Physiological responses to air exposure: acid-base balance and the role of branchial water stores

SYNOPSIS. Intertidal organisms usually exhibit one of two behaviors when they are air exposed. They either isolate themselves from the aerial environment or they interact with the aerial environment. Among the animals using the first behavior, body fluid acid-base balance is partially maintained by dissolution of the calcium carbonate shell in the bivalve molluscs to buffer the metabolic acids produced anaerobically. Calcium ions compensate for the acidosis by increasing the strong ion difference. The release of carbonate from the shell causes an increase in molecular CO2 which offsets the compensation somewhat, but this effect is minimized by distributing the CO2 among the other fluids within the shell and/or by venting the shell to the air. In animals which have a fairly high concentration of a respiratory pigment, such as a lugworm, an anaerobically induced acidosis is minimized by a large Haldane effect. Among the animals which interact with the aerial environment, the decapod crustaceans by and large maintain their metabolism aerobically, although it may be greatly reduced. A respiratory acidosis due to elevated hemolymph Pco2 may be either fully compensated or not at all. Compensation involves an increase in hemolymph calcium, probably from the calcium carbonate exoskeleton. Compensatory mechanisms may also include branchial water stores which accumulate a titratable base. It is suggested that the alkalinization of the branchial water maintains a steeper Pco2 gradient across the gill and reduces the magnitude of the acidosis for a short period of time. The ability to use branchial water stores in this way may be tied to the ability of the animal to osmoregulate.

[1]  L. F. Ayvaziann.  The Regulation of Acid-Base Balance , 1989 .

[2]  P. Defur.  Systemic Respiratory Adaptations to Air Exposure in Intertidal Decapod Crustaceans , 1988 .

[3]  J. Widdows,et al.  Calorimetric studies of behavior, metabolism and energetics of sessile intertidal animals , 1988 .

[4]  M. Grieshaber,et al.  The Regulation of Haemocyanin Oxygen Affinity During Emersion of the Crayfish Austropotamobius Pallipes: III. The Dependence of Ca2+-Haemocyanin Binding on the Concentration Of L-Lactate , 1987 .

[5]  L. Burnett,et al.  Gas Exchange, Hemolymph Acid-Base Status, and the Role of Branchial Water Stores during Air Exposure in Three Littoral Crab Species , 1987, Physiological Zoology.

[6]  J. Widdows,et al.  Activity and Metabolism in the Mussel Mytilus edulis L. during Intertidal Hypoxia and Aerobic Recovery , 1986, Physiological Zoology.

[7]  C. Mangum,et al.  THE CO2 SENSITIVITY OF THE HEMOCYANINS AND ITS RELATIONSHIP TO Cl- SENSITIVITY , 1986 .

[8]  E. Taylor,et al.  THE REGULATION OF HAEMOCYANIN OXYGEN AFFINITY DURING EMERSION OF THE CRAYFISH AUSTROPOTAMOBIUS PALLIPES II. AN INVESTIGATION OF IN VIVO CHANGES IN OXYGEN AFFINITY , 1986 .

[9]  J. N. Cameron.  Compensation of hypercapnic acidosis in the aquatic blue crab, Callinectes sapidus: the predominance of external sea water over carapace carbonate as the proton sink , 1985 .

[10]  L. Burnett,et al.  The CO2‐specific sensitivity of hemocyanin oxygen affinity in the decapod crustaceans , 1984 .

[11]  B. Mcmahon,et al.  Physiological Compensation to Short-Term Air Exposure in Red Rock Crabs, Cancer productus Randall, from Littoral and Sublittoral Habitats. II. Acid-Base Balance , 1984, Physiological Zoology.

[12]  B. Mcmahon,et al.  Physiological Compensation to Short-Term Air Exposure in Red Rock Crabs, Cancer productus Randall, from Littoral and Sublittoral Habitats. I. Oxygen Uptake and Transport , 1984, Physiological Zoology.

[13]  T. Wolcott.  Uptake of Interstitial Water from Soil: Mechanisms and Ecological Significance in the Ghost Crab Ocypode quadrata and Two Gecarcinid Land Crabs , 1984, Physiological Zoology.

[14]  P. Walsh,et al.  Acid-base balance in the sea mussel, Mytilus edulis. II: Effects of hypoxia and air-exposure on intracellular acid-base status , 1984 .

[15]  R. Full,et al.  RESPIRATION OF CRABS IN AIR AND WATER , 1984 .

[16]  B. Mcmahon,et al.  ANALYSIS OF HEMOLYMPH OXYGEN LEVELS AND ACID-BASE STATUS DURING EMERSION 'IN SITU' IN THE RED ROCK CRAB, CANCER PRODUCTUS. , 1983, The Biological bulletin.

[17]  R. Henry,et al.  The Role of Carbonic Anhydrase in Respiration, Ion Regulation and Acid-Base Balance in the Aquatic Crab Calunectes Sapidus and the Terrestrial Crab Gecarcinus Lateraus , 1983 .

[18]  J. Truchot.  8 – Regulation of Acid—Base Balance , 1983 .

[19]  A. Zwaan.  4 – Carbohydrate Catabolism in Bivalves , 1983 .

[20]  M. Wheatly,et al.  Responses to Hypersaline Exposure in the Euryhaline Crayfish Pacifastacus Leniusculus: I. The Interaction between Ionic and Acid-base Regulation , 1982 .

[21]  H. J. Fyhn,et al.  The influence of aerial exposure upon respiratory and osmotic properties of haemolymph from two intertidal mussels, Mytilus edulis L. and Modiolus modiolus L. , 1982 .

