Individual Variation In Behaviour Research Articles

Downstream passage performance of silver eel at an angled rack: effects of behavior and morphology

AbstractThe European eel is critically endangered due to heavy impact of anthropogenic factors, such as habitat fragmentation, overexploitation and climate change. During downstream migration, silver eels may encounter hydropower plants, which often result in delay or mortality from impingement on trash-racks or turbine passage. These problems can be mitigated with downstream passage solutions, such as angled racks that guide downstream-migrating eels to safe passage routes. The importance of bar spacing and phenotypic diversity for passage performance is, however, largely unknown. In this study, we investigated how morphological parameters (body mass, eye and fin indices) and behavioral score (open field test) influenced passage rate at an experimental intake equipped with a bypass and angled racks with either 15 or 30 mm bar spacing. Both racks were efficient in guiding eels into a bypass. There was a strong positive effect of body mass and a weak positive effect of open field test score on passage rate. Other factors such as eye and fin indices played a minor role. These results demonstrate the performance of angled racks with bypasses and form a useful starting point for further research regarding the relationships between individual variation in behavior, morphology and passage solutions for silver eels.

Warming, but not infection with Borrelia burgdorferi, increases off-host winter activity in the ectoparasite, Ixodes scapularis

Warming winters will change patterns of behaviour in temperate and polar arthropods, but we know little about the drivers of winter activity in animals such as ticks. Any changes in behaviour are likely to arise from a combination of both abiotic (e.g. temperature) and biotic (e.g. infection) drivers, and will have important consequences for survival and species interactions. Blacklegged ticks, Ixodes scapularis, have invaded Atlantic Canada and high proportions (30–50%) are infected with the bacteria causing Lyme disease, Borrelia burgdorferi. Infection is correlated with increased overwintering survival of adult females, and ticks are increasingly active in the winter, but it is unclear if infection is associated with activity. Further, we know little about how temperature drives the frequency of winter activity. Here, we exposed wild-caught, adult, female Ixodes scapularis ticks to three different winter temperature regimes (constant low temperatures, increased warming, and increased warming + variability) to determine the thermal and infection conditions that promote or suppress activity. We used automated behaviour monitors to track daily activity in individual ticks and repeated the study with fresh ticks over three years. Following exposure to winter conditions we determined whether ticks were infected with the bacteria B. burgdorferi and if infection was responsible for any patterns in winter activity. Warming conditions promoted increased activity throughout the overwintering period but infection with B. burgdorferi had no impact on the frequency or overall number of ticks active throughout the winter. Individual ticks varied in their levels of activity throughout the winter, such that some were largely dormant for several weeks, while others were active almost daily; however, we do not yet know the drivers behind this individual variation in behaviour. Overall, warming winters will heighten the risk of tick-host encounters.

Unraveling the hierarchical structure of posture and muscle activity changes during mating of Caenorhabditis elegans.

One goal of neurobiology is to explain how decision-making in neuromuscular circuits produces behaviors. However, two obstacles complicate such efforts: individual behavioral variability and the challenge of simultaneously assessing multiple neuronal activities during behavior. Here, we circumvent these obstacles by analyzing whole animal behavior from a library of Caenorhabditis elegans male mating recordings. The copulating males express the GCaMP calcium sensor in the muscles, allowing simultaneous recording of posture and muscle activities. Our library contains wild type and males with selective neuronal desensitization in serotonergic neurons, which include male-specific posterior cord motor/interneurons and sensory ray neurons that modulate mating behavior. Incorporating deep learning-enabled computer vision, we developed a software to automatically quantify posture and muscle activities. By modeling, the posture and muscle activity data are classified into stereotyped modules, with the behaviors represented by serial executions and transitions among the modules. Detailed analysis of the modules reveals previously unidentified subtypes of the male's copulatory spicule prodding behavior. We find that wild-type and serotonergic neurons-suppressed males had different usage preferences for those module subtypes, highlighting the requirement of serotonergic neurons in the coordinated function of some muscles. In the structure of the behavior, bi-module repeats coincide with most of the previously described copulation steps, suggesting a recursive "repeat until success/give up" program is used for each step during mating. On the other hand, the transition orders of the bi-module repeats reveal the sub-behavioral hierarchy males employ to locate and inseminate hermaphrodites.

