Respiratory Adaptations in Intertidal Fish

SYNOPSIS: Intertidal rockpools provide a challenging environmentfor rockpool fish with rapid changes taking place in many environmentalparameters over a tidal cycle. Intertidal fish exhibit a numberof behavioural adaptations such as the avoidance of hypoxicsituations or remaining inactive during aerial "stranding."Other species are, however, well adapted to breathe air andexhibit morphological adaptations such as smaller gill areas,specialized buccopharyngeal epithelia and a proliferation ofcutaneous blood vessels in the skin. Oxygen consumption in rockpoolfish is comparable to non-intertidal fish and responds in asimilar manner to temperature changes. The ability to regulateoxygen consumption down to oxygen tensions below 40 Torr is,however, marked in rockpool species. Aerial and aquatic ratesof respiration are similar in those species which are able tobreathe air and the respiratory quotient normally remains between0.7 and 0.9. A number of intertidal fish are well adapted forcutaneous respiration, satisfying over half of their oxygenand carbon dioxide exchange through the skin. Ventilatory responsesto increased temperature, hyperoxia and hypoxia are similarto those of other fish but cardiac responses may differ in thatno change in heart rate is seen under hypoxia or hyperoxia.Ventilatory and cardiac responses to aerial respiration arewell adapted in some species maintaining ventilation and perfusionduring aerial exposure. A marked Bohr effect, low temperaturesensitivity and a temperature dependent Haldane effect havebeen measured in the haemoglobolin of some intertidal fish.These properties may assist oxygen transport and carbon dioxideexchange during cyclical changes in environmental parameterswithin an intertidal rockpool.     

Effect of Fluctuations in Temperature on the Metabolism of Intertidal Invertebrates

SVNOPSIS. Compensatory changes in the level of activity of intertidalorganisms may occur in response to the thermal conditions whichprevail in the habitat. These involve an increase in the activityof those animals subjected to low temperatures and a correspondingsuppression of activity in those animals subjected to high temperaturesso that organisms with a wide geographical range, or livingover a range of shore levels, have comparable rates of activity.However, the rate of activity varies markedly with short-termfluctuations in temperature, as does the rate of respirationof active animals. Recent evidence suggests that the rate ofrespiration of quiescent animals is relatively independent oftemperature over the normal environmental range, and thus themetabolism is well-suited to an environment where rapid fluctuationsin temperature occur. Further, the extent of the thermal rangeover which metabolism is relatively independent of temperatureis modifiable according to season and storage-temperature. Similarchanges occur in the respiration of cell-free homogenates ofcertain intertidal organisms. Acclimation in such organismsinvolves not only a modification in the level of the activeand standard rates of metabolism but also an alteration in theform of the rate/temperature curve such that the range of temperature-independentmetabolism is appropriate to the thermal conditions prevailingin the habitat.     

Integrated Responses to Salinity Fluctuation

SYNOPSIS. Continuous and discontinuous intertidal waters bothexperience fluctuations in salinity frequently accompanied bychanges in temperature, levels of dissolved gases and pH. Amultidimensional approach to environmental fitness may thereforebe more representative of natural conditions in the intertidalzone. Various combinations of environmental factors are discussedincluding salinity/temperature, and salinity/O₂ tension. Salinitywill directly affect osmotic and ionic regulation and indirectlyaffect acid-base balance and various components of the respiratorysystem including ventilation, gas exchange, perfusion, O₂ transportby the respiratory pigment and utilization at the tissues. Additionalareas covered in this review article are osmotic problems facedby air-breathing aquatic intertidal animals, ontogenetic changesin osmoregulatory capabilities and a comparison of steady state(square wave) versus sinusoidal (cyclical) salinity changes.     

