Shrinking body size and climate warming: Many freshwater salmonids do not follow the rule

Declining body size is believed to be a universal response to climate warming and has been documented in numerous studies of marine and anadromous fishes. The Salmonidae are a family of coldwater fishes considered to be among the most sensitive species to climate warming; however, whether the shrinking body size response holds true for freshwater salmonids has yet to be examined at a broad spatial scale. We compiled observations of individual fish lengths from long-term surveys across the Northern Hemisphere for 12 species of freshwater salmonids and used linear mixed models to test for spatial and temporal trends in body size (fish length) spanning recent decades. Contrary to expectations, we found a significant increase in length overall but with high variability in trends among populations and species. More than two-thirds of the populations we examined increased in length over time. Secondary regressions revealed larger-bodied populations are experiencing greater increases in length than smaller-bodied populations. Mean water temperature was weakly predictive of changes in body length but overall minimal influences of environmental variables suggest that it is difficult to predict an organism's response to changing temperatures by solely looking at climatic factors. Our results suggest that declining body size is not universal, and the response of fishes to climate change may be largely influenced by local factors. It is important to know that we cannot assume the effects of climate change are predictable and negative at a large spatial scale.     

Ecophysiological patterns of distribution and behavior of freshwater fish in thermal gradients

Species-specific, intraspecific, age-related, seasonal, diel, physiological, and behavioral aspects of the distribution of freshwater fishes along experimental and natural thermal gradients are analyzed. It is shown that thermal selection response is displayed by all fish species. The thermal selection process includes two stages: the choice of the current selected temperature, which reflects the acclimation process, and the choice of the final selected temperature, which follows the stabilization of thermal selection. Thermoregulation behavior is an inherent reaction and can be placed among the most shared and earliest adaptations of aquatic animals. The final selected temperature of fishes reflects the ecophysiological optimum of the existence of the species, and the upper lethal temperature reflects the limit of potential existence.     

Reproduction of the annual fish Austrolebias nigrofasciatus (Rivulidae) maintained at different temperatures

Temperature is considered an important factor that influences the life cycle of annual fishes, however the thermal preferendum of Austrolebias nigrofasciatus reproduction has not been defined. The hypothesis was that the fecundity of the species would respond differently within the temperature range analyzed, presenting an optimum thermal. Thus, this study investigated the effect of temperature on the fecundity of A. nigrofasciatus for the first time under laboratory conditions. Pairs of A. nigrofasciatus were placed in aquaria with temperatures of 17, 21 or 25°C, with four pairs (replicates) for each treatment, for five weeks. Contrary to expected, there were no significant differences in fecundity parameters among the tested temperatures, but the weekly fecundity at 17 and 21°C was more homogeneous than at 25°C. During the experimental period, the temperature did not affect male growth; however, at 25°C there was a negative effect on female body growth. The condition factor was also influenced, suggesting a tendency towards a negative energy balance with increasing temperature. The results indicate that temperatures between 17 and 21°C are suitable for broodstock maintenance.     

Seed dormancy in Rumex species in response to environmental factors

Abstract. Many Rumex species show similar seed dormancy characteristics but there is more information concerning R. crispus and R. obtusifolius than other species. These species respond positively to red or white light. Far-red light applied for short periods may promote or inhibit germination depending on the timing of the irradiation in relation to temperature change; but long periods of far-red inhibit germination. Seeds may also be stimulated to germinate in the dark by low-temperature stratification at 15°C or less providing the temperature of the seeds is subsequently raised to a minimum of about 15°C. Seeds can, however, germinate at lower temperatures providing they have received other appropriate stimulatory treatment. Seeds also respond to alternating temperatures. In a diurnal cycle the minimum upper temperature required is about 15°C and the maximum lower temperature is about 25°C. The optimum period spent at the upper temperature is about 8 h when it is 15–25°C but the optimum period decreases as the upper temperature is increased above this range so that at 45°C, for example, it is only about 30 min. The period spent at the lower temperature in a diurnal cycle is not critical. Providing these criteria are met, the percentage germination increases with the number and amplitude of the cycles. The warming part of the cycle is necessary for the response but so far there is no convincing evidence that cooling itself is important. Secondary dormancy is induced at constant temperatures at a rate dependent on temperature, but apparently only in the presence of oxygen. This feature affects the optimum timing of a temperature change or exposure to light. Strong positive interactions are shown between stimulatory temperature treatments and white or red light. Unlike many other weed species the seeds respond only slightly to nitrate ions. The implications of these responses are discussed in relation to field behaviour.     