Understanding the differential impacts of two antidepressants on locomotion of freshwater snails (Lymnaea stagnalis)

There is growing evidence of negative impacts of antidepressants on behavior of aquatic non-target organisms. Accurate environmental risk assessment requires an understanding of whether antidepressants with similar modes of action have consistent negative impacts. Here, we tested the effect of acute exposure to two antidepressants, fluoxetine and venlafaxine (0–50 µg/L), on the behavior of non-target organism, i.e., freshwater pond snail, Lymnaea stagnalis. As compounds interact with chemical cues in the aquatic ecosystems, we also tested whether the effects altered in the presence of bile extract containing 5α-cyprinol sulfate (5α-CPS), a characterized kairomone of a natural predator, common carp (Cyprinus carpio). Behavior was studied using automated tracking and analysis of various locomotion parameters of L. stagnalis. Our results suggest that there are differences in the effects on locomotion upon exposure to venlafaxine and fluoxetine. We found strong evidence for a non-monotonic dose response on venlafaxine exposure, whereas fluoxetine only showed weak evidence of altered locomotion for a specific concentration. Combined exposure to compounds and 5α-CPS reduced the intensity of effects observed in the absence of 5α-CPS, possibly due to reduced bioavailability of the compounds. The results highlight the need for acknowledging different mechanisms of action among antidepressants while investigating their environmental risks. In addition, our results underline the importance of reporting non-significant effects and acknowledging individual variation in behavior for environmental risk assessment.Graphical

Insight into differing decision-making strategies that underlie cognitively effort-based decision making using computational modeling in rats.

The rat cognitive effort task (rCET), a rodent model of cognitive rather than physical effort, requires animals to choose between an easy or hard visuospatial discrimination, with a correct hard choice more highly rewarded. Like in humans, there is stable individual variation in choice behavior. In previous reports, animals were divided into two groups-workers and slackers-based on their mean preference for the harder option. Although these groups differed in their response to pharmacological challenges, the rationale for using this criterion for grouping was not robust. We collated experimental data from multiple cohorts of male and female rats performing the rCET and used a model-based framework combining drift diffusion modeling with cluster analysis to identify the decision-making processes underlying variation in choice behavior. We verified that workers and slackers are statistically different groups but also found distinct intra-group profiles. These subgroups exhibited dissociable performance during the attentional phase, linked to distinct decision-making profiles during choice. Reanalysis of previous pharmacology data using this model-based framework showed that serotonergic drug effects were explained by changes in decision boundaries and non-decision times, while scopolamine's effects were driven by changes in decision starting points and rates of evidence accumulation. Modeling revealed the decision-making processes that are associated with cognitive effort costs, and how these differ across individuals. Reanalysis of drug data provided insight into the mechanisms through which different neurotransmitter systems impact cognitively effortful attention and decision-making processes, with relevance to multiple psychiatric disorders.

Spatiotemporal-social association predicts immunological similarity in rewilded mice.

Environmental influences on immune phenotypes are well-documented, but our understanding of which elements of the environment affect immune systems, and how, remains vague. Behaviors, including socializing with others, are central to an individual's interaction with its environment. We therefore tracked behavior of rewilded laboratory mice of three inbred strains in outdoor enclosures and examined contributions of behavior, including associations measured from spatiotemporal co-occurrences, to immune phenotypes. We found extensive variation in individual and social behavior among and within mouse strains upon rewilding. In addition, we found that the more associated two individuals were, the more similar their immune phenotypes were. Spatiotemporal association was particularly predictive of similar memory T and B cell profiles and was more influential than sibling relationships or shared infection status. These results highlight the importance of shared spatiotemporal activity patterns and/or social networks for immune phenotype and suggest potential immunological correlates of social life.

Behavioral syndromes in paper wasps: Links between social and non-social personality in Polistes fuscatus

Abstract Although much work has focused on non-social personality traits such as activity, exploration, and neophobia, there is a growing appreciation that social personality traits play an important role in group dynamics, disease transmission, and fitness and that social personality traits may be linked to non-social personality traits. These relationships are important because behavioral syndromes, defined here as correlated behavioral phenotypes, can constrain evolutionary responses. However, the strength and direction of relationships between social and non-social personality traits remain unclear. In this project, we examine social and non-social personality traits, and the relationships between them, in the paper wasp Polistes fuscatus. With a novel assay, we identify 5 personality traits, 2 non-social (exploration and activity), and 3 social (aggression, affiliation, and antennation) personality traits. We also find that social and non-social personality traits are phenotypically linked. We find a positive correlation between aggression and activity and a negative correlation between affiliation and activity. We also find a positive correlation between exploration and activity. Our work is an important step in understanding how phenotypic linkage between social and non-social behaviors may influence behavioral evolution. As a burgeoning model system for the study of genetic and neurobiological mechanisms of social behavior, Polistes fuscatus has the potential to add to this work by exploring the causes and consequences of individual behavioral variation.

