded as a metabolic acidosis. The strong ion difference decreased markedly because the concentration of plasma chloride increased more than that of sodium. Arterial O₂ content (C _aO₂) gradually decreased (by 38% at 24 h) simultaneously with the decrease in pH_a, while the ratio P _aO₂/C _aO₂ increased. In parallel, seawater exposure induced a marked decrease in affinity of haemoglobin for O₂, so that at 24 h, P₅₀ was increased by 26% above the value obtained in freshwater-adapted trout. The increase in could be ascribed initially (at 1 h) to the decrease of P _aO₂ and later to a stimulation of respiratory neurons resulting from the lowered medullary interstitial pH. The decrease in C _aO₂ could be interpreted mainly as a consequence of a decreased affinity of haemoglobin for O₂, likely to be due to the blood acidosis and a predictable increase in chloride concentration within erythrocytes. Cardiac output ( ) slightly decreased at 1 h, then progressively increased by 30% at 24 h. Branchial vascular resistance increased at 1 h by 28%, then decreased by 18% of the control value at 24 h. Systemic vascular resistance decreased markedly by 40% at 24 h. As heart rate (HR) remained significantly unchanged, the cardiac stroke volume initially decreased then increased in relation to the changes in . The increase of , allowing compensation for the effect of decreased C _aO₂ in tissue O₂ supply, was interpreted as a passive consequence of the decrease in total vascular resistance occurring during seawater exposure.Abbreviations a.u. arbitrary units - C _aO₂ arterial oxygen content - pH₅₀ arterial pH at P₅₀ - C _vO₂ venous oxygen content - Hb haemoglobin - HR heart rate - Hct hematocrit - n_Hill Hill coefficient - O₂ standard oxygen consumption - P _aCO₂ arterial partial pressure of carbon dioxide - P _aO₂ arterial partial pressure of oxygen - P _vO₂ oxygen partial pressure in mixed venous blood - P₅₀ oxygen tension at half saturation of haemoglobin - P _VA, P _DA blood pressure in ventral and dorsal aorta - pH_a arterial pH - PIO₂, PEO₂ oxygen partial pressure of inspired and expired water - PO₂ oxygen partial pressure - cardiac output - SEM standard error of mean - S.I.D. strong ion difference - SV cardiac stroke volume - TO₂ gill oxygen transfer factor - U oxygen extraction coefficient - VA ventilatory amplitude - VF ventilatory frequency - VR_G, VR_S branchial and systemic vascular resistances - ventilatory flow - ventilatory oxygen convection requirement     

Haemoglobin function and respiratory status of the Port Jackson shark,<Emphasis Type="Italic"> Heterodontus portusjacksoni</Emphasis>, in response to lowered salinity

Haemoglobin function and respiratory status of sub-adult sharks, Heterodontus portusjacksoni was investigated for up to 1 week following transfer from 100% to either 75% or 50% seawater. Metabolic rates were unusually low and arterial–venous differences in blood O₂ small. Haemodilution from osmotic inflow lowered haematocrit and reduced blood O₂ content by up to 50%. There was no change in O₂ consumption rate, blood O₂ partial pressure, cardiac output, or the arterial-venous O₂ content difference, and thus O₂ delivery was maintained. Ventilation was acutely elevated but returned to normal within 24 h. The O₂ delivery to the tissues was facilitated by decreased blood O₂-affinity that could not be simply ascribed to changes in the osmolyte concentration. The Hb was unaffected by changes in intra-erythrocyte fluid urea or trimethylamine-N-oxide (TMAO) but was sensitive to changes in NaCl. The Bohr shifts in whole blood were low and there was little role for pH in modulating O₂ transport. Venous Hb saturation remained close to 65%, at the steepest part of the in vivo O₂ equilibrium curve, such that O₂ unloading could be facilitated by small reductions in pressure without increasing cardiac or ventilatory work. H. portusjacksoni tolerated 50% seawater for at least 1 month, but there was little evidence of respiratory responses being adaptive which instead appeared to be consequential on changes in osmotic and ionic status.Abbreviations a–v arterial–venous - CO ₂ CO₂ content - C _a O ₂ content of O₂ in arterial blood - C _v O ₂ content of O₂ in venous blood - %E branchial O₂ extraction efficiency - f _v ventilatory frequency - GTP guanosine triphosphate - Hct haematocrit - [Hb] haemoglobin concentration - ITP inosine triphosphate - met[Hb] methaemoglobin - oxygen consumption - NTP nucleoside triphosphate - OEC oxygen equilibrium curve - P _a O ₂ partial pressure of O₂ in arterial blood - P _e O ₂ partial pressure of expired O₂ - P _i O ₂ partial pressure of inspired O₂ - P _in O ₂ inflow partial pressure of O₂ - PO ₂ partial pressure of O₂ - P _out O ₂ outflow partial pressure of O₂ - pH _a arterial blood pH - pH _pl whole blood pH - PV plasma volume - P _v CO ₂ partial pressure of CO₂in venous blood - P _v O ₂ partial pressure of O₂in venous blood - cardiac output - SW seawater - TMAO trimethylamine-N-oxide - ventilation volume Communicated by G. Heldmaier     

