L. Fredrik Sundström

Review of Elasmobranch Behavioral Studies Using Ultrasonic Telemetry with Special Reference to the Lemon Shark, Negaprion Brevirostris, Around Bimini Islands, Bahamas

A review of past behavioral ultrasonic telemetry studies of sharks and rays is presented together with previously unpublished material on the behavior of the lemon shark, Negaprion brevirostris, around the Bimini Islands, Bahamas. The review, focusing on movement behaviors of 20 shark and three ray species, reveals that elasmobranchs exhibit a variety of temporal and spatial patterns in terms of rates-of-movement and vertical as well as horizontal migrations. The lack of an apparent pattern in a few species is probably attributable to the scarcity of tracking data. Movements are probably governed by several factors, some still not studied, but data show that food, water temperature, bottom type, and magnetic gradient play major roles in a sharks decision of where and when to swim. A few species exhibit differences in behavior between groups of sharks within the same geographical area. This interesting finding warrants further research to evaluate the causes of these apparent differences and whether these groups constitute different subpopulations of the same species. The lack of telemetry data on batoids and some orders of sharks must be addressed before we can gain a more comprehensive understanding of the behavior of elasmobranch fishes. Previously unpublished data from 47 smaller and 38 larger juvenile lemon sharks, collected over the decade 1988–1998, provide new results on movement patterns, habitat selection, activity rhythms, swimming speed, rate-of-movement, and homing behavior. From these results we conclude that the lemon shark is an active predator with a strong, apparently innate homing mechanism. This species shows ontogenetic differences in habitat selection and behavior, as well as differences in movements between groups of individuals within the same area. We suggest three hypotheses for future research on related topics that will help to understand the enigmatic behavior of sharks.

Effects of egg yolk testosterone on growth and immunity in a precocial bird

In oviparous vertebrates, maternal steroid allocation to eggs can have important fitness consequences for the offspring. However, elevated testosterone levels are not only associated with beneficial postnatal effects, such as enhanced growth and high social status, but may also entail costs by suppressing the immune system. In this study, testosterone levels in eggs of Chinese painted quail (Coturnix chinensis) were experimentally manipulated to evaluate its effects on growth and immunocompetence. Testosterone did not affect embryonic development, body size or growth during the first 20 days. However, elevated testosterone levels during embryonic development were immunosuppressive for chicks with inherently higher growth rate. Adaptive scenarios where only beneficial effects of increased testosterone levels are considered may therefore need to be re‐evaluated.

Boldness Predicts Social Status in Zebrafish (Danio rerio)

This study explored if boldness could be used to predict social status. First, boldness was assessed by monitoring individual zebrafish behaviour in (1) an unfamiliar barren environment with no shelter (open field), (2) the same environment when a roof was introduced as a shelter, and (3) when the roof was removed and an unfamiliar object (Lego® brick) was introduced. Next, after a resting period of minimum one week, social status of the fish was determined in a dyadic contest and dominant/subordinate individuals were determined as the winner/loser of two consecutive contests. Multivariate data analyses showed that males were bolder than females and that the behaviours expressed by the fish during the boldness tests could be used to predict which fish would later become dominant and subordinate in the ensuing dyadic contest. We conclude that bold behaviour is positively correlated to dominance in zebrafish and that boldness is not solely a consequence of social dominance.

GROWTH AND SURVIVAL TRADE‐OFFS AND OUTBREEDING DEPRESSION IN RAINBOW TROUT (ONCORHYNCHUS MYKISS)

Abstract The purpose of this study was to examine, using a rainbow trout (Oncorhynchus mykiss) model system, the fitness consequences of three generations of introgression of genotypes adapted to two different environments (culture and nature). The experiments also isolated the influence of competitive interactions and risk of predation on the relative growth and survival of the wild and backcrossed lines. Line crosses representing fast-growing pure domestic (D), slow-growing pure wild (W), domestic × wild hybrids (F1), F1 × wild backcrosses (B1), and B1 × wild backcrosses (B2) were generated and reared under (1) culture conditions, (2) seminatural conditions with competition among genotypes, and (3) seminatural conditions under risk of predation. Survival of the fry in a seminatural environment with competition fit an additive model of gene action with the domestic fish having the highest survival and the wild fish the lowest, but under risk of predation outbreeding depression was suggested by low survival of the B2 lines. Evidence of a trade-off in growth and survival under risk of predation along with observations of genetically determined behavioral differences among the strains may provide some explanation for the observed differences in survival among the strains. This information is relevant to improving our evolutionary understanding of the interaction among genomes, and the influence of environment, during hybridization events. Results from this experiment indicate that alteration of phenotype likely played a prominent role in the reduced fitness experienced by progeny produced after three generations of introgression, supporting the theory that disruption of genotypes selected for adaptation to local conditions may be a primary cause of outbreeding depression in species such as salmon.

