Innate Predator Recognition and the Problem of Introduced Trout

Innate predator recognition typically only occurs when there is an evolutionary history between predator and prey. Predator introductions thus can pose a substantial threat to native fauna that rely heavily on inherent identification of predators. In permanent aquatic habitats prey often encounter a variety of predatory and non-predatory fish species, and the ability to distinguish between the two is essential to avoid wasted time and energy spent in unnecessary antipredatory efforts. Here, we present a study evaluating the ability of lab-reared larvae of an endangered fully aquatic salamander (hellbenders: Cryptobranchus alleganiensis ) to recognize chemical cues from native and introduced fish predators. We recorded responses of hellbender larvae to chemical stimuli from native and non-native predatory fishes, a non-predatory fish and a blank control. Eastern hellbender larvae ( C. a. alleganiensis ) significantly reduced activity in response to chemical stimuli from native predators ( Micropterus salmoides , Micropterus dolomieu , Ambloplites rupestris , Sander vitreus , and Cottus carolinae ), but responses to non-native rainbow ( Oncorhynchus mykiss ) and brown ( Salmo trutta ) trout were not significantly different from responses to the non-predatory control (redhorse sucker, Moxostoma spp.). Responses of larval Ozark hellbenders ( C. a. bishopi ) to brown trout were similar to that of the native fishes and different from the blank control, but responses to rainbow trout did not differ from the blank control. The generally weak responses of larval hellbenders to chemical cues from introduced predatory trout could lead to increased predation in the wild, which may have exacerbated the decline of hellbender populations.     

Costly plastic morphological responses to predator specific odour cues in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)

Predation risk is one of the major forces affecting phenotypic variation among and within animal populations. While fixed anti-predator morphologies are favoured when predation level is consistently high, plastic morphological responses are advantageous when predation risk is changing temporarily, spatially, or qualitatively. Three-spined sticklebacks (Gasterosteus aculeatus) are well known for their substantial variability in morphology, including defensive traits. Part of this variation might be due to phenotypic plasticity. However, little is known about sticklebacks’ plastic ability to react morphologically to changing risks of predation and about the proximate cues involved. Using a split-clutch design we show that odour of a predatory fish induces morphological changes in sticklebacks. Under predation risk, i.e., when exposed to odour of a predator, fish grew faster and developed a different morphology, compared to fish reared under low predation risk, i.e., exposed to odour of a non-predatory fish, or in a fish-free environment. However, fast growing comes at cost of increased body asymmetries suggesting developmental constraints. The results indicate that sticklebacks are able to distinguish between predatory and non-predatory fishes by olfactory cues alone. As fishes were fed on invertebrates, this reaction was not induced by chemical cues of digested conspecifics, but rather by predator cues themselves. Further, the results show that variation in body morphology in sticklebacks has not only a strong genetical component, but is also based on plastic responses to different environments, in our case different predation pressures, thus opening new questions for this model species in ecology and evolution.     

Impact of intentionally injected carbon dioxide hydrate on deep-sea benthic foraminiferal survival

Sequestration of carbon dioxide (CO₂) in the ocean is being considered as a feasible mechanism to mitigate the alarming rate in its atmospheric rise. Little is known, however, about how the resulting hypercapnia and ocean acidification may affect marine fauna. In an effort to understand better the protistan reaction to such an environmental perturbation, the survivorship of benthic foraminifera, which is a prevalent group of protists, was studied in response to deep-sea CO₂ release. The survival response of calcareous, agglutinated, and thecate foraminifera was determined in two experiments at ∼3.1 and 3.3 km water depth in Monterey Bay (California, USA). Approximately 5 weeks after initial seafloor CO₂ release, in situ incubations of the live–dead indicator CellTracker Green were executed within seafloor-emplaced pushcores. Experimental treatments included direct exposure to CO₂ hydrate, two levels of lesser exposure adjacent to CO₂ hydrate, and controls, which were far removed from the CO₂ hydrate release. Results indicate that survivorship rates of agglutinated and thecate foraminifera were not significantly impacted by direct exposure but the survivorship of calcareous foraminifera was significantly lower in direct exposure treatments compared with controls. Observations suggest that, if large scale CO₂ sequestration is enacted on the deep-sea floor, survival of two major groups of this prevalent protistan taxon will likely not be severely impacted, while calcareous foraminifera will face considerable challenges to maintain their benthic populations in areas directly exposed to CO₂ hydrate.     