Osmotic,sodium, carbon dioxide and acid-base state of the Port Jackson shark,<Emphasis Type="Italic"> Heterodontus portusjacksoni</Emphasis>, in response to lowered salinity

In marine elasmobranch fish the consequences for CO₂ and acid–base state of moving into low salinity water are not well described. Sub-adult Port Jackson sharks, Heterodontus portusjacksoni, occasionally enter brackish water and survive in 50% seawater (SW). The unidirectional Na efflux and content, plasma volume, glomerular filtration rate (GFR), body mass, as well as CO₂ and acid-base state in H. portusjacksoni were investigated following transfer from 100% SW to 75% SW and then to 50% SW. A rapid water influx resulted in a doubling of the plasma volume within 24 h in sharks in 75% SW and an 11% increase in body weight. Osmotic water influx was only partially offset by a doubling of the GFR. There was a ~40% decrease in plasma [Na] through a transiently elevated Na clearance and haemodilution. The result was a decrease in the inward gradient for Na⁺ together with reductions of nearly 50% in CO₂ and buffer capacity. The sharks remained hypo-natric to 50% SW by partially conforming to the decrease in external osmotic pressure and avoided the need for active Na⁺ uptake. The gradient for Na⁺ efflux would by extrapolation approach zero at ~27% SW which may of itself prove a lethal internal dilution. In sharks transferred to 75% SW, a small transient hypercapnia and a later temporary metabolic alkalosis were all largely explained through anaemia promoting loss of CO₂ and buffer capacity. In sharks transferred to 50% SW the metabolic alkalosis persisted until the end of the 1-week trial. Within the erythrocytes, increased pH was consequent on the large decrease in haemoglobin content exhibited by the sharks, which caused a large reduction in intracellular buffer. In water as dilute as 50% SW there was no evidence of specific effects on the mechanisms of management of CO₂ or H⁺ excretion but rather significant and indirect effects of the severe haemodilution.Abbreviations a–v arterial–venous - CA carbonic anhydrase - C _a CO ₂ content of CO₂ in arterial blood - CCO ₂ CO₂ content - ⁵¹ Cr-EDTA ⁵¹chromium-ethylenediaminetetraactic acid - C _v CO ₂ content of CO₂ in venous blood - FW freshwater - GFR glomerular filtration rate - Hct haematocrit - J _out Na flux rate - MCHC mean cell haemoglobin concentration - OP osmotic pressure - P _a CO ₂ partial pressure of CO₂in arterial blood - PCO ₂ partial pressure of CO₂ - pH _a arterial blood pH - pH _er intra-erythrocyte fluid - pH _pl whole blood pH - pH _v venous blood pH - P _v CO ₂ partial pressure of CO₂in venous blood - SID strong ion difference - SW seawater - TMAO trimethylamine-N-oxide - UFR urinary flow rate Communicated by G. Heldmaier     

Effects of rapid transfer from sea water to fresh water on respiratory variables,blood acid-base status and O2 affinity of haemoglobin in Atlantic salmon (Salmo salar L.)