Foraging fidelity and individual specialisation in a temperate bat Myotis nattereri

This study explores whether Natterer’s bats exhibit fidelity to their foraging sites and individual specialisation in their use of habitat types. The research tracked 34 individual bats in two different landscapes, with some bats tracked twice over varying time intervals. The study found that Natterer’s bats show consistency in their fidelity to foraging sites across different landscapes and intervals between observations. The bats repeatedly exploited specific foraging areas, and some showed individual specialisation in their habitat use. The study also found considerable variation in individual behaviour. The research suggests that a single full night of radio-tracking data can serve as a useful proxy for longer periods and contribute to more robust descriptions of resource requirements by Natterer’s bats in their breeding seasons. This study emphasises the importance of maximising the number of individuals from which movement data is obtained to inform conservation and management of temperate bats, whilst ensuring that the data represent a meaningful measure of behaviour. The results also suggest that protecting a mosaic of habitats rather than a single habitat may be important for Natterer’s bats, given their individual specialisation in habitat use.

Examining individual behavioural variation in wild adult bull sharks (Carcharhinus leucas) suggests divergent personalities

Abstract Although many animal species demonstrate individual personalities, studying these traits in wild sharks has proven challenging. Past research focused mainly on captive or juvenile sharks. Our ethological study of 31 wild adult bull sharks (Carcharhinus leucas) utilized an artificial provisioning site, amassing 2813 observations over 651 dives from October 2015 to January 2018 (27 months). Behavioural traits, including boldness-shyness and aggressiveness-placidity, were assessed using an ad hoc ethogram and an influencing factors table. This innovative approach not only allowed us to characterize individual shark behaviours but also to quantify their changes over time. Our findings suggest that adult bull sharks likely possess distinct personalities, spanning from extreme shyness to pronounced boldness, with varying levels of plasticity among individuals. Further exploration of shark personalities holds promise for advancing our comprehension of human–shark interactions and refining the management of potential aggressive behaviours exhibited by large shark species toward humans.

Explorative behavior allows the successful finding of ephemeral food resources in the wild

AbstractThe repeatable among-individual variation in behavior (animal personality) is considered to affect fitness. The variation in personality traits is shaped by complex evolutionary mechanisms involving energy allocation and resource acquisition. However, an association between personality and individual performance in energy gain in the natural environment still seems underexplored. The aim of our study was to test a hypothesis that consistent individual variation in behavior affects the energy acquisition of unpredictable food resources. We predicted that more explorative individuals would be finding ephemeral food resources with higher probability than those less explorative. The quasi-experiment with ‘random-walking-feeders’ was performed to assess the use of unpredictable food resources in the natural habitat occupied by free-ranging arboreal rodents — edible dormice Glis glis. A repeated open field test at laboratory conditions was conducted to assess whether male dormice show consistent among-individual variation in behavior. The open field test analysis indicated two main components of dormice behavior: exploration and boldness, from which only the first one was repeatable and thus considered as a component of animal personality. The probability of finding a feeder increased with exploration, independently of dormice age and body size. The results indicate that exploration can be considered an adaptive trait that improves energy acquisition. We suggest that the behavioral among-individual variation within populations persists due to the usage of distinct foraging strategies in the face of fluctuations in food resources availability.

Dolphin social phenotypes vary in response to food availability but not the North Atlantic Oscillation index.

