Diel vertical migration of freshwater fishes – proximate triggers,ultimate causes and research perspectives

1. Diel vertical migrations (DVM) are typical for many cold‐water fish species such as Pacific salmons (Oncorhynchus spp.) and coregonids (Coregonus spp.) inhabiting deep lakes. A comprehensive recent overview of DVM in freshwater fish has not been available, however. 2. The main proximate trigger of DVM in freshwater fish is the diel change in light intensity, with declining illumination at dusk triggering the ascent and the increase at dawn triggering the descent. Additional proximate cues are hydrostatic pressure and water temperature, which may guide fish into particular water layers at night. 3. Ultimate causes of DVM encompass bioenergetics efficiency, feeding opportunities and predator avoidance. None of these factors alone can explain the DVM in all cases. Multi‐factorial hypotheses, such as the ‘antipredation window’ combined with the thermal niche hypothesis, are more likely to explain DVM. It is suggested that planktivorous fish move within a layer sufficiently well illuminated to capture zooplankton, but too dark for predators to feed upon the migrating fish. In complete darkness, fish seek layers with a temperature that optimises bioenergetics efficiency. The strength of each factor may differ from lake to lake, and hence system‐specific individual analyses are needed. 4. Mechanistic details that are still poorly explored are the costs of buoyancy regulation and migration, the critical light thresholds for feeding of planktivorous and piscivorous fish, and predator assessment by (and size‐dependent predation risk of) the prey fish. 5. A comprehensive understanding of the adaptive value of DVM can be attained only if the behaviour of individual fish within migrating populations is explicitly taken into account. Size, condition and reproductive value differ between individuals, suggesting that migrating populations should split into migrants and non‐migrants for whom the balance between mortality risk and growth rate can differ. There is increasing evidence for this type of partial DVM within populations. 6. Whereas patterns of DVM are well documented, the evolution of DVM is still only poorly understood. Because experimental approaches at realistic natural scales remain difficult, a combination of comprehensive data sets with modelling is likely to resolve the relative importance of different proximate and ultimate causes behind DVM in fish.     

The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton

Many pelagic animal species in the marine environment and in lakes migrate to deeper water layers before sunrise and return around sunset. The amplitude of these diel vertical migrations (DVM) varies from several hundreds of metres in the oceans to approx. 5–20 m in lakes. DVM can be studied from a proximate and an ultimate point of view. A proximate analysis is intended to reveal the underlying behavioural mechanism and the factors that cause the daily displacements. The ultimate analysis deals with the adaptive significance of DVM and the driving forces that were responsible for the selection of the traits essential to the behavioural mechanism. The freshwater cladoceran Daphnia is the best studied species and results can be used to model migration behaviour in general. Phototaxis in Daphnia spp., which is defined as a light-oriented swimming towards (positive phototaxis) or away (negative phototaxis) from a light source, is considered the most important mechanism basic to DVM. A distinction has been made between primary phototaxis which occurs when light intensity is constant, and secondary phototaxis which is caused by changes in light intensity. Both types of reaction are superimposed on normal swimming. This swimming of Daphnia spp. consists of alternating upwards and downwards displacements over small distances. An internal oscillator seems to be at the base of these alternations. Primary phototaxis is the result of a dominance of either the upwards or the downwards oscillator phase, and the direction depends on internal and external factors: for example, fish-mediated chemicals or kairomones induce a downwards drift. Adverse environmental factors may produce a persistent primary phototaxis. Rare clones of D. magna have been found that show also persistent positive or negative primary phototaxis and interbreeding of the two types produces intermediate progeny: thus a genetic component seems to be involved. Also secondary phototaxis is superimposed on normal swimming: a continuous increase in light intensity amplifies the downwards oscillator phase and decreases the upwards phase. A threshold must be succeeded which depends on the rate and the duration of the relative change in light intensity. The relation between both is given by the stimulus strength versus stimulus duration curve. An absolute threshold or rheobase exists, defined as the minimum rate of change causing a response if continued for an infinitely long time. DVM in a lake takes place during a period of 1-5-2 h when light changes are higher than the rheobase threshold. Accelerations in the rate of relative increase in light intensity strongly enhance downwards swimming in Daphnia spp. and this enhancement increases with increasing fish kairomone and food concentration. This phenomenon may represent a ‘decision-making mechanism’ to realize the adaptive goal of DVM: at high fish predator densities, thus high kairomone concentrations, and sufficiently high food concentrations, DVM is profitable but not so at low concentrations. Body axis orientation in Daphnia spp. is controlled with regard to light-dark boundaries or contrasts. Under water, contrasts are present at the boundaries of the illuminated circular window which results from the maximum angle of refraction at 48–9° with the normal (Snell's window). Contrasts are fixed by the compound eye and appropriate turning of the body axis orients the daphnid in an upwards or an obliquely downwards direction. A predisposition for a positively or negatively phototactic orientation seems to be the result of a disturbed balance of the two oscillators governing normal swimming. Some investigators have tried to study DVM at a laboratory scale during a 24 h cycle. To imitate nature, properties of a natural water column, such as a large temperature gradient, were compressed into a few cm. With appropriate light intensity changes, vertical distributions looking like DVM were obtained. The results can be explained by phototactic reactions and the artificial nature of the compressed environmental factors but do not compare with DVM in the field. A mechanistic model of DVM based on phototaxis is presented. Both, primary and secondary phototaxis is considered an extension of normal swimming. Using the light intensity changes of dawn and the differential enhancement of kairomones and food concentrations, amplitudes of DVM could be simulated comparable to those in a lake. The most important adaptive significance of DVM is avoidance of visual predators such as juvenile fish. However, in the absence of fish kairomones, small-scale DVMs are often present, which were probably evolved for UV-protection, and are realized by not enhanced phototaxis. In addition, the ‘decision-making mechanism’ was probably evolved as based on the enhanced phototactic reaction to accelerations in the rate of relative changes in light intensity and the presence of fish kairomones.     

