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1) Through phenotypic plasticity, individual genotypes can produce multiple phenotypes dependent on the environment. In the modern world, anthropogenic influences such as man-made pharmaceuticals are increasingly prevalent. They might alter observable patterns of plasticity and distort our conclusions regarding the adaptive potential of natural populations. 2) Antibiotics are nowadays nearly ubiquitous in aquatic environments and prophylactic antibiotic use is also becoming more common to optimize animal survival and reproductive output in artificial settings. In the well-studied plasticity model system Physella acuta, prophylactic erythromycin treatment acts against gram-positive bacteria and thereby reduces mortality. 3) Here, we study its consequences for inducible defence formation in the same species. In a 2*2 split-clutch design, we reared 635 P. acuta in either the presence or absence of this antibiotic, followed by 28-day exposure to either high or low predation risk as perceived through conspecific alarm cues. 4) Under antibiotic treatment, risk-induced increases in shell thickness, a well-known plastic response in this model system, were larger and consistently detectable. Antibiotic treatment reduced shell thickness in low-risk individuals, suggesting that in controls, undiscovered pathogen infection increased shell thickness under low risk. Family variation in risk-induced plasticity was low, but the large variation in responses to antibiotics among families suggests different pathogen susceptibility between genotypes. Lastly, individuals that developed thicker shells had reduced total mass, which highlights resource trade-offs. 5) Antibiotics thus have the potential to uncover a larger extent of plasticity, but might counterintuitively distort plasticity estimates for natural populations where pathogens are a part of natural ecology. This article is protected by copyright. All rights reserved.

Vertebrate cellular immunity displays substantial variation among taxa and environments. Hematological parameters such as white blood-cell counts have emerged as a valuable tool to understand this variation by assessing the immunological status of individuals. These tools have long revealed that vertebrate cellular immune systems are highly plastic and respond to injury and infection. However, cellular immune systems may also be able to anticipate a high risk of injury from environmental cues (e.g., predation-related cues) and respond plastically ahead of time. We studied white blood-cell (leukocyte) profiles in African cichlids Pelvicachromis taeniatus that were raised for 4 years under different levels of perceived predation risk. In a split-clutch design, we raised fish from hatching onwards under chronic exposure to either conspecific alarm cues (communicating high predation risk) or a distilled water control treatment. Differential blood analysis revealed that alarm cue-exposed fish had twice as many lymphocytes in peripheral blood as did controls, a condition called lymphocytosis. The presence of a higher number of lymphocytes makes the cellular immune response more potent, which accelerates the removal of invading foreign antigens from the bloodstream, and, therefore, may be putatively beneficial in the face of injury. This observed lymphocytosis after long-term exposure to conspecific alarm cues constitutes first evidence for an anticipatory and adaptive plastic response of the cellular immune system to future immunological challenges. Electronic supplementary material The online version of this article (10.1007/s00442-020-04781-y) contains supplementary material, which is available to authorized users.

Antidepressant (AD) drugs are widely prescribed for the treatment of psychiatric disorders, including depression and anxiety disorders. The continuous use of ADs causes significant quantities of these bioactive chemicals to enter the aquatic ecosystems mainly through wastewater effluent discharge. This may result in many aquatic organisms being inadvertently affected by these drugs. Fluoxetine (FLX) and venlafaxine (VEN) are currently among the most widely detected ADs in aquatic systems. A growing body of experimental evidence demonstrates that FLX and VEN have a substantial capacity to induce neurotoxicity and cause behavioral dysfunctions in a wide range of teleost species. At the same time, these studies often report seemingly contradictory results that are confounding in nature. Hence, we clearly require comprehensive reviews that attempt to find overarching patterns and establish possible causes for these variable results. This review aims to explore the current state of knowledge regarding the neurobehavioral effects of FLX and VEN on fishes. This study also discusses the potential mechanistic linkage between the neurotoxicity of ADs and behavioral dysfunction and identifies key knowledge gaps and areas for future research. Copyright © 2021 Elsevier B.V. All rights reserved.