Respiration and Vitality of Binucleate and Trinucleate Pollen

The respiration and vitality of ungerminated bi- and trinucleate pollen were studied in order to determine the influence of relative humidity and temperature on metabolic activity. The gas exchange, germination capacity and staining with tetrazolium bromide were followed under standardized conditions. A constant respiration rate occurred under conditions of high relative humidity (97%). Per mg pollen, the trinucleate grains of Compositae and Gramineae respired 2 to 3 times as intense as 6 species of binucleate grains. Per unit of pollen protein the differences were even larger. In contrast to binucleate pollen, the longevity of trinucleate pollen was very short and the ability to germinate was lost twice as fast as the respiration capacity. This limits the use of tetrazolium bromide as an indicator of viability. At reduced relative humidities respiration was strongly restricted, but the longevity of bi- and trinucleate pollen considerably increased. Pollen of Gramineae, however, was very sensitive to changes in relative humidity; short exposure to low relative humidity decreased both the vitality and the capacity to respire.     

Environmental correlates of phenotypic variation: do variable tidal regimes influence morphology in intertidal seaweeds?

Seaweed morphology is often shaped by the hydrodynamic environment. However, exposure to air at low tide represents an additional factor potentially affecting the morphology of intertidal species. Here, we examined the relationships between the morphology of Hormosira banksii, an important intertidal habitat‐forming seaweed in southern Australia, and environmental factors across multiple spatial scales around the island of Tasmania, Australia. Tasmania is surrounded by a diverse coastline with differences in wave exposure, tidal parameters, and temperature. We sampled Hormosira from four regions (100s km apart), three sites (10s km apart) within each region, and two zones (meters apart; eulittoral and sublittoral) at each site, and measured multiple morphological variables to test for differences in morphology at those different spatial scales. Thirteen environmental variables reflecting wave exposure, tidal conditions, and temperature for each site were generated to assess the relationship between Hormosira morphology and environmental variation. Morphology varied at all spatial scales examined. Most notably, north coast individuals had a distinct morphology, generally having smaller vesicles and shorter fronds, compared to other regions. Tidal conditions were the main environmental factors separating north coast sites from other sites and tidal regime was identified as the best predictor of morphological differences between regions. In contrast to other studies, we found little evidence that wave exposure was associated with morphological variation. Overall, our study emphasizes the role of tidal conditions, associated with emersion stress during low tide, in affecting the morphology of intertidal seaweeds.     

Soil and sediment bacteria capable of aerobic nitrate respiration.

Several laboratory strains of gram-negative bacteria are known to be able to respire nitrate in the presence of oxygen, although the physiological advantage gained from this process is not entirely clear. The contribution that aerobic nitrate respiration makes to the environmental nitrogen cycle has not been studied. As a first step in addressing this question, a strategy which allows for the isolation of organisms capable of reducing nitrate to nitrite following aerobic growth has been developed. Twenty-nine such strains have been isolated from three soils and a freshwater sediment and shown to comprise members of three genera (Pseudomonas, Aeromonas, and Moraxella). All of these strains expressed a nitrate reductase with an active site located in the periplasmic compartment. Twenty-two of the strains showed significant rates of nitrate respiration in the presence of oxygen when assayed with physiological electron donors. Also isolated was one member of the gram-positive genus Arthrobacter, which was likewise able to respire nitrate in the presence of oxygen but appeared to express a different type of nitrate reductase. In the four environments studied, culturable bacteria capable of aerobic nitrate respiration were isolated in significant numbers (10(4) to 10(7) per g of soil or sediment) and in three cases were as abundant as, or more abundant than, culturable bacteria capable of denitrification. Thus, it seems likely that the corespiration of nitrate and oxygen may indeed make a significant contribution to the flux of nitrate to nitrite in the environment.     