Social behaviours can allow individuals to flexibly respond to environmental change, potentially buffering adverse effects. However, individuals may respond differently to the same environmental stimulus, complicating predictions for population-level response to environmental change. Here, we show that bottlenose dolphins (Tursiops truncatus) alter their social behaviour at yearly and monthly scales in response to a proxy for food availability (salmon abundance) but do not respond to variation in a proxy for climate (the North Atlantic Oscillation index). There was also individual variation in plasticity for gregariousness and connectedness to distant parts of the social network, although these traits showed limited repeatability. By contrast, individuals showed consistent differences in clustering with their immediate social environment at the yearly scale but no individual variation in plasticity for this trait at either timescale. These results indicate that social behaviour in free-ranging cetaceans can be highly resource dependent with individuals increasing their connectedness over short timescales but possibly reducing their wider range of connection at longer timescales. Some social traits showed more individual variation in plasticity or mean behaviour than others, highlighting how predictions for the responses of populations to environmental variation must consider the type of individual variation present in the population.

Exploratory Study on Individual Locomotor Activity in Local Dual-Purpose and Commercial Breeder Pullets.

Improving animal welfare is a prerequisite for the societal acceptance of poultry production. Support for improvements requires practical tools to quantify animal welfare and identify predispositions at the individual level, where possible. In this study, the activities and behavior of dual-purpose chickens (N = 245) and commercial breeders (N = 224) were analyzed. The general locomotor activity (GLA) data were collected using an RFID system over five days with 9-to-14-week-old animals. The results show that the animals of comparable age and stocking density differed from each other in their activity (p ≤ 0.001) according to breed, but no sex differences were observed (p = 0.159). No correlations were found between GLA and plumage condition (p > 0.05). The individual variations within the breeds are presented and discussed on an animal-by-animal level, providing new insights into the individual behavioral variability of chickens. The RFID systems can reliably generate GLA data that help to understand the potential interplay between behavior and animal welfare. The technology is also suitable for creating individual (personality) profiles that can be used for breeding. With a better understanding of the role of activity, husbandry and management practices can be adapted to improve animal welfare.

Graduate Student Literature Review: Sociability, fearfulness, and coping style—Impacts on individual variation in the social behavior of dairy cattle

Dairy cattle typically live in groups, but individuals within these groups vary in their social behavior. An improved understanding of factors affecting the expression of social behavior may help refine management practices on farms to better accommodate the needs of all individuals within the herd. In this paper, we review (1) some examples of how social behavior is expressed in cattle, (2) commonly assessed personality traits in this species (i.e., sociability and fearfulness) as well as coping style, and (3) how these can affect the expression of social behavior of dairy cattle and in turn their welfare. We also identify understudied social behaviors that personality might influence (social learning, social stress, and social buffering of negative emotions), and that could inform how to improve the welfare of intensively housed dairy cattle.

Not so fast: Paradoxically increased variability in the glucose tolerance test due to food withdrawal in continuous glucose-monitored mice

ObjectiveThis study was performed to determine the effect of fasting on reproducibility of the glucose tolerance test. Due to individual variation in animal feeding behaviors, fasting animals prior to metabolic and behavioral experiments is widely held to reduce inter-subject variation in glucose and metabolic parameters of preclinical rodent models. Reducing variability is especially important for studies where initial metabolite levels can influence the magnitude of experimental interventions, but fasting also imposes stress that may distort the variables of interest. One such intervention is the glucose tolerance test (GTT) which measures the maximum response and recovery following a bolus of exogenous glucose. We sought to investigate how fasting affects the response of individual mice to a GTT. MethodsUsing simultaneous continuous glucose monitoring (CGM) and indirect calorimetry, we quantified blood glucose, physical activity, body temperature, metabolic rates, and food consumption levels on a minute-to-minute basis in adult male mice for 4 weeks. We tested the effects of a 4-h or 18-h fast on the GTT to examine the effect of food withdrawal in light or dark photoperiods. Studies were also performed with 4-h fasting in additional mice without implanted CGM probes. ResultsContrary to our expectations, a 4-h fast during the light photoperiod promotes a paradoxical increase in inter-animal variation in metabolic rate, physical activity, body temperature, glycemia, and glucose tolerance. This hyperglycemic and hyper-metabolic phenotype promotes increased corticosterone levels and is consistent with a behavioral stress response to food deprivation, even in well-fed mice. We find that mice undergoing an 18-h fast entered torpor, a hibernation-like state. In addition to low body temperature and metabolic rate, torpor is also associated with glucose levels 56 mg/dl lower than those seen in mice with ad libitum access to food. Moreover, the time spent in torpor affects the response to a GTT. ConclusionOur results suggest fasting mice before glucose tolerance testing, and perhaps other experiments, can have the opposite of the intended effect where fasting can increase, rather than decrease, experimental variability.