Recent studies show that bisphenol S (BPS) induces multiple adverse effects in exposed organisms; however, the maternal effects of BPS exposure remain poorly understood. Here, we expose adult female zebrafish to environmentally relevant concentrations of BPS (0, 1, 10, 30 μg/L) and 1 μg/L of 17-β-estradiol (E2) as a positive control for 60 days. Females were then paired with BPS-unexposed males and their offspring were raised in control water for 6 months. Maternal exposure to BPS was found to alter social behavior and anxiety response in a dose-specific manner in male offspring. Group preferences and social cohesion were significantly reduced by maternal exposure to 1 and 10 μg/L BPS, respectively. Additionally, maternal exposure to 1 and 30 μg/L BPS and E2 decreased offspring stress responses during the novel tank test. The impaired social behavior was associated with elevated arginine-vasotocin (AVT) level as well as with the altered expression of genes involved in AVT signaling pathway (AVT, avpr1aa) and enzymatic antioxidant genes (cat and Mn-sod) in the brain. Collectively, these results suggest that maternal exposure to environmentally relevant concentrations of BPS alters social behavior in zebrafish offspring, which is likely mediated by oxidative stress and disruption of neuropeptide signaling pathways in the brain.

Inclusive fitness theory predicts that individuals can increase their indirect fitness by grouping with kin. However, kin grouping also increases competition between kin, which potentially outweighs its benefits. The level of kin competition is contingent on environmental conditions and thus highly variable. Hence, individuals should benefit from plastically adjusting kin discrimination according to the expected level of kin competition. Here, we investigate whether perceived high competition affects juvenile kin-shoaling preferences in the cichlid Pelvicachromis taeniatus. Juveniles were given the choice between two shoals consisting of either kin or nonkin. Levels of perceived competition were manipulated through food limitation in the face of the differential energy expenditure of differently sized fish. The preference to shoal with kin decreased with increasing levels of perceived competition; small food-deprived individuals avoided kin. Shoaling with kin under strong competition may reduce individual indirect fitness. Hence, individuals can likely improve their inclusive fitness by plastically adjusting their kin-grouping preferences.

High predation risk during development induces phenotypic changes in animals. However, little is known about how these plastic responses affect signalling and competitiveness during contests. Herein, we have studied the consequences of anti-predator plasticity during the intra-sexual competition of Pelvicachromis taeniatus, a cichlid fish with mutual mate choice. We staged contests between adult size-matched siblings of the same sex derived from different environments: one fish was regularly exposed to conspecific alarm cues since the larval stage (simulating predator presence), the other fish to control conditions. Rearing environment did not affect the winner of contests or total aggression within a fight. However, contest behaviour differed between treatments. The effects were especially pronounced in alarm cue-exposed fish that lost a contest: they generally displayed lower aggression than winners but also lower aggression than losers of the control treatment. Thus, perceived predator presence modulates intra-sexual competition behaviour by increasing the costs associated with fighting.

Exposure to predation risk can induce a fearful baseline state, as well as fear reactions toward novel situations (i.e., neophobia). Some research indicates that risk exposure during sensitive periods makes adults more prone to acquiring long-term fearful phenotypes. However, chronic risk can also lead to ignoring threats in order to maintain other activities. We sought to assess how a relatively long period of low risk, experienced either early in life or by the previous generation, influences fear behaviour acquired from a short period of high risk as adults. We used fathead minnows as study subjects and simulated predation risk with repeated exposures to conspecific chemical alarm cues. The period of high risk experienced by adults induced typical fear behaviour (baseline freezing and neophobia), whereas the early-life low-risk period 1 year prior caused only a reduction in baseline foraging. We found no evidence that the early-life risk significantly altered the fear acquired from the adult-risk period. However, in a second experiment, a low-risk period during the parental generation interacted with a high-risk period experienced by the adult offspring. The combination of both risk periods heightened baseline freezing despite parental risk having little effect independently. Hence, our study provides evidence that parental risk exposure can lead to an additive intergenerational effect on fear acquisition in minnows.

Although comprehending the significance of phenotypic plasticity for evolution is of major interest in biology, the pre-requirement for that, the understanding of variance in plasticity, is still in its infancy. Most researchers assess plastic traits at single developmental stages and pool results between sexes. Here, we study variation among sexes and developmental stages in inducible morphological defences, a well-known instance of plasticity. We raised fathead minnows, Pimephales promelas, under different levels of background predation risk (conspecific alarm cues or distilled water) in a split-clutch design and studied morphology in both juveniles and adults. In accordance with the theory that plasticity varies across ontogeny and sexes, geometric morphometry analyses revealed significant shape differences between treatments that varied across developmental stages and sexes. Alarm cue-exposed juveniles and adult males developed deeper heads, deeper bodies, longer dorsal fin bases, shorter caudal peduncles and shorter caudal fins. Adult alarm cue-exposed males additionally developed a larger relative eye size. These responses represent putative adaptive plasticity as they are linked to reduced predation risk. Perhaps most surprisingly, we found no evidence for inducible morphological defences in females. Understanding whether similar variation occurs in other taxa and their environments is crucial for modelling evolution.