By-product runaway evolution by adaptive mate choice: A behavioural aspect of sexual selection

Summary From a behavioural perspective on adaptive female choice, I developed a by-product runaway model of adaptive mate choice. The model illustrates the evolution of the tail size of peacocks. I consider the causal mechanisms of adaptive female choice: (1) why (ultimate reasons); (2) how (proximate mechanisms). Assumptions are developed based on these behavioural aspects. For (1) ultimate reasons, I assume that many male losers (low-fitness males) always occur due to genetic and environmental uncertainty (A-1). For (2) proximate mechanisms, I assume that losers tend to differ in the expression of a fitness-sensitive trait (an ultimate target, e.g. body size; A-2), that the fitness-sensitive trait correlates with a secondary sexual trait (a proximate cue, e.g. allometry in body size and tail size; (A-3), and that the cue trait has a genetic basis that is independent of the target trait (e.g. a genetic basis in tail ratio to body size; A-4). The model's results are: persistent female choice by means of a proximate cue (R-1); by-product selection on the independent genetic basis of the cue (R-2); and the non-adaptive or maladaptive runaway evolution of the male proximate cue (R-3). In this model, female mate preferences are non-arbitrary and adaptive, whereas the resulting evolution of male secondary sexual traits is non-adaptive in the sense of survival selection.     

Social plasticity in fish: integrating mechanisms and function

Social plasticity is a ubiquitous feature of animal behaviour. Animals must adjust the expression of their social behaviour to the nuances of daily social life and to the transitions between life‐history stages, and the ability to do so affects their Darwinian fitness. Here, an integrative framework is proposed for understanding the proximate mechanisms and ultimate consequences of social plasticity. According to this framework, social plasticity is achieved by rewiring or by biochemically switching nodes of the neural network underlying social behaviour in response to perceived social information. Therefore, at the molecular level, it depends on the social regulation of gene expression, so that different brain genomic and epigenetic states correspond to different behavioural responses and the switches between states are orchestrated by signalling pathways that interface the social environment and the genotype. At the evolutionary scale, social plasticity can be seen as an adaptive trait that can be under positive selection when changes in the environment outpace the rate of genetic evolutionary change. In cases when social plasticity is too costly or incomplete, behavioural consistency can emerge by directional selection that recruits gene modules corresponding to favoured behavioural states in that environment. As a result of this integrative approach, how knowledge of the proximate mechanisms underlying social plasticity is crucial to understanding its costs, limits and evolutionary consequences is shown, thereby highlighting the fact that proximate mechanisms contribute to the dynamics of selection. The role of teleosts as a premier model to study social plasticity is also highlighted, given the diversity and plasticity that this group exhibits in terms of social behaviour. Finally, the proposed integrative framework to social plasticity also illustrates how reciprocal causation analysis of biological phenomena (i.e. considering the interaction between proximate factors and evolutionary explanations) can be a more useful approach than the traditional proximate–ultimate dichotomy, according to which evolutionary processes can be understood without knowledge on proximate causes, thereby black‐boxing developmental and physiological mechanisms.     