The influence of body size on the diving behaviour and physiology of the bimodally respiring turtle, Elseya albagula

In aquatic vertebrates that acquire oxygen aerially dive duration scales positively with body mass, i.e. larger animals can dive for longer periods, however in bimodally respiring animals the relationship between dive duration and body mass is unclear. In this study we investigated the relationships between body size, aquatic respiration, and dive duration in the bimodally respiring turtle, Elseya albagula. Under normoxic conditions, dive duration was found to be independent of body mass. The dive durations of smaller turtles were equivalent to that of larger individuals despite their relatively smaller oxygen stores and higher mass specific metabolic rates. Smaller turtles were able to increase their dive duration through the use of aquatic respiration. Smaller turtles had a relatively higher cloacal bursae surface area than larger turtles, which allowed them to extract a relatively larger amount of oxygen from the water. By removing the ability to respire aquatically (hypoxic conditions), the dive duration of the smaller turtles significantly decreased restoring the normal positive relationship between body size and dive duration that is seen in other air-breathing vertebrates.     

在水生与气生状态下石莼光合作用对光照和温度的响应

如何对付由于高潮时的水生状态与低潮时的气生状态高频率循环所导致的不同环境条件,是潮间带海藻的光合作用所面临的独特问题。对采自汕头沿岸的石莼(Ulva lactuca)在水生和气生不同状态下光合作用对光照和温度的响应特性进行了测定,以探讨这种常见的潮间带绿藻在潮汐循环背景下的光合特性。在气生状态下,光饱和净光合速率(Pmax)随气生暴露时间的变化模式可以很好地用三次方程进行描述,而温度影响方程的系数;当水分损失为15％时,石莼的Pmax增加至最大值,然后Pmax随进一步脱水而下降,在水分损失为80％时下降至0。温度对Pmax的影响在水生状态下比在气生状态下更大。气生状态下（充分水化）Pmax在10℃时显著小于水生状态下的值,而在30℃时则相反。在10℃时,气生干出时间在6 h 以内,或在20℃时,气生干出时间在2.2 h 以内,石莼的净碳固定量在气生状态下比在水生状态下要大;而在30℃时,在气生状态下的净碳固定量比总是小于在水生状态下的净碳固定量。认为石莼在低潮气生状态下与在高潮水生状态下光合特性及净碳固定存在差异,但这种差异与环境温度及叶状体的水分状态有关。     

在水生与气生状态下石莼光合作用对光照和温度的响应

如何对付由于高潮时的水生状态与低潮时的气生状态高频率循环所导致的不同环境条件,是潮间带海藻的光合作用所面临的独特问题。对采自汕头沿岸的石莼(Ulva lactuca)在水生和气生不同状态下光合作用对光照和温度的响应特性进行了测定,以探讨这种常见的潮间带绿藻在潮汐循环背景下的光合特性。在气生状态下,光饱和净光合速率(Pmax)随气生暴露时间的变化模式可以很好地用三次方程进行描述,而温度影响方程的系数;当水分损失为15％时,石莼的Pmax增加至最大值,然后Pmax随进一步脱水而下降,在水分损失为80％时下降至0。温度对Pmax的影响在水生状态下比在气生状态下更大。气生状态下(充分水化)Pmax在10℃时显著小于水生状态下的值,而在30％时则相反。在10℃时,气生干出时间在6h以内,或在20％时,气生干出时间在2．2h以内,石莼的净碳固定量在气生状态下比在水生状态下要大;而在30％时,在气生状态下的净碳固定量比总是小于在水生状态下的净碳固定量。认为石莼在低潮气生状态下与在高潮水生状态下光合特性及净碳固定存在差异,但这种差异与环境温度及叶状体的水分状态有关。     

Seasonal variations in the rates of aquatic and aerial respiration and ammonium excretion of the ribbed mussel, Geukensia demissa (Dillwyn)