Bisphenol S (BPS), considered to be a safe alternative to Bisphenol A, is increasingly used in a wide variety of consumer and industrial products. However, mounting evidence suggests that BPS can act as a xenoestrogen targeting a wide range of neuro-endocrine functions in animals. At present, very little is known about the impacts of BPS on social behaviors and/or the potential underlying mechanisms. To this end, we exposed adult male and female zebrafish to environmentally relevant concentrations of BPS (0 (control), 1, 10, and 30 μg/L), as well as to 17β-estradiol (E2; 1 μg/L; as positive control) for 75 days. Subsequently, alterations in social behaviors were evaluated by measuring shoal cohesion, group preferences, and locomotor activity. Furthermore, to elucidate the possible molecular mechanism underlying the neuro-behavioral effects of BPS, we also quantified the changes in the mRNA abundance of arginine vasotocin (AVT), isotocin (IT), and their corresponding receptors in the zebrafish brain. The results showed that E2 and BPS (30 μg/L) decreased shoal cohesion in both males and females. Moreover, a marked decline in group preferences was observed in all treatment groups, while locomotor activity remained unaffected. Alterations in the social behaviors were associated with sex-specific changes in the mRNA expression of genes involved in IT and AVT signaling. Taken together, the results of this study suggest that chronic exposure to BPS can impair zebrafish social behaviors via disruption of isotocinergic and vasotocinergic neuro-endocrine systems.

In the face of anthropogenic change, we require a better understanding of how adaptive behavioural changes emerge from the interaction between personality and phenotypic plasticity, to be able to predict population persistence. Predation is an important context where this interaction occurs. Sampling wild fish populations revealed that boldness is size dependent in habitats with low predation risk but not in high predation habitats. This is predicted to occur due to a trade-off between a prey's metabolic demand, as per the metabolic hypothesis (smaller fish have higher metabolic requirements and thus need to be bolder) and their risk of predation, as per the predation risk hypothesis (smaller fish are subject to higher predation risk and should thus be shyer). However, since selective predation on specific individuals, age effects or the extent of direct experience with predators were not controlled for in these studies, the role of body size in modulating boldness remains unclear. Here, we tested the relationship between body size, boldness and predation risk in a widespread North American prey fish, the fathead minnow, Pimephales promelas. From hatching onwards, we manipulated perceived risk by exposing minnows to either conspecific alarm cues (indicative of predation risk) or distilled water in a split-clutch design. When fish were 4 months old, we measured body size and determined boldness using an emergence assay. Different levels of perceived predation risk did not directly affect body size or general boldness in fathead minnows. However, as predicted by the metabolic and predation risk hypotheses, high perceived predation risk cancelled out the negative correlation between boldness and body size present in controls through phenotypic plasticity. This effect was independent of selective predation, direct experience with predators and age. By adjusting personality through phenotypic plasticity, perceived predation risk alone may thus be sufficient for individuals to maximize their fitness in high-risk habitats.

To cope with the heterogeneous nature of predation and the trade-off between predator avoidance and foraging, prey animals have evolved several cognitive rules. One of these is the risk allocation hypothesis, which predicts that in environments with long periods of sustained high risk, individuals should decrease their antipredator effort to satisfy their metabolic requirements. The neophobia hypothesis, in turn, predicts increased avoidance of novel cues in high-risk habitats. Despite the recent interest in predator-induced neophobia across different sensory channels, tests of such generalized neophobia are restricted to a single fish taxon, the Cichlidae. Hence, we retested the generalized neophobia hypothesis in fathead minnows Pimephales promelas, a small schooling North American cyprinid fish. From hatching onward, minnows were exposed to conspecific alarm cues, which indicate predation risk, or distilled water in a split-clutch design. After 1 month, shoaling behavior was examined prior and subsequent to a mechanical predator disturbance. Fish previously exposed to elevated background risk formed compact shoals for a shorter time interval after the stimulus compared with controls. These results contrast previous studies of generalized neophobia but match the risk allocation hypothesis. Consequently, risk allocation and generalized neophobia are not ubiquitous cognitive rules but instead evolved adaptations of different taxa to their respective environments.