Summary Oxygen consumption, gill ventilation, blood acid-base/ionic status and haemoglobin oxygen affinity were studied in seawater-adapted adult salmon (Salmo salar) during five weeks after transfer into fresh water. Freshwater exposure induced the following changes: Standard oxygen consumption ( ) and ventilatory flow ( ) decreased markedly during the first days after transfer, then decreased more gradually until a new steady-state was achieved at which and were about 80% and 56% of the control values, respectively. The marked increase in oxygen extraction coefficient (Ew _{O 2}) and the marked decrease in the oxygen convection requirement ( ) were associated with a reduction in the partial pressure of carbon dioxide in arterial blood (Pa _{CO 2}), in spite of a decrease of both ventilatory flow and water CO₂ capacitance. These results suggested that transfer into fresh water induced an increase in branchial diffusive conductance. A biphasic pattern was observed in the time-course of the changes in both plasma ion concentration and acid-base status. During the first 10 days, plasma Na⁺, K⁺, and Cl^– concentrations fell abruptly, then more gradually. [Cl^–] decreased more than [Na⁺] resulting in a progressive increase in the [Na⁺]/[Cl^–] ratio. During the second phase of acclimation to fresh water plasma Na⁺, K⁺, and Cl^– concentrations progressively increased. [Cl^–] increased more than [Na⁺], so that [Na⁺]/[Cl^–] ratio decreased. Transfer into fresh water did not significantly change plasma lactate concentration. Upon exposure to fresh water, blood pH increased from 7.94±0.04 to 8.43±0.06 at day 10 and then decreased to 8.08±0.03 at day 34. The increase in blood pH induced by transfer to fresh water initially represented a mixed metabolic/respiratory alkalosis. However, after 15 days Pa _{CO 2} had returned to pretransfer values and the alkalosis was purely metabolic. The metabolic component of the alkalosis was associated with appropriate changes in the plasma strong ion difference (S.I.D.). Blood alkalosis moved the oxygen dissociation curve to the left, so that P₅₀ was decreased by 30% below the value in seawater for the maximal increase in blood pH. This rise in haemoglobin affinity for O₂, associated with a marked increase in blood buffer capacity, are regarded as adaptative processes allowing the salmon to cope with the markedly increased energy expenditure required for upstream migration.     

Physiological changes in the Norway lobster Nephrops norvegicus (L.) escaping and discarded from commercial trawls on the West Coast of Scotland: II. Disturbances in haemolymph respiratory gases, tissue metabolites and swimming performance after capture and during recovery

The physiological states of trawled and creel-caught (control) Norway Lobsters (Nephrops norvegicus (L.)) captured on grounds off the West Coast of Scotland (120–150 m) were compared. Undersized “discards” (<25–35 mm carapace length) were sampled directly from the cod-end and following recovery in underwater (u/w) cages at a mean depth of 24 m. “Escaped” animals which had passed through the cod-end meshes and collected in a small-meshed net “cover” were transferred without emersion (air-exposure) for sampling on-board. Some of these individuals were also transferred by SCUBA divers to u/w cages. Haemolymph PO₂, PCO₂ and pH measurements showed that both discarded and escaped animals experienced moderate internal hypoxia, hypercapnia and acidosis which became severe after 1 h emersion of the former on deck. A marked handling effect was evident in which haemolymph PO₂ declined and PCO₂ became elevated in both controls and recovering trawled animals. In both discarded and escaped animals haemolymph l-lactate and d-glucose concentrations were high compared to controls, but with levels significantly lower in escapes suggesting less tail-flip swimming activity within the cod-end cover. Further emersion had little effect on haemolymph l-lactate in discards. Recovery to control levels of both metabolites occurred within 24 h in u/w cages but the exercising of captured individuals (by tactile stimulation) produced further significant increases. Abdominal flexor muscle concentrations of l-lactate were also elevated in discards and escapes and positively correlated with haemolymph levels. Muscle glycogen showed severe depletion in both groups compared to unexercised controls and 1 h emersion reduced levels drastically (to 20% of normal concentrations). High haemolymph ammonia (T_amm) was characteristic of both trawled groups but was reduced rapidly during recovery. These metabolite changes were accompanied by reductions in the number of escape swimming tail-flips that could be elicited before exhaustion, particularly in discards. This reduction in performance was evident in discards even after 24 h recovery, but escapes showed almost normal responses. The severity of the physiological stresses experienced during trawling, and to a lesser extent in escaped animals, and their effects on recovery of undersized discards regaining the seabed, is discussed. These findings may help us predict the survival, longer-term recovery and fitness of fished N. norvegicus, and their potential contribution to the regeneration of exploited populations.