Effects of ocean acidification on learning in coral reef fishes

Ocean acidification has the potential to cause dramatic changes in marine ecosystems. Larval damselfish exposed to concentrations of CO(2) predicted to occur in the mid- to late-century show maladaptive responses to predator cues. However, there is considerable variation both within and between species in CO(2) effects, whereby some individuals are unaffected at particular CO(2) concentrations while others show maladaptive responses to predator odour. Our goal was to test whether learning via chemical or visual information would be impaired by ocean acidification and ultimately, whether learning can mitigate the effects of ocean acidification by restoring the appropriate responses of prey to predators. Using two highly efficient and widespread mechanisms for predator learning, we compared the behaviour of pre-settlement damselfish Pomacentrus amboinensis that were exposed to 440 μatm CO(2) (current day levels) or 850 μatm CO(2), a concentration predicted to occur in the ocean before the end of this century. We found that, regardless of the method of learning, damselfish exposed to elevated CO(2) failed to learn to respond appropriately to a common predator, the dottyback, Pseudochromis fuscus. To determine whether the lack of response was due to a failure in learning or rather a short-term shift in trade-offs preventing the fish from displaying overt antipredator responses, we conditioned 440 or 700 μatm-CO(2) fish to learn to recognize a dottyback as a predator using injured conspecific cues, as in Experiment 1. When tested one day post-conditioning, CO(2) exposed fish failed to respond to predator odour. When tested 5 days post-conditioning, CO(2) exposed fish still failed to show an antipredator response to the dottyback odour, despite the fact that both control and CO(2)-treated fish responded to a general risk cue (injured conspecific cues). These results indicate that exposure to CO(2) may alter the cognitive ability of juvenile fish and render learning ineffective.     

Ocean acidification alters temperature and salinity preferences in larval fish

Ocean acidification alters the way in which animals perceive and respond to their world by affecting a variety of senses such as audition, olfaction, vision and pH sensing. Marine species rely on other senses as well, but we know little of how these might be affected by ocean acidification. We tested whether ocean acidification can alter the preference for physicochemical cues used for dispersal between ocean and estuarine environments. We experimentally assessed the behavioural response of a larval fish (Lates calcarifer) to elevated temperature and reduced salinity, including estuarine water of multiple cues for detecting settlement habitat. Larval fish raised under elevated CO₂ concentrations were attracted by warmer water, but temperature had no effect on fish raised in contemporary CO₂ concentrations. In contrast, contemporary larvae were deterred by lower salinity water, where CO₂-treated fish showed no such response. Natural estuarine water—of higher temperature, lower salinity, and containing estuarine olfactory cues—was only preferred by fish treated under forecasted high CO₂ conditions. We show for the first time that attraction by larval fish towards physicochemical cues can be altered by ocean acidification. Such alterations to perception and evaluation of environmental cues during the critical process of dispersal can potentially have implications for ensuing recruitment and population replenishment. Our study not only shows that freshwater species that spend part of their life cycle in the ocean might also be affected by ocean acidification, but that behavioural responses towards key physicochemical cues can also be negated through elevated CO₂ from human emissions.     