Biobehavioral susceptibility for obesity in childhood: Behavioral, genetic and neuroimaging studies of appetite

Modern food environments are conducive to overeating and weight gain, but not everyone develops obesity. One reason for this may be that individuals differ in appetitive characteristics, or traits, that manifest early in life and go on to influence their behavioral susceptibility to gain and maintain excess weight. Classic studies showing that eating behavior in children can be measured by behavioral paradigms such as tests of caloric compensation and eating in the absence of hunger inspired the development and validation of psychometric instruments to assess appetitive characteristics in children and infants. A large body of evidence now suggests that food approach traits increase obesity risk, while food avoidant traits, such as satiety responsiveness, decrease obesity risk. Twin studies and genetic association studies have demonstrated that appetitive characteristics are heritable, consistent with a biological etiology. However, family environment factors are also influential, with mounting evidence suggesting that genetic and environmental risk factors interact and correlate with consequences for child eating behavior and weight. Further, neuroimaging studies are revealing that individual differences in responses to visual food cues, as well as to small tastes and larger amounts of food, across a number of brain regions involved in reward/motivation, cognitive control and other functions, may contribute to individual variation in appetitive behavior. Growing evidence also suggests that variation on psychometric measures of appetite is associated with regional differences in brain structure, and differential patterns of resting state functional connectivity. Large prospective studies beginning in infancy promise to enrich our understanding of neural and other biological underpinnings of appetite and obesity development in early life, and how the interplay between genetic and environmental factors affects appetitive systems. The biobehavioral susceptibility model of obesity development and maintenance outlined in this narrative review has implications for prevention and treatment of obesity in childhood.

The influence of flow on movement of a headwater specialist in an intermittent urban headwater stream

AbstractFish movement in freshwater streams is often tied to patterns of flow, especially when the stream is intermittent, flashy, or some combination thereof. Relationships between flow and fish movement are exacerbated in small urban streams in impervious surface‐dominated watersheds that lead to extreme flow conditions. Here we examined the response of Creek Chub (Semotilus atromaculatus), a common pioneer fish species in eastern North America, to flow events of various sizes in a small intermittent urban headwater stream. Our primary goals were to determine (1) the level of Creek Chub movement occurring in an intermittent headwater stream, (2) the influence of flow on movement direction and magnitude, (3) the degree of variation in individual movement behaviour, and (4) mortality rates in the system. The study used PIT tagging and telemetry surveys to monitor Creek Chub movement over a six‐month period. We found that individual Creek Chub in our study system were generally mobile (71%), but usually moved short distances only; however, a few individuals moved much farther, with a maximum single movement of 521 m. Moderate or major flow events increased the probability and magnitude of Creek Chub movement, but there was no detectable asymmetry towards upstream or downstream directionality. Movement was variable among individual fish and fish were able to respond to large and small flow events to move among habitats. Finally, we observed relatively high, but episodic, mortality in our study site, potentially related to pollution events. Evidence of elevated mortality points to the potential importance of movement in highly disturbed streams, as it may enable recolonization after local extirpation.

Disparate behavioral types in wild and reared juveniles of gilthead seabream

Fish differ consistently in behavior within the same species and population, reflecting distinct behavioral types (BTs). Comparing the behavior of wild and reared individuals provides an excellent opportunity to delve into the ecological and evolutionary consequences of BTs. In this work, we evaluated the behavioral variation of wild and reared juvenile gilthead seabreams, Sparus aurata, a highly relevant species for aquaculture and fisheries. We quantified behavioral variation along the five major axes of fish behavioral traits (exploration-avoidance, aggressiveness, sociability, shyness-boldness, and activity) using standardized behavioral tests and a deep learning tracking algorithm for behavioral annotation. Results revealed significant repeatability in all five behavior traits, suggesting high consistency of individual behavioral variation across the different axes in this species. We found reared fish to be more aggressive, social and active compared to their wild conspecifics. Reared individuals also presented less variance in their aggressiveness, lacking very aggressive and very tame individuals. Phenotypic correlation decomposition between behavioral types revealed two different behavioral syndromes: exploration-sociability and exploration-activity. Our work establishes the first baseline of repeatability scores in wild and reared gilthead seabreams, providing novel insight into the behavior of this important commercial species with implications for fisheries and aquaculture.