Aquatic and aerial rates of oxygen consumption and ammonium excretion of ribbed mussels, Geukensia demissa (Dillwyn), collected from the mid-intertidal zone of a mid-Atlantic salt marsh, were measured under ambient conditions of food, temperature, and salinity over five seasons. Rates of aquatic respiration covaried with body size and season, as the rates were high and strongly related to mussel tissue weight in spring and summer but low and weight independent in winter. There was a significant interannual difference between summer of 1995 and 1996. Rates of aerial respiration also varied seasonally, with high rates of oxygen consumption in summer and low rates in winter. The magnitude of these seasonal variations were greater than those for aquatic respiration, and as a result, the ratio of aerial to aquatic respiration was higher in summer (0.61 and 0.52) than in winter (0.11). This indicates that G. demissa was able to actively regulate aerial respiration, thereby permitting high aerobic metabolism during prolonged periods of air exposure in summer. We hypothesize that such high rates of aerial respiration, during the seasons of high metabolic activity, are required to provide sufficient energy for mussels to facilitate food digestion during air exposure at low tide. Rates of ammonium excretion varied seasonally and increased with mussel weight in all seasons. The atomic ratio of oxygen to nitrogen (O:N), calculated from aquatic respiration vs. ammonium excretion, was significantly lower in autumn (26) than in other seasons (46–60) among which the O:N did not vary significantly.     

Exposure to elevated temperature and Pco(2) reduces respiration rate and energy status in the periwinkle Littorina littorea

In the future, marine organisms will face the challenge of coping with multiple environmental changes associated with increased levels of atmospheric Pco(2), such as ocean warming and acidification. To predict how organisms may or may not meet these challenges, an in-depth understanding of the physiological and biochemical mechanisms underpinning organismal responses to climate change is needed. Here, we investigate the effects of elevated Pco(2) and temperature on the whole-organism and cellular physiology of the periwinkle Littorina littorea. Metabolic rates (measured as respiration rates), adenylate energy nucleotide concentrations and indexes, and end-product metabolite concentrations were measured. Compared with values for control conditions, snails decreased their respiration rate by 31% in response to elevated Pco(2) and by 15% in response to a combination of increased Pco(2) and temperature. Decreased respiration rates were associated with metabolic reduction and an increase in end-product metabolites in acidified treatments, indicating an increased reliance on anaerobic metabolism. There was also an interactive effect of elevated Pco(2) and temperature on total adenylate nucleotides, which was apparently compensated for by the maintenance of adenylate energy charge via AMP deaminase activity. Our findings suggest that marine intertidal organisms are likely to exhibit complex physiological responses to future environmental drivers, with likely negative effects on growth, population dynamics, and, ultimately, ecosystem processes.     

Thermal plasticity of diving behavior, aquatic respiration, and locomotor performance in the Mary River turtle Elusor macrurus

Locomotion is a common measure of performance used in studies of thermal acclimation because of its correlation with predator escape and prey capture. However, for sedentary animals such as freshwater turtles, we propose that diving behavior may be a more ecologically relevant measure of performance. Increasing dive duration in hatchling turtles reduces predator exposure and therefore functions as an ecological benefit. Diving behavior is thermally dependent, and in some species of freshwater turtles, it is also reliant on aquatic respiration. This study examined the influence of thermal acclimation on diving behavior, aquatic respiration, and locomotor performance in the endangered, bimodally respiring Mary River turtle Elusor macrurus. Diving behavior was found to partially acclimate at 17 degrees C, with turtles acclimated to a cold temperature (17 degrees C) having a significantly longer dive duration than hatchlings acclimated to a warm temperature (28 degrees C). This increase in dive duration at 17 degrees C was not a result of physiological alterations in metabolic rate but was due instead to an increase in aquatic oxygen consumption. Increasing aquatic oxygen consumption permitted cold-acclimated hatchlings to remain submerged for significantly longer periods, with one turtle undertaking a dive of over 2.5 d. When burst-swimming speed was used as the measure of performance, thermal acclimation was not detected. Overall, E. macrurus demonstrated a partial ability to acclimate to changes in environmental temperature.     