Two hundred years of zooplankton vertical migration research

Vertical migration is a geographically and taxonomically widespread behaviour among zooplankton that spans across diel and seasonal timescales. The shorter-term diel vertical migration (DVM) has a periodicity of up to 1 day and was first described by the French naturalist Georges Cuvier in 1817. In 1888, the German marine biologist Carl Chun described the longer-term seasonal vertical migration (SVM), which has a periodicity of ca. 1 year. The proximate control and adaptive significance of DVM have been extensively studied and are well understood. DVM is generally a behaviour controlled by ambient irradiance, which allows herbivorous zooplankton to feed in food-rich shallower waters during the night when light-dependent (visual) predation risk is minimal and take refuge in deeper, darker waters during daytime. However, DVMs of herbivorous zooplankton are followed by their predators, producing complex predator–prey patterns that may be traced across multiple trophic levels. In contrast to DVM, SVM research is relatively young and its causes and consequences are less well understood. During periods of seasonal environmental deterioration, SVM allows zooplankton to evacuate shallower waters seasonally and take refuge in deeper waters often in a state of dormancy. Both DVM and SVM play a significant role in the vertical transport of organic carbon to deeper waters (biological carbon sequestration), and hence in the buffering of global climate change. Although many animal migrations are expected to change under future climate scenarios, little is known about the potential implications of global climate change on zooplankton vertical migrations and its impact on the biological carbon sequestration process. Further, the combined influence of DVM and SVM in determining zooplankton fitness and maintenance of their horizontal (geographic) distributions is not well understood. The contrasting spatial (deep versus shallow) and temporal (diel versus seasonal) scales over which these two migrations occur lead to challenges in studying them at higher spatial, temporal and biological resolution and coverage. Extending the largely population-based vertical migration knowledge base to individual-based studies will be an important way forward. While tracking individual zooplankton in their natural habitats remains a major challenge, conducting trophic-scale, high-resolution, year-round studies that utilise emerging field sampling and observation techniques, molecular genetic tools and computational hardware and software will be the best solution to improve our understanding of zooplankton vertical migrations.     

The levels of analysis revisited

The term levels of analysis has been used in several ways: to distinguish between ultimate and proximate levels, to categorize different kinds of research questions and to differentiate levels of reductionism. Because questions regarding ultimate function and proximate mechanisms are logically distinct, I suggest that distinguishing between these two levels is the best use of the term. Integrating across levels in research has potential risks, but many benefits. Consideration at one level can help generate novel hypotheses at the other, define categories of behaviour and set criteria that must be addressed. Taking an adaptationist stance thus strengthens research on proximate mechanisms. Similarly, it is critical for researchers studying adaptation and function to have detailed knowledge of proximate mechanisms that may constrain or modulate evolutionary processes. Despite the benefits of integrating across ultimate and proximate levels, failure to clearly identify levels of analysis, and whether or not hypotheses are exclusive alternatives, can create false debates. Such non-alternative hypotheses may occur between or within levels, and are not limited to integrative approaches. In this review, I survey different uses of the term levels of analysis and the benefits of integration, and highlight examples of false debate within and between levels. The best integrative biology reciprocally uses ultimate and proximate hypotheses to generate a more complete understanding of behaviour.     