Ocean acidification affects prey detection by a predatory reef fish

Changes in olfactory-mediated behaviour caused by elevated CO(2) levels in the ocean could affect recruitment to reef fish populations because larval fish become more vulnerable to predation. However, it is currently unclear how elevated CO(2) will impact the other key part of the predator-prey interaction--the predators. We investigated the effects of elevated CO(2) and reduced pH on olfactory preferences, activity levels and feeding behaviour of a common coral reef meso-predator, the brown dottyback (Pseudochromis fuscus). Predators were exposed to either current-day CO(2) levels or one of two elevated CO(2) levels (～600 μatm or ～950 μatm) that may occur by 2100 according to climate change predictions. Exposure to elevated CO(2) and reduced pH caused a shift from preference to avoidance of the smell of injured prey, with CO(2) treated predators spending approximately 20% less time in a water stream containing prey odour compared with controls. Furthermore, activity levels of fish was higher in the high CO(2) treatment and feeding activity was lower for fish in the mid CO(2) treatment; indicating that future conditions may potentially reduce the ability of the fish to respond rapidly to fluctuations in food availability. Elevated activity levels of predators in the high CO(2) treatment, however, may compensate for reduced olfactory ability, as greater movement facilitated visual detection of food. Our findings show that, at least for the species tested to date, both parties in the predator-prey relationship may be affected by ocean acidification. Although impairment of olfactory-mediated behaviour of predators might reduce the risk of predation for larval fishes, the magnitude of the observed effects of elevated CO(2) acidification appear to be more dramatic for prey compared to predators. Thus, it is unlikely that the altered behaviour of predators is sufficient to fully compensate for the effects of ocean acidification on prey mortality.     

Ontogenetic changes in responses to settlement cues by Anemonefish

Population connectivity for most marine species is dictated by dispersal during the pelagic larval stage. Although reef fish larvae are known to display behavioral adaptations that influence settlement site selection, little is known about the development of behavioral preferences throughout the larval phase. Whether larvae are attracted to the same sensory cues throughout their larval phase, or exhibit distinct ontogenetic shifts in sensory preference is unknown. Here, we demonstrate an ontogenetic shift in olfactory cue preferences for two species of anemonefish, a process that could aid in understanding both patterns of dispersal and settlement. Aquarium-bred na?ve Amphiprion percula and A. melanopus larvae were tested for olfactory preference of relevant reef-associated chemical cues throughout the 11-day pelagic larval stage. Age posthatching had a significant effect on the preference for olfactory cues from host anemones and live corals for both species. Preferences of olfactory cues from tropical plants of A. percula, increased by approximately ninefold between hatching and settlement, with A. percula larvae showing a fivefold increase in preference for the olfactory cue produced by the grass species. Larval age had no effect on the olfactory preference for untreated seawater over the swamp-based tree Melaleuca nervosa, which was always avoided compared with blank seawater. These results indicate that reef fish larvae are capable of utilizing olfactory cues early in the larval stage and may be predisposed to disperse away from reefs, with innate olfactory preferences drawing newly hatched larvae into the pelagic environment. Toward the end of the larval phase, larvae become attracted to the olfactory cues of appropriate habitats, which may assist them in identification of and navigation toward suitable settlement sites.     

Behavioural crypsis in a South African galaxiid fish induced by predatory and non-predatory heterospecifics

This study investigated recognition of, and behavioural responses to, predatory and non-predatory heterospecifics by a small cryptically coloured fish species, Galaxias ‘nebula’. Nebula recognised and differentiated between predatory and non-predatory heterospecifics and altered its behaviour facultatively. With both predatory and non-predatory fishes, the proportion of time spent motionless increased, whereas refugia use was affected only by predators and neither heterospecific affected the time spent active. Although nebula appeared to face no conflict, in that their responses to predatory and non-predatory heterospecifics varied in the same direction and differed only in intensity, the presence of both heterospecifics together induced responses midway between those for each heterospecific separately. Non-predatory heterospecifics thus modified nebula's responses towards predators, potentially making time available for other essential activities such as foraging. This modified predator response may aid its survival in an increasingly threatened habitat.     