Temperature‐dependent oxygen limitation and the rise of Bergmann's rule in species with aquatic respiration

Bergmann's rule is the propensity for species‐mean body size to decrease with increasing temperature. Temperature‐dependent oxygen limitation has been hypothesized to help drive temperature–size relationships among ectotherms, including Bergmann's rule, where organisms reduce body size under warm oxygen‐limited conditions, thereby maintaining aerobic scope. Temperature‐dependent oxygen limitation should be most pronounced among aquatic ectotherms that cannot breathe aerially, as oxygen solubility in water decreases with increasing temperature. We use phylogenetically explicit analyses to show that species‐mean adult size of aquatic salamanders with branchial or cutaneous oxygen uptake becomes small in warm environments and large in cool environments, whereas body size of aquatic species with lungs (i.e., that respire aerially), as well as size of semiaquatic and terrestrial species do not decrease with temperature. We argue that oxygen limitation drives the evolution of small size in warm aquatic environments for species with aquatic respiration. More broadly, the stronger decline in size with temperature observed in aquatic versus terrestrial salamander species mirrors the relatively strong plastic declines in size observed previously among aquatic versus terrestrial invertebrates, suggesting that temperature‐dependent oxygen availability can help drive patterns of plasticity, micro‐ and macroevolution.     

Climate extremes are associated with invertebrate taxonomic and functional composition in mountain lakes

Climate change is expected to increase climate variability and the occurrence of extreme climatic events, with potentially devastating effects on aquatic ecosystems. However, little is known about the role of climate extremes in structuring aquatic communities or the interplay between climate and local abiotic and biotic factors. Here, we examine the relative influence of climate and local abiotic and biotic conditions on biodiversity and community structure in lake invertebrates. We sampled aquatic invertebrates and measured environmental variables in 19 lakes throughout California, USA, to test hypotheses of the relationship between climate, local biotic and environmental conditions, and the taxonomic and functional structure of aquatic invertebrate communities. We found that, while local biotic and abiotic factors such as habitat availability and conductivity were the most consistent predictors of alpha diversity, extreme climate conditions such as maximum summer temperature and dry‐season precipitation were most often associated with multivariate taxonomic and functional composition. Specifically, sites with high maximum temperatures and low dry‐season precipitation housed communities containing high abundances of large predatory taxa. Furthermore, both climate dissimilarity and abiotic dissimilarity determined taxonomic turnover among sites (beta diversity). These findings suggest that while local‐scale environmental variables may predict alpha diversity, climatic variability is important to consider when projecting broad‐scale aquatic community responses to the extreme temperature and precipitation events that are expected for much of the world during the next century.     

Thermal plasticity is independent of environmental history in an intertidal seaweed

Organisms inhabiting the intertidal zone have been used to study natural ecophysiological responses and adaptations to thermal stress because these organisms are routinely exposed to high‐temperature conditions for hours at a time. While intertidal organisms may be inherently better at withstanding temperature stress due to regular exposure and acclimation, they could be more vulnerable to temperature stress, already living near the edge of their thermal limits. Strong gradients in thermal stress across the intertidal zone present an opportunity to test whether thermal tolerance is a plastic or canalized trait in intertidal organisms. Here, we studied the intertidal pool‐dwelling calcified alga, Ellisolandia elongata, under near‐future temperature regimes, and the dependence of its thermal acclimatization response on environmental history. Two timescales of environmental history were tested during this experiment. The intertidal pool of origin was representative of long‐term environmental history over the alga's life (including settlement and development), while the pool it was transplanted into accounted for recent environmental history (acclimation over many months). Unexpectedly, neither long‐term nor short‐term environmental history, nor ambient conditions, affected photosynthetic rates in E. elongata. Individuals were plastic in their photosynthetic response to laboratory temperature treatments (mean 13.2°C, 15.7°C, and 17.7°C). Further, replicate ramets from the same individual were not always consistent in their photosynthetic performance from one experimental time point to another or between treatments and exhibited no clear trend in variability over experimental time. High variability in climate change responses between individuals may indicate the potential for resilience to future conditions and, thus, may play a compensatory role at the population or species level over time.     