Ultimate explanations concern the adaptive rationale for organism design

My understanding is that proximate explanations concern adaptive mechanism and that ultimate explanations concern adaptive rationale. Viewed in this light, the two kinds of explanation are quite distinct, but they interact in a complementary way to give a full understanding of biological adaptations. In contrast, Laland et al. (2013)—following a literal reading of Mayr (Science 134:1501–1506, 1961)—have characterized ultimate explanations as concerning any and all mechanisms that have operated over the course of an organism’s evolutionary history. This has unfortunate consequences, such as allowing random drift to form the basis for ultimate explanations, and allowing proximate and ultimate explanations to bleed into each other until their distinction is meaningless. Here, I suggest Laland et al’s explanatory framework of “reciprocal causation” is not conducive to successful biological science, and that they have misunderstood key elements of the theory of Darwinian adaptation.     

Morphology and behaviour: functional links in development and evolution

Development and evolution of animal behaviour and morphology are frequently addressed independently, as reflected in the dichotomy of disciplines dedicated to their study distinguishing object of study (morphology versus behaviour) and perspective (ultimate versus proximate). Although traits are known to develop and evolve semi-independently, they are matched together in development and evolution to produce a unique functional phenotype. Here I highlight similarities shared by both traits, such as the decisive role played by the environment for their ontogeny. Considering the widespread developmental and functional entanglement between both traits, many cases of adaptive evolution are better understood when proximate and ultimate explanations are integrated. A field integrating these perspectives is evolutionary developmental biology (evo-devo), which studies the developmental basis of phenotypic diversity. Ultimate aspects in evo-devo studies--which have mostly focused on morphological traits--could become more apparent when behaviour, 'the integrator of form and function', is integrated into the same framework of analysis. Integrating a trait such as behaviour at a different level in the biological hierarchy will help to better understand not only how behavioural diversity is produced, but also how levels are connected to produce functional phenotypes and how these evolve. A possible framework to accommodate and compare form and function at different levels of the biological hierarchy is outlined. At the end, some methodological issues are discussed.     

The primate palette: The evolution of primate coloration

Flip through The Pictorial Guide to the Living Primates¹ and you will notice a striking yet generally underappreciated aspect of primate biology: primates are extremely colorful. Primate skin and pelage coloration were highlighted examples in Darwin's² original discussions of sexual selection but, surprisingly, the topic has received little research attention since. Here we summarize the patterns of color variation observed across the primate order and examine the selective forces that might drive and maintain this aspect of primate phenotypic diversity. We discuss how primate color patterns might be adaptive for physiological function, crypsis, and communication. We also briefly summarize what is known about the genetic basis of primate pigmentation and argue that understanding the proximate mechanisms of primate coloration will be essential, not only for understanding the evolutionary forces shaping phenotypic variation, but also for clarifying primate taxonomies and conservation priorities.     

Dose‐dependent effects of an immune challenge at both ultimate and proximate levels in Drosophila melanogaster

Immune responses are highly dynamic. The magnitude and efficiency of an immune response to a pathogen can change markedly across individuals, and such changes may be influenced by variance in a range of intrinsic (e.g. age, genotype, sex) and external (e.g. abiotic stress, pathogen identity, strain) factors. Life history theory predicts that up‐regulation of the immune system will come at a physiological cost, and studies have confirmed that increased investment in immunity can reduce reproductive output and survival. Furthermore, males and females often have divergent reproductive strategies, and this might drive the evolution of sex‐specific life history trade‐offs involving immunity, and sexual dimorphism in immune responses per se. Here, we employ an experiment design to elucidate dose‐dependent and sex‐specific responses to exposure to a nonpathogenic immune elicitor at two scales – the ‘ultimate’ life history and the underlying ‘proximate’ immune level in Drosophila melanogaster. We found dose‐dependent effects of immune challenges on both male and female components of reproductive success, but not on survival, as well as a response in antimicrobial activity. These results indicate that even in the absence of the direct pathogenic effects that are associated with actual disease, individual life histories respond to a perceived immune challenge – but with the magnitude of this response being contingent on the initial dose of exposure. Furthermore, the results indicate that immune responses at the ultimate life history level may indeed reflect underlying processes that occur at the proximate level.     