Predicting patterns of variation in sexual traits requires understanding how developmental environments influence mate choice. Here, we studied how long-term perceived high predation risk affects mutual mate choice in the cichlid Pelvicachromis taeniatus. From hatching onwards, fish were exposed regularly to either conspecific alarm cues signalling high risk or a control treatment consisting of exposure to water. Alarm cue-exposed and control adult fish were then offered the dichotomous choice between opposite-sex fish from different treatment groups. Under chronic predation risk, lower investment in mate choice and assortative preferences for mates from the same environment may be optimal. As predicted, alarm cue-exposed males assessed potential mates less than controls whereas females did not. Mate preferences differed as well but were independent of sex. While alarm cue-exposed fish were unselective, against expectations, control fish displayed disassortative preferences for alarm cue-exposed fish. This indicates that perceived high predation risk during development may contribute to maintaining variation in mating preferences and sexual traits.

The ability of organisms to plastically respond to changing environments is well studied. However, variation in phenotypic plasticity during ontogeny is less well understood despite its relevance of being an important source of phenotypic variation in nature. Here, we comprehensively study ontogenetic variation in morphological antipredator plasticity across multiple traits in Pelvicachromis taeniatus, a western African cichlid fish with sexually dimorphic ornamentation. In a split-clutch design, fish were raised under different levels of perceived predation risk (conspecific alarm cues or distilled water). Morphological plasticity varied substantially across ontogeny: it was first observable at an early juvenile stage where alarm cue-exposed fish grew faster. Subsequently, significant plasticity was absent until the onset of sexual maturity. Here, alarm cue-exposed males were larger than control males, which led to deeper bodies, longer dorsal spines, larger caudal peduncles, and increased eye diameters. Sexual ornamentation emerged delayed in alarm cue-exposed males. In later adulthood, the plastic responses receded. Despite small effect sizes, these responses represent putative adaptive plasticity, as they are likely to reduce predation risk. In females, we did not observe any plasticity. In accordance with theory, these results suggest fine-tuned expression of plasticity that potentially increases defenses during vulnerable developmental stages and reproductive output.

Many animal species are able to perceive light wavelengths beyond those visible to humans. While numerous species are additionally sensitive to short wavelengths (UV), long wavelengths such as the near-infrared spectrum (NIR) are supposed to be unsuitable for visual perception. Here, we experimentally show that under exclusive NIR illumination, the cichlid fish Pelvicachromis taeniatus displays a clear foraging response towards NIR reflecting prey. Additional control experiments without prey indicate that the observed behavior is not a mere response to the NIR environment. These results give first evidence for NIR visual sensitivity in a functional context and thus challenge the current view about NIR perception.

Mating preferences for genetic compatibility strictly depend on the interplay of the genotypes of potential partners and are therein fundamentally different from directional preferences for ornamental secondary sexual traits. Thus, the most compatible partner is on average not the one with most pronounced ornaments and vice versa . Hence, mating preferences may often conflict. Here, we present a solution to this problem while investigating the interplay of mating preferences for relatedness (a compatibility criterion) and large body size (an ornamental or quality trait). In previous experiments, both sexes of Pelvicachromis taeniatus , a cichlid fish with mutual mate choice, showed preferences for kin and large partners when these criteria were tested separately. In the present study, test fish were given a conflicting choice between two potential mating partners differing in relatedness as well as in body size in such a way that preferences for both criteria could not simultaneously be satisfied. We show that a sex-specific trade-off occurs between mating preferences for body size and relatedness. For females, relatedness gained greater importance than body size, whereas the opposite was true for males. We discuss the potential role of the interplay between mating preferences for relatedness and body size for the evolution of inbreeding preference.

A 76-year old patient was referred to our Oncology Unit due to hepatosplenomegaly, anemia, thrombocytopenia, eosinophilia, elevation of lactat dehydrogenase and leucocytosis with occurrence of myeloid and erythroid precursors in the peripheral blood. Histopathological examination of the bone marrow and molecular genetics showing a c-kit-D816V-mutation confirmed the diagnosis of systemic mastocytosis without evidence of idiopathic myelofibrosis. After starting of prednisone treatment anemia, thromocytopenia and the patient's performance status rapidly improved.