How Nemo finds home: the neuroecology of dispersal and of population connectivity in larvae of marine fishes

Nearly all demersal teleost marine fishes have pelagic larval stages lasting from several days to several weeks, during which time they are subject to dispersal. Fish larvae have considerable swimming abilities, and swim in an oriented manner in the sea. Thus, they can influence their dispersal and thereby, the connectivity of their populations. However, the sensory cues marine fish larvae use for orientation in the pelagic environment remain unclear. We review current understanding of these cues and how sensory abilities of larvae develop and are used to achieve orientation with particular emphasis on coral-reef fishes. The use of sound is best understood; it travels well underwater with little attenuation, and is current-independent but location-dependent, so species that primarily utilize sound for orientation will have location-dependent orientation. Larvae of many species and families can hear over a range of ~100-1000 Hz, and can distinguish among sounds. They can localize sources of sounds, but the means by which they do so is unclear. Larvae can hear during much of their pelagic larval phase, and ontogenetically, hearing sensitivity, and frequency range improve dramatically. Species differ in sensitivity to sound and in the rate of improvement in hearing during ontogeny. Due to large differences among-species within families, no significant differences in hearing sensitivity among families have been identified. Thus, distances over which larvae can detect a given sound vary among species and greatly increase ontogenetically. Olfactory cues are current-dependent and location-dependent, so species that primarily utilize olfactory cues will have location-dependent orientation, but must be able to swim upstream to locate sources of odor. Larvae can detect odors (e.g., predators, conspecifics), during most of their pelagic phase, and at least on small scales, can localize sources of odors in shallow water, although whether they can do this in pelagic environments is unknown. Little is known of the ontogeny of olfactory ability or the range over which larvae can localize sources of odors. Imprinting on an odor has been shown in one species of reef-fish. Celestial cues are current- and location-independent, so species that primarily utilize them will have location-independent orientation that can apply over broad scales. Use of sun compass or polarized light for orientation by fish larvae is implied by some behaviors, but has not been proven. Use of neither magnetic fields nor direction of waves for orientation has been shown in marine fish larvae. We highlight research priorities in this area.     

Altered kelp (Laminariales) phlorotannins and growth under elevated carbon dioxide and ultraviolet-B treatments can influence associated intertidal food webs

Due to the importance of brown algae, such as kelp (Laminariales, Phaeophyta), within most cool nearshore environments, any direct responses of kelp to multiple global changes could alter the integrity of future coastal marine systems. Fifty-five-day manipulation of carbon dioxide (CO₂) and ultraviolet light (UVB) within outdoor sea-tanks, approximating past, present and two predicted future levels, examined the direct influences on Saccharina latissima (= Laminaria saccharina ) and Nereocystis luetkeana development and biochemistry, as well as the indirect influences on a marine herbivore ( Tegula funebralis ; Gastropoda, Mollusca) and on naturally occurring intertidal detritivores. Kelp species displayed variable directional (negative and positive growth) and scale responses to CO₂ and UVB manipulations, which was influenced by interactions. Kelp phlorotannin (phenolic) production in blade tissues was induced by elevated UVB levels, and especially enhanced (additively) by elevated CO₂, further suggesting that some actively growing kelp species are carbon limited in typical nearshore environments. Negative indirect effects upon detritivore consumers fed CO₂-manipulated kelp blade tissues were detected, however, no statistical relationships existed among UVB-treated tissues, and test herbivores did not distinguish between phlorotannin-altered CO₂: UVB-treated kelp blade tissues. Results suggest that past and future conditions differentially benefit these kelp species, which implies a potential for shifts in species abundance and community composition. Higher CO₂ conditions can indirectly impede marine decay processes delaying access to recycled trace nutrients, which may be disruptive to the seasonal regrowth of algae and/or higher trophic levels of nearshore ecosystems.     