Long-term Population Trends in White-Footed Mice and the Impact of Supplemental Food and Shelter

Central to most models of population regulation is the ideathat the degree of intraspecific competition is in some wayproportional to the availability of limiting resources. Althoughlaboratory research has demonstrated a number of proximate mechanismsby which behavior might affect population growth, little isknown about the resources that are actually limiting populationdensity in the field or about how animals might compete forthem. Long-term field studies reveal that the white-footed mouse,Peromyscus leucopus, exhibits two types of pronounced changesin density: intra-annual (seasonal) and inter-annual. The formerseem to be due ultimately to climate and the lack of winterbreeding, but populations often decline sharply in late summerin the midst of plentiful food; fall recruitment of new animalsis density-dependent and usually poor. Differences in peak densitiesfrom one year to the next as high as 13-fold have been recorded. Weather, shelter, and food are possible ultimate limiting resources.Food has received themost attention, but supplemental feedingexperiments have yielded mixed results. A review of the socialbehavior of this polygnous species suggests that each sex islimited by different combinations of factors. Females may defenda food source and nest sites; males may search actively forfemales at low adult densities, covering large areas, and defendaccess to females at high adult densities. Additional long-term field studies are needed, both to providedirection to laboratory research on proximate mechanisms andto provide the data base for understanding the role of weather,food, and shelter as ultimate limiting factors. Enough short-termstudies have been published to permit comparisons across habitatsand through time which will give a better perspective on climateand habitat variables. Field experiments are necessary to demonstratethe operation of proximate behavioral, physiological, and geneticmechanisms of population regulation in natural populations.     

Epigenetic rules and Darwinian algorithms: The adaptive study of learning and development

Recent efforts toward a Darwinian psychology of human behavior will profit from taking account of prior investigations of proximate phenomena and adaptive mechanisms conducted within the science of biology, and from realizing that adaptive significance and underlying mechanisms must be investigated in concert. Contrary to some recent arguments, evidence of adaptive design is usually manifested initially and most prominently in the behavior (or other “ultimate” phenotypic expressions) of organisms, human or nonhuman, rather than in underlying psychological, physiological, or developmental mechanisms, which are often obscure, and in any case, as adaptive mechanisms, must be investigated secondarily. The reason is that selection acts most directly on behavior, and on its underlying mechanisms only as they influence the behavior. This is as true for learned and cultural behaviors as for any others. Adaptive significance of behavior, and evidence of its underlying design, is thus examined only by studying the behavior itself, its complexity, the situations in which it is expressed, and its effects in different situations. Biological mechanisms of any kind cannot even be identified with confidence, or understood, until, at the least, reasonable inferences have been made about their adaptive functions, what they are, as mechanisms, designed by selection to accomplish. Moreover, what appear to be adaptive psychological, physiological, or other mechanisms, may, as with some expressions of behavior, be incidental effects of still other mechanisms that are adaptive.Adaptation is not restricted to situations in which genes program specifically for particular behavioral alternatives: natural selection of alternative alleles may also yield abilities and tendencies to engage in conditional strategies, to assess costs and benefits in directly or indirectly reproductive terms. In humans, such cost-benefit assessments may be conducted entirely through mental scenario-building, or even through absorbing and judging the mental scenarios of others, without either admission or cognizance of the reproductive significance of the assessment. The goals actually sought may be secondary, tertiary, or even more distantly removed correlates of reproductive success (e.g., status or reputation, which may correlate with power, which may correlate with wealth, which may correlate with access to the resources of reproduction); reproductive success itself may be a concept alien to the actor's conscious motivations, even denied vehemently as a goal. In learned and cultural behaviors, selection has to be not only for the ability to learn but for its patterning, such as for the machinery enabling development of the ability to learn to make appropriate (cultural) decisions.Kin recognition is reviewed as the most prominent example of a set of extensively studied adaptive mechanisms involving learning, and as a central problem with respect to adaptiveness in social behavior. Arguments that the adaptive mechanisms collected under the concept of learning evolve as special, rather than general-purpose devices, raise provocative questions about the evolution of ontogenetic and physiological preparation to deal with environmental novelty, especially in complex social interactions. Evolution of the human psyche, especially its conscious aspects, is briefly discussed as a problem in understanding the history of sociality. It is argued that the principal environment of natural selection leading to the modern human psyche was social, and that on this account the environment of human behavior has not changed as much since the Pleistocene as is often assumed.     