Temporal variations in microbenthic metabolism and inorganic nitrogen fluxes in sandy and muddy sediments of a tidally dominated bay in the northern Wadden Sea

Factors controlling seasonal variations in benthic metabolism (O₂ flux) and dissolved inorganic nitrogen (DIN) fluxes were examined during a 12–14 month period at three intertidal Wadden Sea stations. Since the flux measurements were made as small-scale laboratory core incubations, the results are primarily related to the microbenthic community (microalgae, bacteria, micro-, meio- and small macrofauna) and cannot be considered representative of the total benthic community in the Wadden Sea. Furthermore, it has to be emphasized that light intensity during day-time simulations were constant and saturating at all times. Benthic primary production and oxygen uptake appeared to be temperature dependent with a ‘seasonal Q₁₀’ of 1.7–1.8 and 2.7–4.3, respectively. Inundation had no effect on oxygen fluxes as evidenced by similar sediment respiration with and without water cover. A stronger temperature dependence of primary production in muddy than in sandy sediment indicated that the overall control in the latter may be complex due to factors like macrofaunal grazing and nutrient availability. Benthic respiration may not be controlled by temperature alone, as sedimentary organic matter content correlated significantly with both temperature and benthic respiration. Annual gross primary production in high intertidal sandy sediment was 10 and 50% higher than in low intertidal sandy and muddy sediments, respectively. Since annual benthic community respiration was 2 times higher in muddy than sandy sediments, the annual net primary production was about 0 in the former and 17–19 mol C m^?2 yr^?1 in the latter. However, heterotrophic contribution by larger faunal components as well as removal of organic carbon by waves and tidal currents, which are not included here, may balance the budget at the sandy stations. There was no or only weak relationships between (light and dark) DIN exchange and factors like temperature, sedimentary organic content, and oxygen fluxes. Factors related to nutrient fluxes, such as denitrification and nutrient concentration in the overlying water, may have hampered any such relationships. In fact, DIN fluxes at all three stations appeared to be strongly controlled by DIN concentrations in the overlying water. On an annual basis, the sediment appeared to be a net sink for DIN.     

Tolerance of forced air emergence by a fish with a broad vertical distribution, the rockpool blenny, Hypsoblennius gilberti (Blenniidae)

Many intertidal fishes, particularly among the Blenniidae and Cottidae, possess amphibious adaptations, including the ability to breathe in air and to avoid desiccation in terrestrial conditions. These traits are absent in subtidal species of blennies and cottids. Hypsoblennius gilberti, the rockpool blenny, is found in shallow rockpools in the mid to high intertidal areas of Southern California, and deeper to 18 m in the subtidal zone. This broad vertical distribution could indicate that this blenny is adapted for tidal air emergence, although H. gilberti has not been observed out of water in its natural habitat. H. gilberti does not emerge voluntarily from hypoxic sea water in the laboratory, but it easily withstands 3 h out of water. The aerial respiratory exchange ratio (CO2 released compared to O2 consumed) is 0.70, similar to that of amphibious intertidal fishes in air, indicating sufficient release of metabolically produced CO2 while emerged. There is no increase in aquatic respiration following emergence. However, unlike other amphibious fishes that maintain aerial oxygen consumption at a level similar to aquatic oxygen consumption, H. gilberti has an aerial oxygen consumption rate one-third that in water. H. gilberti can recover rapidly from terrestrial water loss, and shows no change in evaporative water loss rates at 93% and 77% relative humidities. The amphibious capabilities in H. gilberti, even if rarely used, permit survival in air during tidal emergence. These findings suggest that H. gilberti may demonstrate an intermediate condition between the amphibious species of intertidal fishes that regularly emerge from water, and the subtidal fishes that do not survive air emergence and are completely restricted to an aquatic habitat.     