Characteristics of alternating temperatures which stimulate loss of dormancy in seeds of Rumex obtusifolius L. and Rumex crispus L.

Abstract. Alternating temperatures stimulate the germination of Rumex crispus L. and Rumex obtusifolius L. The optimum period spent at the lower temperature in a diurnal cycle is greater than that spent at the higher temperature. Under most conditions the optimum period at the upper temperatures is about 8 h but, as the upper temperature of a cycle is increased, the optimum period at the upper temperature becomes shorter and more critical. Thus when it is 35°C the optimum period is 2.5–4 h in the light, or about 1 h in the dark. The effect of alternating temperatures is much less in the dark than in the light and in general only extreme alternations with short periods at the higher temperature are effective in the dark. In the light any temperature alternation within the range 1–35°C is effective to at least some extent, providing the temperature difference is 5°C or more and providing the alternation includes one temperature which is above approximately 15°C and one which is below approximately 25°C. The optimum temperature difference is about 15°C. In the light, 4 to 10 cycles saturate the response, but in the dark, where the effect is much less, the response may not be saturated even by 16 cycles. KNO₃ at 10⁻³ M has little effect on the response to alternating temperatures either in the light or the dark. The response to alternating temperature regimes does not appear to vary in quality, i.e., in terms of which particular treatments are best, but it varies in magnitude with site and year of seed collection; and it increases slowly during dry storage, even when stored at a temperature as low as 1.5°C.     

Fish can show emotional fever: stress-induced hyperthermia in zebrafish

Whether fishes are sentient beings remains an unresolved and controversial question. Among characteristics thought to reflect a low level of sentience in fishes is an inability to show stress-induced hyperthermia (SIH), a transient rise in body temperature shown in response to a variety of stressors. This is a real fever response, so is often referred to as ‘emotional fever’. It has been suggested that the capacity for emotional fever evolved only in amniotes (mammals, birds and reptiles), in association with the evolution of consciousness in these groups. According to this view, lack of emotional fever in fishes reflects a lack of consciousness. We report here on a study in which six zebrafish groups with access to a temperature gradient were either left as undisturbed controls or subjected to a short period of confinement. The results were striking: compared to controls, stressed zebrafish spent significantly more time at higher temperatures, achieving an estimated rise in body temperature of about 2–4°C. Thus, zebrafish clearly have the capacity to show emotional fever. While the link between emotion and consciousness is still debated, this finding removes a key argument for lack of consciousness in fishes.     

Immune defense and host life history

Recent interest has focused on immune response in an evolutionary context, with particular attention to disease resistance as a life-history trait, subject to trade-offs against other traits such as reproductive effort. Immune defense has several characteristics that complicate this approach, however; for example, because of the risk of autoimmunity, optimal immune defense is not necessarily maximum immune defense. Two important types of cost associated with immunity in the context of life history are resource costs, those related to the allocation of essential but limited resources, such as energy or nutrients, and option costs, those paid not in the currency of resources but in functional or structural components of the organism. Resource and option costs are likely to apply to different aspects of resistance. Recent investigations into possible trade-offs between reproductive effort, particularly sexual displays, and immunity have suggested interesting functional links between the two. Although all organisms balance the costs of immune defense against the requirements of reproduction, this balance works out differently for males than it does for females, creating sex differences in immune response that in turn are related to ecological factors such as the mating system. We conclude that immune response is indeed costly and that future work would do well to include invertebrates, which have sometimes been neglected in studies of the ecology of immune defense.     