The fish fauna of the Jordan reservoir, one of the oldest man-made lakes in central Europe

The Jordan reservoir, Czechoslovakia, was constructed in 1492 and the last full drainage and fish removal was in 1830. In this 'mature' fish community of 20 species (plus three hybrids), the roach, Rutilus rutilus (L.) (1446 fish ha^–1) and beam, Abramis brama (L.) (1074 fish ha^–1) dominated the fish stock (total of all species = 3628 fish ha^–1). Predatory species made a small contribution to the total (maximum 7.7%, obligatory predators 2%). The contribution (12%) by perch, Perca fluviatilis L., was also low, and its numbers in successive age-groups showed a steady decrease. The high variability and time-synchronization in the year-class strengths of the principal cyprinid species may indicate unstable environmental conditions or intra-community cycles. The growth rates of non-predatory species were low. There was a high fishing rate on predatory species but there was little impact by fishery management on the whole 'mature' community. Intensive eutrophication from both communal waters and agriculture probably caused the high biomass level (all species) and the low abundance of zander, Stizostedion lucioperca (L.).     

The relative importance of lethal and non-lethal effects of fish on insect colonisation of ponds

1. We hypothesised that adult insects actively monitor potential habitats for the presence of fish by means of chemical cues and avoid sites that pose significant risks. This was examined by quantifying colonisation of insects in outdoor pools with no fish (controls), fish (direct predation effect) or caged fish (chemical predator cues).
2. A significant direct effect of predation was found, but no indirect effect (avoidance of chemical cue pools), on the total biomass of colonising insects. However, predatory insects avoided fish-cue pools, thus releasing non-predatory insects from predation. This resulted in significantly greater biomass of non-predatory insects in fish-cue pools than control pools.
3. Fish reduced the number of species of colonising insects in pools through predation. This negative influence of fish implies that caution is necessary when stocking wetlands and ponds with fish if the goal is to maximise biodiversity.
4. Our data suggest that although predatory aquatic insects may use chemical signals to assess the quality of potential habitats with respect to predation risk, direct predation is the main method by which fish affect insect assemblages in ponds. Because fish and invertebrate predators may both have strong effects on prey mortality, behavioural adjustment by insects to the actual predator regime within a habitat should be more important than avoiding colonisation of habitats with fish.     

Face your fears: cleaning gobies inspect predators despite being stressed by them

social stressors typically elicit two distinct behavioural responses in vertebrates: an active response (i.e., "fight or flight") or behavioural inhibition (i.e., freezing). Here, we report an interesting exception to this dichotomy in a Caribbean cleaner fish, which interacts with a wide variety of reef fish clients, including predatory species. Cleaning gobies appraise predatory clients as potential threat and become stressed in their presence, as evidenced by their higher cortisol levels when exposed to predatory rather than to non-predatory clients. Nevertheless, cleaning gobies neither flee nor freeze in response to dangerous clients but instead approach predators faster (both in captivity and in the wild), and interact longer with these clients than with non-predatory clients (in the wild). We hypothesise that cleaners interrupt the potentially harmful physiological consequences elicited by predatory clients by becoming increasingly proactive and by reducing the time elapsed between client approach and the start of the interaction process. The activation of a stress response may therefore also be responsible for the longer cleaning service provided by these cleaners to predatory clients in the wild. Future experimental studies may reveal similar patterns in other social vertebrate species when, for instance, individuals approach an opponent for reconciliation after a conflict.     

Ocean acidification and responses to predators: can sensory redundancy reduce the apparent impacts of elevated CO2 on fish?