Individual variation in preference behavior in sailfin fish refines the neurotranscriptomic pathway for mate preference.

Social interactions can drive distinct gene expression profiles which may vary by social context. Here we use female sailfin molly fish (Poecilia latipinna) to identify genomic profiles associated with preference behavior in distinct social contexts: male interactions (mate choice) versus female interactions (shoaling partner preference). We measured the behavior of 15 females interacting in a non-contact environment with either two males or two females for 30 min followed by whole-brain transcriptomic profiling by RNA sequencing. We profiled females that exhibited high levels of social affiliation and great variation in preference behavior to identify an order of magnitude more differentially expressed genes associated with behavioral variation than by differences in social context. Using a linear model (limma), we took advantage of the individual variation in preference behavior to identify unique gene sets that exhibited distinct correlational patterns of expression with preference behavior in each social context. By combining limma and weighted gene co-expression network analyses (WGCNA) approaches we identified a refined set of 401 genes robustly associated with mate preference that is independent of shoaling partner preference or general social affiliation. While our refined gene set confirmed neural plasticity pathways involvement in moderating female preference behavior, we also identified a significant proportion of discovered that our preference-associated genes were enriched for 'immune system' gene ontology categories. We hypothesize that the association between mate preference and transcriptomic immune function is driven by the less well-known role of these genes in neural plasticity which is likely involved in higher-order learning and processing during mate choice decisions.

Boldness and physiological variation in round goby populations along their Baltic Sea invasion front

The round goby (Neogobius melanostomus) is a fish native to the Ponto-Caspian region that is highly invasive through freshwater and brackish habitats in northern Europe and North America. Individual behavioural variation appears to be an important factor in their spread, for example a round goby's personality traits can influence their dispersal tendency, which may also produce variation in the behavioral composition of populations at different points along their invasion fronts. To further analyze the drivers of behavioral variation within invasive round goby populations, we focused on two populations along the Baltic Sea invasion front with closely comparable physical and community characteristics. Specifically, this study measured personality within a novel environment and predator response context (i.e., boldness), and directly analyzed links between individuals’ personality traits and their physiological characteristics and stress responses (i.e., blood cortisol and lactate, brain neurotransmitters). In contrast to previous findings, the more recently established population had similar activity levels but were less bold in response to a predator cue than the older population, which suggests that behavioral compositions within our study populations may be more driven by local environmental conditions rather than being a result of personality-biased dispersal. Furthermore, we found that both populations showed similar physiological stress responses, and there also appeared to be no detectable relationship between physiological parameters and behavioral responses to predator cues. Instead, body size and body condition were important factors influencing individual behavioral responses. Overall, our results reinforce the importance of boldness traits as a form of phenotypic variation in round goby populations in the Baltic Sea. We also highlight the importance of these traits for future studies specifically testing for effects of invasion processes on phenotypic variation in the species. Nonetheless, our results also highlight that the physiological mechanisms underpinning behavioural variation in these populations remain unclear.

Mid‐summer fish behavior in a high‐latitude twilight zone

AbstractThe behavior of the mesopelagic fish Benthosema glaciale was studied at 60°N in mid‐summer. We hypothesized that diel vertical migration (DVM) is constrained by short and dusk nights (surface illumination > 10−2 μmol m−2 s−1) and that individuals are active at depth during the long summer days. Submerged echosounders provided high‐resolution data throughout the water column. During the day, a part of the population ascended toward the increasing daylight. Short vertical relocations were followed by minutes of vertical inactivity. Swimming included horizontal turns and loops associated with the vertical steps. Normal DVM was initiated ~ 4 h before sunset and reflected independent individual decisions. The fish initially ascended stepwise but switched to mostly straight upwards swimming attaining 3–4 cm s−1. Their vertical speed was faster than the slow ascent of isolumes and even the deepest living fish potentially could reach upper layers shortly after sunset. However, many individuals aborted their ascent and returned to depth before the darkest time of the night, while others returned downward closer to sunrise. The daytime swimming and individual variability in diel migration behavior have implications for encounters with prey and predators in the twilight zone and the biological carbon pump. A principal conclusion is that mesopelagic fishes can modify their behavior and migration patterns to suit a wide range of changing conditions.