Sex-specific effects of carotenoid intake on the immunological response to allografts in guppies (Poecilia reticulata)

Rarely are the evolutionary origins of mate preferences known, but, recently, the preference of female guppies (Poecilia reticulata) for males with carotenoid-based sexual coloration has been linked to a sensory bias that may have originally evolved for detecting carotenoid-rich fruits. If carotenoids enhance the immune systems of these fishes, as has been suggested for other species, this could explain the origin of the attraction to orange fruits as well as the maintenance of the female preference for orange males. We used the classic immunological technique of tissue grafting to assay a component of the immune response of guppies raised on two different dietary levels of carotenoids. Individual scales were transplanted between pairs of unrelated fishes, creating reciprocal allografts. Transplanted scales were scored on a six-point rejection scale every day for 10 days. Five days later, the same pairs of fishes received a second set of allografts and were scored again. Compared with low-carotenoid-diet males, high-carotenoid-diet males mounted a significantly stronger rejection response to the second allograft but not to the first allograft. High-carotenoid-diet females, however, showed no improvement in graft rejection compared with low-carotenoid-diet females. To our knowledge, this is the first experimental evidence for sex-specific effects of carotenoid consumption on the immune system of a species with carotenoid-based sexual coloration. These results are consistent with the hypothesis that the mate preference for carotenoid coloration is maintained by the benefits to females of choosing healthy mates, but they cast doubt on the idea that the benefits of carotenoid consumption, per se, could account for the origin of the preference. The sex-specificity of carotenoid effects on allograft rejection in guppies provides indirect support for the general hypothesis that males pay an immunological cost for sexual ornamentation.     

Temperature Tolerances of North American Freshwater Fishes Exposed to Dynamic Changes in Temperature

Traditionally lower and upper temperature tolerances of fishes have been quantified in the laboratory via three different experimental approaches: the Fry or incipient lethal temperature (ILT), critical thermal (CTM) and chronic lethal (CLM) methodologies. Although these three experimental laboratory approaches generate endpoints which are quantitatively expressed as a temperature, are determined experimentally with random samples of fish acclimated to specific temperatures, and involve both time and temperature as major test variables, they do not quantify the same response. All three approaches generate valuable, albeit different, information concerning the temperature tolerance of a species. In this review we have summarized published research concerning the tolerance of North American freshwater fishes to dynamic changes in temperature, i.e., tolerance is tested by methods that gradually change temperatures until biological stress is observed. We found more than 450 individual temperature tolerances listed in 80 publications which present original dynamic temperature tolerance data for 116 species, 7 subspecies and 7 hybrids from 19 families of North American freshwater fishes. This total represents about 1/3 of the families and 1/6 of the known North American freshwater species. Temperature tolerance data were partitioned by experimental approach, i.e., critical thermal method (CTM) and chronic lethal method (CLM), and direction of temperature change. Although both CTM and CLM expose fish to dynamic changes in water temperature, these two methods differ in temperature change rates and test endpoints, and hence measure different aspects of thermal stress. A majority of the 80 studies employed CTM to assess temperature tolerance, in particular determination of CTmaxima. One or more CTmaxima has been reported for 108 fishes. Twenty-two fishes have reported highest CTmaxima of 40°C or higher. Several species in the family Cyprinodontidae have generated some of the highest CTmaxima reported for any ectothermic vertebrate. For a variety of reasons, data concerning tolerance of low temperatures are less plentiful. Low temperature tolerance quantified as either CTminima or CLminima were found for a total of 37 fishes. Acclimation temperature exerts a major effect on the temperature tolerance of most North American fish species and it is usually strongly linearly related to both CTmaxima and CTminima. Although we uncovered dynamic temperature tolerance data for 130 fishes, only a single dynamic, temperature tolerance polygon has been published, that for the sheepshead minnow, Cyprinodon variegatus.     