Investment in territorial defence depends on rearing environment in brown trout (Salmo trutta)

In many animals, territoriality will arise or cease depending on environmental factors such as intruder rate and resource availability. We investigated the effect of rearing environment on territorial behaviour in ~1.5-month-old brown trout. In the laboratory, wild-caught (reared at a low density) and hatchery-reared (high density) trout were allowed to defend a territory against a size-matched intruder reared in the same or the other environment. Because territorial behaviour should be relaxed at high-rearing densities, we hypothesized that hatchery-reared trout should value their territories less and therefore invest less in defence compared with wild-caught trout. However, in all cases, territory owners were more likely to win the contest and hatchery-reared trout were just as likely as wild-reared to win mixed contests. Furthermore, pairs of hatchery-reared trout initiated contests sooner, fought longer and were more aggressive during the contest compared with pairs of wild trout. When hatchery-reared owners met wild intruders, the contest ended sooner compared with when the roles were reversed. We conclude that territorial behaviour in brown trout is largely innate, but that the hatchery environment has promoted more aggressive individuals. These results suggest that hatchery-reared trout invest more time and energy to obtain the same contest success as wild trout. In conclusion, the lack of experience of territorial defence in a high-density rearing environment seems to reduce the efficiency of territorial behaviour. In turn, this may have negative consequences for the performance of released hatchery fish in the wild.

Leptin depresses food intake in great tits (Parus major).

Food availability for wild organisms typically varies both in time and space, requiring a mechanism that regulates the storage of excess energy and makes it possible to use stores during energy shortfall. Leptin, a protein hormone encoded by an obesity gene, has been suggested to be the signal mediator for this flux of energy. In a controlled laboratory experiment on caged great tits (Parus major) we evaluated the effect of leptin on food intake and behaviour. Experimental birds were given an intramuscular injection of 10 microg leptin dissolved in phosphate buffered saline (PBS), while the control birds were injected with PBS only at 09:00 h after a nights fasting. Within the first 20 min after injections we observed a significant difference in food intake between groups: control birds initially fed at higher rates compared to leptin treated birds. The cumulative food intake suggested that the effect of leptin disappeared after approximately 40-50 min post-injections. Similar results have previously been found in domesticated chickens. To our knowledge, this is the first study to show that leptin depresses food intake in wild birds.

Using speed-sensing transmitters to construct a bioenergetics model for subadult lemon sharks, Negaprion brevirostris (Poey), in the field

Several methods to indirectly estimate metabolism of aquatic animals have been reported, including heart rate, electromyograms, video recording, and averaging velocity of an animal moving between two or more points. The present study carried out in the lagoon at Bimini Islands, Bahamas, used acoustic, speed-sensing transmitters to indirectly estimate energy consumption of 1.5–2 m subadult lemon sharks (Negaprion brevirostris). Speed records from three sharks tracked a total of 170 h, yielded average swimming speeds of 0.44–0.71 m s-1. These speeds were converted into energy consumption to obtain metabolism.

Physiological constraints to climate warming in fish follow principles of plastic floors and concrete ceilings

Understanding the resilience of aquatic ectothermic animals to climate warming has been hindered by the absence of experimental systems experiencing warming across relevant timescales (for example, decades). Here, we examine European perch (Perca fluviatilis, L.) from the Biotest enclosure, a unique coastal ecosystem that maintains natural thermal fluctuations but has been warmed by 5–10 °C by a nuclear power plant for over three decades. We show that Biotest perch grow faster and display thermally compensated resting cardiorespiratory functions compared with reference perch living at natural temperatures in adjacent waters. However, maximum cardiorespiratory capacities and heat tolerance limits exhibit limited or no thermal compensation when compared with acutely heated reference perch. We propose that while basal energy requirements and resting cardiorespiratory functions (floors) are thermally plastic, maximum capacities and upper critical heat limits (ceilings) are much less flexible and thus will limit the adaptive capacity of fishes in a warming climate.

Predation risk and the territory value of cover: an experimental study

In an experiment on territorial brown trout (Salmo trutta), we addressed the novel hypothesis that protective cover increases the value of a territory in relation to the perceived level of predation risk. We predicted that territory holders should invest more resources defending territories with cover than territories without cover, and that defence should increase as predation risk increases. First, trout were allowed to establish ownership in territories with or without overhead cover. Second, predation risk was manipulated by simulating aerial predator attacks in half of the territories of each type, whereupon the preference for cover was estimated. Third, owners of the four types of territories were staged in dyadic contests against size-matched intruders. Territory owners showed a preference for cover, which increased further after simulated predator attacks. In subsequent contests, conflicts over territories with cover were settled faster than conflicts over territories without cover, which may suggest that the value of cover increases the motivational asymmetry between owner and intruder. Consistent with our hypothesis, owners of territories with cover were much more aggressive if they had been subjected to predator attacks the day before the conflict. These results suggest that territory owners are able to estimate the value of protective cover in response to variation in the level of predation risk in the habitat.

Leptin and social environment influence the risk-taking and feeding behaviour of Asian blue quail

Optimality models predict that hungry animals should be more willing to take risks than satiated individuals; thus, the decision whether to forage is influenced by the physiological process mediating the sensing of energy reserves. An important substance involved in this sensing process is leptin, a protein hormone known to down-regulate appetite and increase energy expenditure. However, feeding and risk-taking behaviours are also influenced by the behaviour of surrounding individuals. By increasing leptin levels in a focal bird and/or in its groupmates, we studied the combined effects of physiological and social factors on the risk-taking and feeding behaviour of Asian blue quail, Coturnix chinensis. Risk-taking behaviour, estimated by the time taken to start feeding, was dependent on social factors and a birds own weight but not on whether it was treated with leptin; focal birds with leptin-treated companions took longer before feeding than did focal birds with control companions. Total feeding time, on the other hand, was physiologically governed with no effect of the companions; leptin-treated focal birds spent less time feeding than did control focal birds. To our knowledge, this study is the first to investigate the interaction between leptin and social factors in animal behaviour.