Ecological, Evolutionary, and Physical Factors Influencing Aquatic Animal Respiration

Compared to air-breathers, animals that respire aquaticallyhave limited access to O₂ and their habitats are more subjectto hypoxia. Because O₂ diffuses more slowly through water thanair, animals in water experience greater diffusion boundarylayer effects on respiratory gas diffusion. While ventilationand specialized exchange surfaces mitigate O₂ diffusion limitationson respiration, most animal phyla, particularly those confinedto aquatic habitats, lack these. Diffusion limitation influencesthe ontogeny of aquatic animals and may have also shaped Precambrianmetazoans. In spite of a more limited O₂ access, aquatic animalsdisplay a much greater spectrum of respiratory adaptation, rangingfrom the loss of Hb in icefishes to the independent evolution,invention, and acquisition of Hb in many invertebrates confinedto hypoxic habitats. Three features of aquatic respiratory systemsdistinguishing them from aerial systems are the widespread occurrenceof integumental respiration, the frequent presence of combinedrespiratory and feeding surfaces, and the profound effect ofhypoxia on shaping respiratory adaptation, both in shallow waterand in the deep sea.     

The combined effects of temperature,salinity, and declining oxygen tension on oxygen consumption in the marine pulmonate Amphibola crenata (Gmelin, 1791)

Oxygen consumption of Amphibola crenata (Gmelin) was measured in various salinity-temperature combinations (< 0.1‰ to 41‰ salinity and 5 to 30°C) in air, and following exposure to declining oxygen tensions. In all experimental conditions, respiration varied with the 0.44 power of the body weight (sd = 0.14). The aquatic rate was consistently higher than the aerial rate of oxygen consumption, although at 30 °C the two rates were similar. Oxygen consumption increased with temperature up to 25 °C in all salinities; the lowest values were recorded at temperatures below 10 °C and at 30 °C in the most dilute medium. At all exposure temperatures, the oxygen consumption of Amphibola decreased regularly with salinity down to 0.1 ‰, and following exposure to concentrated sea water (41‰). Salinity had the least effect at 15 °C which was the acclimation temperature. In general, all of the temperature coefficients (Q₁₀ values) were low, < 1.65. However, Q₁₀ values above 2.8 were recorded at a salinity of 17.8‰ between 10 and 15 °C. Oxygen consumption of all size classes of Amphibola was more temperature dependent in air than in water and small individuals show a greater difference between their aerial and aquatic rates than larger snails. The rates of oxygen consumption in declining oxygen tensions were expressed as fractions of the rates in air saturated sea water at each experimental salinity-temperature combination. The quadratic coefficient B₂ becomes increasingly more negative with both decreasing salinity and temperatures up to 20 °C. At higher temperatures (25 and 30 °C) the response is reversed such that O₂ uptake in snails becomes increasingly independent of declining oxygen tensions at higher salinities. On exposure to a salinity of 4‰, Amphibola showed no systematic response to declining oxygen tension with respect to temperature. The ability of Amphibola to maintain its rate of oxygen consumption in a wide range of environmental conditions is discussed in relation to its potential for invading terrestrial habitats and its widespread distribution on New Zealand's intertidal mudflats.     

A Comparative Study of Respiration Rates of Some Aquatic Oligochaetes in Relation to Sublethal Stress

Aquatic oligochaete species selected on the basis of pollution tolerance or intolerance as determined from field studies were used to determine respiration rates under standard conditions, after exposure to sublethal levels of environmental variables (pH, salinity, temperature) or toxicants (Cd, Hg, NaPCP) alone or in pairs. Respiration rates, critical oxygen levels and degree of regulation were not correlated with field-determined tolerance to organic pollution. Stress factors caused increased or lowered respiration rate, loss of regulation (partial or total) and sometimes shifts in critical oxygen level. Regulation was improved in the face of some stresses. Combined stresses produced additional changes in respiration rates and regulation. These responses were complex, which makes them useful indicators of specific stress factors but also indicate complex mechanisms behind the observed changes.