Effects of carotenoids, immune activation and immune suppression on the intensity of chronic coccidiosis in greenfinches

Allocation trade-offs of carotenoids between their use in the immune system and production of integumentary colouration have been suggested as a proximate mechanism maintaining honesty of signal traits. We tested how dietary carotenoid supplementation, immune activation and immune suppression affect intensity of coccidian infection in captive greenfinches Carduelis chloris, a passerine with carotenoid-based plumage. Immune activation with phytohaemagglutinin (PHA) decreased body mass among birds not supplemented with lutein, while among the carotenoid-fed birds, PHA had no effect on mass dynamics. Immune suppression with dexamethasone (DEX) induced loss of body mass and reduced the swelling response to PHA. DEX and PHA increased the concentration of circulating heterophils. Lutein supplementation increased plasma carotenoid levels but had no effect on the swelling response induced by PHA. PHA and DEX treatments did not affect plasma carotenoids. Immune stimulation by PHA suppressed the infection, but only among carotenoid-supplemented birds. Priming of the immune system can thus aid in suppressing chronic infection but only when sufficient amount of carotenoids is available. Our experiment shows the importance of carotenoids in immune response, but also the complicated nature of this impact, which could be the reason for inconsistent results in studies investigating the immunomodulatory effects of carotenoids. The findings about involvement of carotenoids in modulation of an immune response against coccidiosis suggest that carotenoid-based ornaments may honestly signal individuals’ ability to manage chronic infections.     

Ocean warming has a greater effect than acidification on the early life history development and swimming performance of a large circumglobal pelagic fish

Ocean warming and acidification are serious threats to marine life; however, their individual and combined effects on large pelagic and predatory fishes are poorly understood. We determined the effects of projected future temperature and carbon dioxide (CO₂) levels on survival, growth, morphological development and swimming performance on the early life stages of a large circumglobal pelagic fish, the yellowtail kingfish Seriola lalandi. Eggs, larvae and juveniles were reared in cross‐factored treatments of temperature (21 and 25°C) and pCO₂ (500 and 985 μatm) from fertilisation to 25 days post hatching (dph). Temperature had the greatest effect on survival, growth and development. Survivorship was lower, but growth and morphological development were faster at 25°C, with surviving fish larger and more developed at 1, 11 and 21 dph. Elevated pCO₂ affected size at 1 dph, but not at 11 or 21 dph, and did not affect survival or morphological development. Elevated temperature and pCO₂ had opposing effects on swimming performance at 21 dph. Critical swimming speed (U_crit) was increased by elevated temperature but reduced by elevated pCO₂. Additionally, elevated temperature increased the proportion of individuals that responded to a startle stimulus, reduced latency to respond and increased maximum escape speed, potentially due to the more advanced developmental stage of juveniles at 25°C. By contrast, elevated pCO₂ reduced the distance moved and average speed in response to a startle stimulus. Our results show that higher temperature is likely to be the primary driver of global change impacts on kingfish early life history; however, elevated pCO₂ could affect critical aspects of swimming performance in this pelagic species. Our findings will help parameterise and structure fisheries population dynamics models and improve projections of impacts to large pelagic fishes under climate change scenarios to better inform adaptation and mitigation responses.     

Heterochrony and the development of the escape response: prehatching movements in the rainbow trout Oncorhynchus mykiss

Teleost fishes produce coordinated escape responses (C-starts) at hatching. This implies that essential swimming morphologies and motor behaviors develop during the incubation interval while the embryo is in the chorion. We examined prehatching motor behaviors in rainbow trout Oncorhycus mykiss (considered morphologically mature at hatching) and compared this species with zebrafish Danio rerio (considered morphologically immature) and assessed two hypotheses concerning the development of escape behavior. (1) Escape behavior is associated with the formation of key elements of the musculoskeletal and nervous systems; thus, the escape response appears early in ontogeny, when these elements form. (2) Escape behavior is not directly associated with the formation of underlying morphological elements; instead, it appears at hatching (i.e. when needed). We find that rainbow trout, like zebrafish, respond to touch early in the incubation interval, but do not demonstrate a complete C-start (including the second, propulsive stage) until shortly before hatching. At hatching, rainbow trout and zebrafish are similar in the degree of development of the chondocranium, paired fins and visceral arches (which comprise the larval jaw and gill support); however, rainbow trout have incipient rays in their unpaired fins (dorsal, anal and caudal), whereas zebrafish retain the embryonic fin fold. Although rainbow trout are more mature in axial swimming morphology at hatching, the essential neural and musculoskeletal systems that produce a coordinated escape response are functional at hatching in both species. This finding supports the evolutionary hypothesis that an effective escape response is critical for the survival of newly hatched teleost fishes.     