Developmental responses to early‐life adversity: Evolutionary and mechanistic perspectives

Adverse ecological and social conditions during early life are known to influence development, with rippling effects that may explain variation in adult health and fitness. The adaptive function of such developmental plasticity, however, remains relatively untested in long‐lived animals, resulting in much debate over which evolutionary models are most applicable. Furthermore, despite the promise of clinical interventions that might alleviate the health consequences of early‐life adversity, research on the proximate mechanisms governing phenotypic responses to adversity have been largely limited to studies on glucocorticoids. Here, we synthesize the current state of research on developmental plasticity, discussing both ultimate and proximate mechanisms. First, we evaluate the utility of adaptive models proposed to explain developmental responses to early‐life adversity, particularly for long‐lived mammals such as humans. In doing so, we highlight how parent‐offspring conflict complicates our understanding of whether mothers or offspring benefit from these responses. Second, we discuss the role of glucocorticoids and a second physiological system—the gut microbiome—that has emerged as an additional, clinically relevant mechanism by which early‐life adversity can influence development. Finally, we suggest ways in which nonhuman primates can serve as models to study the effects of early‐life adversity, both from evolutionary and clinical perspectives.     

Brood Value and Life Expectancy as Determinants of Parental Investment in Male Three-spined Sticklebacks,Gasterosteus aculeatus

29 breeding male three-spined sticklebacks (Gasterosteus aculeatus) caught in the wild at different times in the breeding season were exposed simultaneously to a potential threat to their nest (a conspecific male) and to two hunting trout (potential predators of adult sticklebacks, seen through a transparent partition). By promoting a variety of protective responses, the presence of the predators reduced both the time spent confronting the intruder and the rate at which it was attacked. Subjects with a clutch of eggs in their nest maintained higher levels of territorial defence in the presence of predators than did those with an empty nest. However, those breeding later in the season took fewer risks to defend their nest. Possible proximate mechanisms responsible for these results are discussed. In addition, the adaptive significance of the behavioural changes is considered in the light of the value of the brood and the expected future reproductive output of the sticklebacks breeding at different times in the season.     

Immune function across generations: integrating mechanism and evolutionary process in maternal antibody transmission

The past 30 years of immunological research have revealed much about the proximate mechanisms of maternal antibody transmission and utilization, but have not adequately addressed how these issues are related to evolutionary and ecological theory. Much remains to be learned about individual differences within a species in maternal antibody transmission as well as differences among species in transmission or utilization of antibodies. Similarly, maternal-effects theory has generally neglected the mechanisms by which mothers influence offspring phenotype. Although the environmental cues that generate maternal effects and the consequent effects for offspring phenotype are often well characterized, the intermediary physiological and developmental steps through which the maternal effect is transmitted are generally unknown. Integration of the proximate mechanisms of maternal antibody transmission with evolutionary theory on maternal effects affords an important opportunity to unite mechanism and process by focusing on the links between genetics, environment and physiology, with the ultimate goal of explaining differences among individuals and species in the transfer of immune function from one generation to the next.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Diel vertical migration of larval and early‐juvenile burbot optimises survival and growth in a deep,pre‐alpine lake

1. Burbot larvae (Lota lota) perform a substantial diel vertical migration (DVM) of increasing amplitude in the pelagic zone during a 3‐month period before migrating to the littoral zone as early‐juveniles. We hypothesised that feeding in the warm surface layers at night and then spending the day in cold water below the thermocline reduces metabolic costs and earns burbot larvae an energetic advantage. 2. To test our hypothesis, we mimicked the temperature conditions experienced by vertically migrating burbot in the pelagic zone. We also simulated three further scenarios, in which temperature remained constant. 3. Burbot showed the best performance (defined as specific growth rate multiplied by the probability of survival) in the treatments simulating DVM. The high temperature treatment, simulating permanent residence in the warm epilimnion, resulted in high growth combined with high mortality. At a permanently low temperature, simulating life in the hypolimnion, growth was poor and activity reduced. 4. In a deep, temperature‐stratified lake, where the apparently beneficial overall medium temperature is found in a restricted layer within the thermocline, DVM optimises performance in young burbot. Various ultimate factors might act synergistically in selecting for DVM in larval and early‐juvenile burbot.     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.