Carbon dioxide (CO₂) levels in the atmosphere and surface ocean are rising at an unprecedented rate due to sustained and accelerating anthropogenic CO₂ emissions. Previous studies have documented that exposure to elevated CO₂ causes impaired antipredator behavior by coral reef fish in response to chemical cues associated with predation. However, whether ocean acidification will impair visual recognition of common predators is currently unknown. This study examined whether sensory compensation in the presence of multiple sensory cues could reduce the impacts of ocean acidification on antipredator responses. When exposed to seawater enriched with levels of CO₂ predicted for the end of this century (880 μatm CO₂), prey fish completely lost their response to conspecific alarm cues. While the visual response to a predator was also affected by high CO_2, it was not entirely lost. Fish exposed to elevated CO_2, spent less time in shelter than current‐day controls and did not exhibit antipredator signaling behavior (bobbing) when multiple predator cues were present. They did, however, reduce feeding rate and activity levels to the same level as controls. The results suggest that the response of fish to visual cues may partially compensate for the lack of response to chemical cues. Fish subjected to elevated CO₂ levels_, and exposed to chemical and visual predation cues simultaneously, responded with the same intensity as controls exposed to visual cues alone. However, these responses were still less than control fish simultaneously exposed to chemical and visual predation cues. Consequently, visual cues improve antipredator behavior of CO₂ exposed fish, but do not fully compensate for the loss of response to chemical cues. The reduced ability to correctly respond to a predator will have ramifications for survival in encounters with predators in the field, which could have repercussions for population replenishment in acidified oceans.     

Carbon dioxide-concentrating mechanism and the development of extracellular carbonic anhydrase in the marine picoeukaryote Micromonas pusilla

A range of marine photosynthetic picoeukaryote phytoplankton species grown in culture were screened for the presence of extracellular carbonic anhydrase (CA_ext), a key enzyme in inorganic carbon acquisition under carbon- limiting conditions in some larger marine phytoplankton species. Of the species tested, extracellular carbonic anhydrase was detected only in Micromonas pusilla Butcher. The rapid, light-dependent development of CA_ext when cells were transferred from carbon-replete to carbon-limiting conditions was regulated by the available free- CO₂ concentration and not by total dissolved inorganic carbon. Kinetic studies provided support for a CO₂- concentrating mechanism in that the K _0.5[CO₂] (i.e. the CO₂ concentration required for the half-maximal rate of photosynthesis) was substantially lower than the K _m[CO₂] of Rubisco from related taxa, whilst the intracellular carbon pool was at least seven fold greater than the extracellular DIC concentration, for extracellular DIC values 1.0 m m .
It is proposed that when the flux of CO₂ into the cell is insufficient to support the photosynthetic rate at an optimum photon irradiance, the development of CA_ext increases the availability of CO₂ at the plasma membrane. This ensures rapid acclimation to environmental change and provides an explanation for the central role of M. pusilla as a carbon sink in oligotrophic environments.     

Auditory and olfactory abilities of larvae of the Indo-Pacific coral trout Plectropomus leopardus (Lacepède) at settlement

Auditory and olfactory abilities of settlement-stage larvae of the coral trout Plectropomus leopardus (Pisces: Serranidae) were tested electrophysiologically to determine if these senses are sufficiently developed to aid larvae in detection of settlement habitats on coral reefs. Plectropomus leopardus larvae detected sounds from 100 to 2000 Hz with hearing most sensitive at the frequencies of 100, 200 and 600 Hz. The olfactory response of P. leopardus was similar for the two amino acids tested and for the water conditioned by conspecifics. Auditory and olfactory abilities of P. leopardus are well developed at settlement-stage, and apparently sufficient to detect auditory and olfactory cues from reefs.     

AN ECOLOGICAL BASIS FOR EXTRACELLULAR CARBONIC ANHYDRASE IN MARINE UNICELLULAR ALGAE

Extracellular carbonic anhydrase (CA_e) is expressed by many, but not all, autotrophic species of aquatic unicellular protists. We measured CA_e activities in unicellular marine algae characteristic of either high nutrient spring, fall, and winter blooms or low nutrient summer populations to provide ecological/evolutionary information about the enzyme. Highest activities occurred in spring bloom and opportunistic diatoms exposed to long photoperiods (16 h) when pH was highest and CO₂ was lowest. Lower activities were recorded for a fall-bloom diatom exposed to the long photoperiod, and lowest values were found under all culture conditions for one diatom and a number of flagellated species typical of summer low nutrient environments. Other potential sources of variance in measurements of CA_e activity were examined. Maximum activities of CA_e were recorded for the diatom, Skeletonema costatum (Greville) Cleve, during late exponential phase of growth and within 8 h of the beginning of the photoperiod. We concluded that ecological factors are important in determining CA_e activities in marine unicellular protists. Potential functions of CA_e in the metabolism of marine unicellular algae are discussed.     