Stress and welfare in ornamental fishes: what can be learned from aquaculture?

The ornamental fish trade is estimated to handle up to 1·5 billion fishes. Transportation and handling of fishes imposes a range of stressors that can result in mortality at rates of up to 73%. These rates vary hugely, however, and can be as low as 2%, because they are generally estimated rather than based on experimental work. Given the numbers of ornamental fishes traded, any of the estimated mortality rates potentially incur significant financial losses and serious welfare issues. Industry bodies, such as the Ornamental Aquatic Trade Association (OATA), have established standards and codes of best practice for handling fishes, but little scientific research has been conducted to understand the links between stress, health and welfare in ornamental species. In aquaculture, many of the same stressors occur as those in the ornamental trade, including poor water quality, handling, transportation, confinement, poor social and physical environment and disease and in this sector directed research and some resulting interventions have resulted in improved welfare standards. This review considers the concept of welfare in fishes and evaluates reported rates of mortality in the ornamental trade. It assesses how the stress response can be quantified and used as a welfare indicator in fishes. It then analyses whether lessons from aquaculture can be usefully applied to the ornamental fish industry to improve welfare. Finally, this analysis is used to suggest how future research might be directed to help improve welfare in the ornamental trade.     

Effects of temperature on the food processing of three species of seaweed-eating fishes from European coastal waters

Food consumption, gut transit times and assimilation efficiencies of the temperate fishes Lipophrys pholis and Parablennius sanguinolentus (Blenniidae) and Gobius cobitis (Gobiidae. when fed the green alga Ulva lactuca were determined at four temperatures in the laboratory. Food consumption increased and gut transit times decreased with increase in temperature in all three species. Assimilation efficiencies for algal carbon and nitrogen did not vary with temperature in G. cobitis . Carbon but not nitrogen assimilation efficiency varied with temperature in P. sanguinolentus , and both carbon and nitrogen assimilation efficiencies changed broadly with temperature in L. pholis , the most omnivorous of the three fishes. Assimilation efficiencies of all three species were generally within the previously reported ranges for herbivorous marine fishes. These species are thus clearly capable of extracting energy from algal food sources.     

Metabolic cold adaptation in fishes occurs at the level of whole animal, mitochondria and enzyme

Metabolic cold adaptation (MCA), the hypothesis that species from cold climates have relatively higher metabolic rates than those from warm climates, was first proposed nearly 100 years ago and remains one of the most controversial hypotheses in physiological ecology. In the present study, we test the MCA hypothesis in fishes at the level of whole animal, mitochondria and enzyme. In support of the MCA hypothesis, we find that when normalized to a common temperature, species with ranges that extend to high latitude (cooler climates) have high aerobic enzyme (citrate synthase) activity, high rates of mitochondrial respiration and high standard metabolic rates. Metabolic compensation for the global temperature gradient is not complete however, so when measured at their habitat temperature species from high latitude have lower absolute rates of metabolism than species from low latitudes. Evolutionary adaptation and thermal plasticity are therefore insufficient to completely overcome the acute thermodynamic effects of temperature, at least in fishes.     