[22]  R. Henry,et al.  THE ROLE OF CaCO3 DISSOLUTION AS A SOURCE OF HCO3- FOR THE BUFFERING OF HYPERCAPNIC ACIDOSIS IN AQUATIC AND TERRESTRIAL DECAPOD CRUSTACEANS , 1981 .

[23]  C. Wood,et al.  Carbon dioxide excretion in the land crab (Cardisoma carnifex) , 1981 .

[24]  C. Wood,et al.  Oxygen and carbon dioxide exchange during exercise in the land crab (Cardisoma carnifex) , 1981 .

[25]  C. Wood,et al.  Haemolymph gas transport, acid‐base regulation, and anaerobic metabolism during exercise in the land crab (Cardisoma carnifex) , 1981 .

[26]  E. Taylor,et al.  The Effect of Long-Term Aerial Exposure on Heart Rate, Ventilation, Respiratory Gas Exchange and Acid-Base Status in the Crayfish Austropotamobius Pallipes , 1981 .

[27]  L. Burnett,et al.  Crab Gill Intra-Epithelial Carbonic Anhydrase Plays a Major Role in Haemolymph CO2 and Chloride Ion Regulation , 1981 .

[28]  B. Mcmahon,et al.  Non-equilibrium acid-base status in C. productus: role of exoskeletal carbonate buffers. , 1980, Respiration physiology.

[29]  J. Truchot.  Mechanisms of the compensation of blood respiratory acid‐base disturbances in the shore crab, Carcinus maenas (L.) , 1979 .

[30]  J. Grant,et al.  Desiccation tolerance ofEurypanopeus depressus (smith) (Decapoda: Xanthidae) and the exploitation of microhabitat , 1979 .

[31]  C. Mangum,et al.  The influence of inorganic ions and pH on oxygenation properties of the blood in the gastropod mollusc Busycon canaliculatum , 1979 .

[32]  C. Mangum,et al.  Oxygen Uptake and Transport in the Lamellibranch Mollusc Modiolus demissus , 1978, Physiological Zoology.

[33]  R. Gleeson,et al.  The determination of hemolymph volume in the blue crab, Callinectes sapidus, utilizing 14C-thiocyanate , 1977 .

[34]  J. Truchot.  Carbon dioxide combining properties of the blood of the shore crab Carcinus maenas (L): carbon dioxide solubility coefficient and carbonic acid dissociation constants. , 1976, The Journal of experimental biology.

[35]  J. Bernards.  Principles of comparative respiratory physiology , 1976 .

[36]  J. Truchot.  Blood acid-base changes during experimental emersion and reimmersion of the intertidal crab Carcinus maenas (L.). , 1975, Respiration physiology.

[37]  C. Mangum,et al.  The quantitative role of hemocyanin in aerobic respiration of limulus polyphemus. , 1975, The Journal of experimental zoology.

[38]  J. V. Bannister.  The respiration in air and in water of the limpets Patella caerulea (L.) and Patella lusitanica (Gmelin). , 1974, Comparative biochemistry and physiology. A, Comparative physiology.

[39]  A. Toulmond.  Tide-related changes of blood respiratory variables in the lugworm Arenicola marina (l.). , 1973, Respiration physiology.

[40]  K. Johansen,et al.  Gas exchange in the giant sea cradle Cryptochiton stelleri (Middendorff) , 1973 .

[41]  M. Crenshaw.  THE INORGANIC COMPOSITION OF MOLLUSCAN EXTRAPALLIAL FLUID. , 1972, The Biological bulletin.

[42]  J. Neff,et al.  Decalcification at the Mantle-Shell Interface in Molluscs , 1969 .

[43]  J. Riegel.  Analysis of the distribution of sodium, potassium and osmotic pressure in the urine of crayfishes. , 1968, The Journal of experimental biology.

[44]  P. A. Dehnel.  ASPECTS OF OSMOREGULATION IN TWO SPECIES OF INTERTIDAL CRABS , 1962 .

[45]  J. Shaw.  Sodium Balance in Eriocheir Sinensis (M. EDW.). The Adaptation of the Crustacea to Fresh Water , 1961 .

[46]  Launce J. Flemister.  SALT AND WATER ANATOMY, CONSTANCY AND REGULATION IN RELATED CRABS FROM MARINE AND TERRESTRIAL HABITATS , 1958 .

[47]  C. Prosser,et al.  IONIC AND OSMOTIC CONCENTRATIONS IN BLOOD AND URINE OF PACHYGRAPSUS CRASSIPES ACCLIMATED TO DIFFERENT SALINITIES , 1955 .

[48]  W. J. Gross.  Aspects of Osmotic Regulation in Crabs Showing the Terrestrial Habit , 1955, The American Naturalist.

[49]  L. Dugal.  The use of calcareous shell to buffer the product of anaerobic glycolysis in Venus mercenaria , 1939 .

[50]  G. S. Carter.  AQUATIC AND AERIAL RESPIRATION IN ANIMALS , 1931 .

[51]  J. B. Collip.  A FURTHER STUDY OF THE RESPIRATORY PROCESSES IN MYA ARENARIA AND OTHER MARINE MOLLUSCA , 1921 .

[52]  J. B. Collip.  STUDIES ON MOLLUSCAN CELOMIC FLUID EFFECT OF CHANGE IN ENVIRONMENT ON THE CARBON DIOXIDE CONTENT OF THE CELOMIC FLUID. ANAEROBIC RESPIRATION IN MYA ARENARIA , 1920 .