Reef Odor: A Wake Up Call for Navigation in Reef Fish Larvae

The behavior of reef fish larvae, equipped with a complex toolbox of sensory apparatus, has become a central issue in understanding their transport in the ocean. In this study pelagic reef fish larvae were monitored using an unmanned open-ocean tracking device, the drifting in-situ chamber (DISC), deployed sequentially in oceanic waters and in reef-born odor plumes propagating offshore with the ebb flow. A total of 83 larvae of two taxonomic groups of the families Pomacentridae and Apogonidae were observed in the two water masses around One Tree Island, southern Great Barrier Reef. The study provides the first in-situ evidence that pelagic reef fish larvae discriminate reef odor and respond by changing their swimming speed and direction. It concludes that reef fish larvae smell the presence of coral reefs from several kilometers offshore and this odor is a primary component of their navigational system and activates other directional sensory cues. The two families expressed differences in their response that could be adapted to maintain a position close to the reef. In particular, damselfish larvae embedded in the odor plume detected the location of the reef crest and swam westward and parallel to shore on both sides of the island. This study underlines the critical importance of in situ Lagrangian observations to provide unique information on larval fish behavioral decisions. From an ecological perspective the central role of olfactory signals in marine population connectivity raises concerns about the effects of pollution and acidification of oceans, which can alter chemical cues and olfactory responses.     

Effects of graded hypoxia on Atlantic salmon infected with amoebic gill disease

Atlantic salmon Salmo salar with amoebic gill disease (AGD) were exposed to a graded hypoxia (135–40 mmHg water P O₂) and blood samples analysed for respiratory gases and pH at 119, 79·5 and 40 mmHg water P O₂. There were no differences in the rate of oxygen uptake between infected and control fish. However, arterial P O₂, and pH were significantly lower in the infected fish whereas P CO₂ was significantly higher in infected fish compared with controls prior to hypoxia and at 119 mmHg water P O₂. At 79·5 and 40 mmHg water P O₂ saturation, there were no significant differences in blood P O₂ or pH although blood P CO₂ was elevated in AGD affected fish at 50% hypoxia (79·5 mmHg water P O₂). The elevated levels of P CO₂ in fish affected by AGD resulted in a persistent respiratory acidosis even during hypoxic challenge. These data suggest that even though the fish were severely affected by AGD, the presence of AGD while impairing gas transfer under normoxic conditions, did not contribute to respiratory failure during hypoxia.     

The role of the mesophyll in stomatal responses to light and CO2

Stomatal responses to light and CO₂ were investigated using isolated epidermes of Tradescantia pallida , Vicia faba and Pisum sativum . Stomata in leaves of T. pallida and P. sativum responded to light and CO₂, but those from V. faba did not. Stomata in isolated epidermes of all three species could be opened on KCl solutions, but they showed no response to light or CO₂. However, when isolated epidermes of T. pallida and P. sativum were placed on an exposed mesophyll from a leaf of the same species or a different species, they regained responsiveness to light and CO₂. Stomatal responses in these epidermes were similar to those in leaves in that they responded rapidly and reversibly to changes in light and CO₂. Epidermes from V. faba did not respond to light or CO₂ when placed on mesophyll from any of the three species. Experiments with single optic fibres suggest that stomata were being regulated via signals from the mesophyll produced in response to light and CO₂ rather than being sensitized to light and CO₂ by the mesophyll. The data suggest that most of the stomatal response to CO₂ and light occurs in response to a signal generated by the mesophyll.