Aaron J. Zolderdo

Stress, nutrition and parental care in a teleost fish: Exploring mechanisms with supplemental feeding and cortisol manipulation

ABSTRACT Parental care is an essential life-history component of reproduction for many animal species, and it entails a suite of behavioural and physiological investments to enhance offspring survival. These investments can incur costs to the parent, reducing their energetic and physiological condition, future reproductive capabilities and survival. In fishes, relatively few studies have focused on how these physiological costs are mediated. Male smallmouth bass provide parental care for developing offspring until the brood reaches independence. During this energetically demanding life stage, males cease active foraging as they vigorously defend their offspring. Experimental manipulation of cortisol levels (via implantation) and food (via supplemental feeding) in parental males was used to investigate the fitness consequences of parental care. Improving the nutritional condition of nest-guarding males increased their reproductive success by reducing premature nest abandonment. However, supplemental feeding and cortisol treatment had no effect on parental care behaviours. Cortisol treatment reduced plasma lymphocyte numbers, but increased neutrophil and monocyte concentrations, indicating a shift in immune function. Supplemental feeding improved the physiological condition of parental fish by reducing the accumulation of oxidative injury. Specifically, supplemental feeding reduced the formation of 8-hydroxy-2′-deoxyguanosine (8-OHdG) on DNA nucleotides. Increasing the nutritional condition of parental fish can reduce the physiological cost associated with intensive parental activity and improve overall reproductive success, illustrating the importance of nutritional condition as a key modulator of parental fitness. Summary: Stress and nutritional condition are key modulators of physiological condition and reproductive success in a parental teleost fish.

Parental Care in a Stressful World: Experimentally Elevated Cortisol and Brood Size Manipulation Influence Nest Success Probability and Nest-Tending Behavior in a Wild Teleost Fish*

Parental care is an advantageous reproductive behavior, as the fitness of the caregiver is increased through improving the chances of its offspring’s survival. Parental care occurs in a variety of teleost fishes. The body size of parental fish and the size of their brood can affect nest abandonment decisions, where compared with smaller fish with smaller broods, larger fish with larger broods typically invest more energy into reproductive events because they have less future reproductive potential. Although essential for basal metabolism and body maintenance functions, when glucocorticoid hormones (e.g., cortisol) are chronically elevated, as can occur during stress, fish may experience impairments in behavior and immune function, leading to compromised health and condition. Anthropogenic stressors during parental care can lead to elevated stress, therefore making it necessary to understand how stress influences an already-challenging period. Using smallmouth bass as a model, a gradient of body sizes, and experimentally manipulated brood size (i.e., reducing large broods and supplementing small broods) and cortisol levels (i.e., elevated via slow-release intraperitoneal cocoa butter implants containing cortisol versus controls), we tested the hypothesis that the reproductive success and parental care behaviors (i.e., aggression, nest tending) of nest-guarding male smallmouth bass are influenced by parental body size, brood size, and cortisol level. Overall, there was a relationship between cortisol treatment and nest success in which larger fish exhibited lower success when cortisol levels were elevated. Brood size had a significant effect on fish-tending behavior, independent of cortisol level and body size. Lending partial support to our hypothesis, the results of this study indicate that the reproductive success of guarding male smallmouth bass is influenced by cortisol level and that tending behavior is affected by brood size.

Consequences of Different Types of Littoral Zone Light Pollution on the Parental Care Behaviour of a Freshwater Teleost Fish

Ecological light pollution occurs when artificial lights disrupt the natural regimes of individual organisms or their ecosystems. Increasing development of shoreline habitats leads to increased light pollution (e.g., from cottages, docks, automobile traffic), which could impact the ecology of littoral zones of lakes and rivers. Smallmouth bass (Micropterus dolomieu) engage in sole paternal care, guarding their nest continually, day and night, to protect their developing offspring. Any alterations to their behaviour—either directly because of the response to light or indirectly due to changes in nest predator activity and associated response of the bass—could lead to increased energetic demands for fish that have a fixed energy budget and ultimately reduce reproductive success. To examine this issue, tri-axial accelerometer biologgers were externally attached to nesting smallmouth bass during the egg stage to determine whether light pollution (i.e., dock lights with low levels of continuous light and spotlights with high intensity irregular light simulating automobile traffic) altered behaviour of nesting males relative to control fish. Our study revealed that both types of light pollution increased overall bass activity level compared with the control group. The intermittent light treatment group had the highest activity and exhibited large fluctuations between night and day activity levels. Fish in the continual light treatment group displayed statistically higher activity than the control fish but showed limited fluctuations between day and night activity levels. Our results suggest that continuous or intermittent light sources, common in shoreline habitats that have been developed, have the potential to alter the behaviour and thus energy use of nest-guarding fish. This study contributes to the growing body of literature on the ecological consequences of light pollution in aquatic ecosystems.

An experimental test of in-season homing mechanisms used by nest-guarding male Largemouth Bass following displacement.

Through manipulations of sensory functions, researchers have evaluated the various mechanisms by which migratory fish, particularly in lotic systems, locate natal spawning grounds. Comparatively less work has occurred on the ways by which fish in lentic systems locate spawning sites, and more specifically, the ways by which displaced fish in these systems locate their broods post spawning. The primary goal of this research was to determine the sensory mechanisms used by nesting, male Largemouth Bass to navigate back to their brood following displacement. This was accomplished by comparing the ability of visually impaired, olfactory impaired and geomagnetically impaired individuals to return to their nests after a 200 m displacement, relative to control males. All treatments were designed to be temporary and harmless. We analyzed the data using a generalized linear mixed model, and found that the probability of an olfactory impaired individual returning to his nest within a given time interval was significantly lower than the probability of a geomagnetically impaired individual returning. Overall, it appears as though olfaction is the most important sensory mechanism used for homing in Largemouth Bass.

Cortisol treatment affects locomotor activity and swimming behaviour of male smallmouth bass engaged in paternal care: A field study using acceleration biologgers

Paternal care, where the male provides sole care for the developing brood, is a common form of reproductive investment among teleost fish and ubiquitous in the Centrarchidae family. Throughout the parental care period, nesting males expend energy in a variety of swimming behaviours, including routine and burst swimming, vigilantly monitoring the nest area and protecting the brood from predators. Parental care is an energetically demanding period, which is presumably made even more difficult if fish are exposed to additional challenges such as those arising from human disturbance, resulting in activation of the hypothalamic-pituitary-interrenal axis (i.e., elevation of cortisol). To study this situation, we examined the effects of experimental manipulation of the stress hormone cortisol on locomotor activity and behaviour of nest guarding male smallmouth bass (Micropterus dolomieu). We exogenously elevated circulating cortisol levels (via intracoelomic implants) and attached tri-axial accelerometers to wild smallmouth bass for three days. During the recovery period (i.e., ≤4h post-release), cortisol-treated fish exhibited significantly reduced locomotor activity and performed significantly less burst and routine swimming relative to control fish, indicating cortisol uptake was rapid, as were the associated behavioural responses. Post-recovery (i.e., >4h post-release), fish with high cortisol exhibited lower locomotor activity and reduced routine swimming relative to controls. Fish were less active and reduced routine and burst swimming at night compared to daylight hours, an effect independent of cortisol treatment. Collectively, our results suggest that cortisol treatment (as a proxy for anthropogenic disturbance and stress) contributed to altered behaviour, and consequently cortisol-treated males decreased parental investment in their brood, which could have potential fitness implications.