Habitat use by a Midwestern U.S.A. riverine fish assemblage: effects of season, water temperature and river discharge

The hypothesis that temperate stream fishes alter habitat use in response to changing water temperature and stream discharge was evaluated over a 1 year period in the Neosho River, Kansas, U.S.A. at two spatial scales. Winter patterns differed from those of all other seasons, with shallower water used less frequently, and low‐flow habitat more frequently, than at other times. Non‐random habitat use was more frequent at the point scale (4·5 m²) than at the larger reach scale (20–40 m), although patterns at both scales were similar. Relative to available habitats, assemblages used shallower, swifter‐flowing water as temperature increased, and shallower, slower‐flowing water as river discharge increased. River discharge had a stronger effect on assemblage habitat use than water temperature. Proportion of juveniles in the assemblage did not have a significant effect. This study suggests that many riverine fishes shift habitats in response to changing environmental conditions, and supports, at the assemblage level, the paradigm of lotic fishes switching from shallower, high‐velocity habitats in summer to deeper, low‐velocity habitats in winter, and of using shallower, low‐velocity habitats during periods of high discharge. Results also indicate that different species within temperate river fish assemblages show similar habitat use patterns at multiple scales in response to environmental gradients, but that non‐random use of available habitats is more frequent at small scales.     

Plasticity in animal personality traits: does prior experience alter the degree of boldness?

Theoreticians predict that animal 'personality' traits may be maladaptive if fixed throughout different contexts, so the present study aimed to test whether these traits are fixed or plastic. Rainbow trout (Onchorhyncus mykiss) were given emboldening or negative experiences in the forms of watching bold or shy individuals responding to novelty or winning or losing fights to examine whether prior experience affected boldness. Bold individuals that lost fights or watched shy demonstrators became more shy by increasing their latency to approach a novel object, whereas shy observers that watched bold demonstrators remained cautious and did not modify their responses to novelty. Shy winners became bolder and decreased their latency to approach a novel object, but shy losers also displayed this shift. In comparison, control groups showed no change in behaviour. Bold fishes given negative experiences reduced their boldness which may be an adaptive response; however, shy fishes may base their strategic decisions upon self-assessment of their relative competitive ability and increase their boldness in situations where getting to resources more quickly ensures they outcompete better competitors.     

Evaluating the roles of energetic functional constraints on teleost mitochondrial-encoded protein evolution

Mitochondria are the power plant of cells, which play critical roles not only in energy metabolism but also in thermoregulation. These two roles have been individually suggested to influence mitochondrial DNA (mtDNA) evolution, however their relative importance is still rarely considered. Here, we conduct a comparative genomic analysis of 401 teleost complete mitochondrial genomes and test the roles of these dual functional constraints on mitochondria to provide a more complete view of mtDNA evolution. We found that mitochondrial protein-coding genes of migratory fishes have significantly smaller Ka/Ks than nonmigratory fishes. The same data set showed that the genes of fishes living in cold climates have significantly smaller Ka/Ks than tropical fishes. In contrast, these trends were not observed for two nuclear genes that are not involved in energy metabolism. The differences in selection patterns observed between mitochondrial and nuclear genes suggest that the functional constraints acting on mitochondria, due to energy metabolism and/or thermoregulation, influence the evolution of mitochondrial-encoded proteins in teleosts.     

Effects of coral bleaching on the feeding response of two species of coral-feeding fish

Coral bleaching is an increasingly prominent threat to coral reef ecosystems, not only to corals, but also to the many organisms that rely on coral for food and shelter. Coral-feeding fishes are negatively affected by coral loss caused by extensive bleaching, but it is unknown how feeding behaviour of most corallivorous fishes changes in response to coral bleaching. In this study, coral bleaching was experimentally induced in situ to examine the feeding response of two obligate corallivorous fish, Labrichthys unilineatus (Labridae) and Chaetodon baronessa (Chaetodontidae). Feeding rates were monitored before, during, and immediately after experimental bleaching of prey corals. L. unilineatus significantly increased its feeding on impacted corals during bleaching, but showed a steady decline in feeding once corals were fully bleached. Feeding response of L. unilineatus appears to parallel the expected stress-induced mucous production by bleaching colonies. In contrast, C. baronessa preferentially fed from healthy colonies over bleached colonies, although bleached colonies were consumed for five days following manipulation. Feeding by corallivorous fishes can play an important role in determining coral condition and mortality of corals following stress induced bleaching.