Fast acclimatisation of experimental animals to laboratory test conditions is crucial for effective research. During highly standardized experiments, confounding parameters must be mini- mized which might lead to animals behaving unnaturally. The objective of the present study was to develop a simple method to enhance activity of fishes under laboratory test conditions. Many fishes intensively interact with substrate in their natural environment; however, providing substrate during experiments might impair observation or may otherwise confound the results. Therefore, the behav- iour of the cichlid Pelvicachromis taeniatus was examined in water that was exposed to substrate only prior to the experiment. As a control, fish behaviour in tap water was investigated. Fish activity (i.e. covered distance) was evaluated during a period of 2 h. In substrate-treated water fish were signifi- cantly more active than in tap water. Furthermore, males in the substrate-treated water increased their activity over time, whereas males in tap water did not. In contrast, activity of females did not change significantly over time. No significant relationship between fish activity and body size or condition was found. Furthermore, standard chemical water parameters were not altered by the substrate treatment. Thus, substrate is suggested as a means to improve the olfactory environment, promoting fish welfare and activity. In conclusion, this method is suggested as a standard with wide applicability across benthic fish species to improve acclimatisation of fishes to experimental conditions in the laboratory.

Theory predicts that parents should adjust their parental investment to the reproductive value of the brood. Previous studies have mainly investigated the relationship between brood size, brood age and the intensity of care. However, the impact of brood quality traits such as the offspring’s average body size on parental investment has received relatively little attention. We examined the parental response to brood quality in the biparentally brood-caring cichlid fish Pelvicachromis taeniatus. We manipulated offspring quality by manipulating the food availability for the free-swimming fry and measured the parental response over 4 weeks. Generally, care decreased over the 4 weeks suggesting that parents adjust care to decreasing offspring vulnerability. However, parents of relatively low-quality broods, that is, of broods with relatively small average individual body size, showed a greater reduction in care than parents of high-quality broods resulting in a significant difference in care provided in the fourth week. The result suggests that parents adjust their care to brood quality and supports the predictions of the parental investment theory. Furthermore, mothers of high-quality fry were significantly more aggressive towards their partner than mothers of low-quality fry. This result is discussed in the context of parental response to brood value. Generally, females invested more in brood care than males.

Individuals have to respond simultaneously to different environmental factors often making trade-offs between conflicting demands necessary. Many freshwater ecosystems are resource-limited and both intra- and interspecific competitiveness is a common requirement to gain and defend resources necessary for reproduction. Although predation risk is an important selective force affecting behavioral decisions, little is known about the impact of predation risk on interspecific competition. Here, we investigate whether chemically mediated predation risk affects interspecific territorial aggression by the freshwater cichlid Pelvicachromis taeniatus. In our experiments, territorial P. taeniatus males were visually confronted with a territorial intruder: a heterospecific, sympatric cichlid (Benitochromis nigrodorsalis) which generally induced aggression in P. taeniatus. Predation risk for P. taeniatus was simulated by a concurrent release of conspecific chemical alarm cues. In control treatments, no chemical cues, dissolved heterospecific alarm cues, or aliquots of distilled water were provided during these aggressive encounters. The results show that interspecific aggression of territorial male P. taeniatus is significantly decreased under predation risk compared to the control treatments. This suggests that interspecific competition becomes less intense under concurrent predation risk. As this process could hinder competitive exclusion, predation risk may indirectly promote and stabilize biodiversity in natural ecosystems.

The evolution of alarm signals has puzzled evolutionary ecologists for decades. This is particularly true for alarm cues (‘Schreckstoff’) which are present in many fishes. They are passively released through injuries and signal the presence of a predator. Although the benefits for conspecific receivers are obvious (they can adjust their behavior to avoid predation), those for the senders are not which is, however, a necessary requirement for the evolution of alarm signals. Several hypotheses relying on potential direct benefits for the senders have been suggested. Alarm cues might attract secondary predators which in turn might increase the escape probability of the sender. A primary immune enhancing role was suggested as well. An alternative explanation is based on Hamilton’s inclusive fitness theory stating that individuals can indirectly increase their fitness by increasing the survival of genetically related individuals (‘kin selection theory’). If related individuals preferentially benefit from alarm signals, for instance by being more receptive to kin-alarm cues, senders could increase their inclusive fitness. Here, we investigate whether individuals of the cichlid fish Pelvicachromis taeniatus respond differentially to alarm cues derived from kin and non-kin. P. taeniatus possesses alarm cues and is known to adjust its behavior when exposed to alarm cues. We measured the change in activity after the addition of alarm cues (derived from kin- and non-kin) relative to a control treatment. Reduced activity is a widespread behavioral adaptation to reduce predation risk in prey organisms. Fish of the alarm cue treatments significantly reduced their activity relative to control fish. However, fish did not respond differentially to alarm cues derived from kin and non-kin suggesting that potential inclusive fitness benefits are not mediated by responses specific to individual alarm cues. We discuss alternative mechanisms such as kin shoaling and mating preferences potentially leading to kin-biased alarm cue perception.