Climate change alters the reproductive phenology and investment of a lacustrine fish,the three‐spine stickleback

High‐latitude lakes are particularly sensitive to the effects of global climate change, demonstrating earlier ice breakup, longer ice‐free seasons, and increased water temperatures. Such physical changes have implications for diverse life‐history traits in taxa across entire lake food webs. Here, we use a five‐decade time series from an Alaskan lake to explore effects of climate change on growth and reproduction of a widely distributed lacustrine fish, the three‐spine stickleback (Gasterosteus aculeatus). We used multivariate autoregressive state‐space (MARSS) models to describe trends in the mean length for multiple size classes and to explore the influence of physical (date of ice breakup, surface water temperature) and biological (density of con‐ and heterospecifics) factors. As predicted, mean size of age 1 and older fish at the end of the growing season increased across years with earlier ice breakup and warmer temperatures. In contrast, mean size of age 0 fish decreased over time. Overall, lower fish density and warmer water temperatures were associated with larger size for all cohorts. Earlier ice breakup was associated with larger size for age 1 and older fish but, paradoxically, with smaller size of age 0 fish. To explore this latter result, we used mixing models on age 0 size distributions, which revealed an additional cohort in years with early ice breakup, lowering the mean size of age 0 fish. Moreover, early ice breakup was associated with earlier breeding, evidenced by earlier capture of age 0 fish. Our results suggest that early ice breakup altered both timing and frequency of breeding; three‐spine stickleback spawned earlier and more often in response to earlier ice breakup date. While previous studies have shown the influence of changing conditions in northern lakes on breeding timing and growth, this is the first to document increased breeding frequency, highlighting another pathway by which climate change can alter the ecology of northern lakes.     

Phylogeny and ontogeny of fish leucocytes

In contrast to higher vertebrates, most fish species hatch at the embryonic stage of life. Consequently, they have to defend against a variety of micro-organisms living in their aquatic environment. This paper is focussed on the development of leucocytes functioning within this early innate system and later on in the acquired immune system (B and T cells). Most of the data are derived from cyprinid fish (zebrafish, carp), which are excellent models to study early ontogeny. Attention is also paid to the phylogeny of leucocytes, with special attention to early chordates. It is clear that young fish use innate mechanisms during the first weeks/months of their development. In zebrafish, a variety of hematopoietic genes have been sequenced which allow a detailed picture of the development of the distinct leucocytes and their precursors. In cyprinids and sea bass, the thymus is the first lymphoid organ and T cells appear to be selected there much earlier than the first detection of T cell-dependent antibody responses. The first B cells are most probably generated in head kidney. Although T cells are selected earlier than B cells, T cell independent responses occur earlier than the T cell-dependent responses. The very early (pre-thymic) appearance of T-like cells in gut of sea bass and carp suggests an extra-thymic origin of these cells. However, B cells populate the GALT much later than spleen or kidney, indicating a rather late appearance of mucosal immunity. The first plasma cells are found long after the intake of food in cyprinids, but in many marine fish they appear around the first food intake. In general, acquired immunity is not correlated to food intake.     

Food web de‐synchronization in England's largest lake: an assessment based on multiple phenological metrics

Phenological changes have been observed globally for marine, freshwater and terrestrial species, and are an important element of the global biological ‘fingerprint’ of climate change. Differences in rates of change could desynchronize seasonal species interactions within a food web, threatening ecosystem functioning. Quantification of this risk is hampered by the rarity of long‐term data for multiple interacting species from the same ecosystem and by the diversity of possible phenological metrics, which vary in their ecological relevance to food web interactions. We compare phenological change for phytoplankton (chlorophyll a), zooplankton (Daphnia) and fish (perch, Perca fluviatilis) in two basins of Windermere over 40 years and determine whether change has differed among trophic levels, while explicitly accounting for among‐metric differences in rates of change. Though rates of change differed markedly among the nine metrics used, seasonal events shifted earlier for all metrics and trophic levels: zooplankton advanced most, and fish least, rapidly. Evidence of altered synchrony was found in both lake basins, when combining information from all phenological metrics. However, comparisons based on single metrics did not consistently detect this signal. A multimetric approach showed that across trophic levels, earlier phenological events have been associated with increasing water temperature. However, for phytoplankton and zooplankton, phenological change was also associated with changes in resource availability. Lower silicate, and higher phosphorus, concentrations were associated with earlier phytoplankton growth, and earlier phytoplankton growth was associated with earlier zooplankton growth. The developing trophic mismatch detected between the dominant fish species in Windermere and important zooplankton food resources may ultimately affect fish survival and portend significant impacts upon ecosystem functioning. We advocate that future studies on phenological synchrony combine data from multiple phenological metrics, to increase confidence in assessments of change and likely ecological consequences.     

Inhibition of optimal behavior by social transmission in the guppy depends on shoaling

Previous research has suggested social learning of foragingbehavior can inhibit learning of the optimal behavior pattern.Based on their transmission chain design, we used small groupsof guppies (Poecilia reticulata) to determine the degree towhich the optimal behavior pattern was inhibited by sociallylearned information. A founder group was trained to take a long, energetically costly route to a food source. The members ofthis group were gradually replaced with naive conspecifics.Replicating the findings of the earlier researchers, it wasclear that the behavior of the founders strongly influencedthe behavior of the naive fish, probably through a process oflocal enhancement. When tested as a group, the naive fish chosethe long route to the food source significantly more oftenthan chance. Each naive fish was also tested in isolation.When tested alone, there was a significant tendency to choosethe short route despite following the long route when testedas a group. These results suggest social learning does notinhibit learning of optimal behavior patterns but that a trade-offoccurs when tested in the group condition. It is possible thatthe advantages for an individual fish of swimming with theshoal, and thus following the socially learned route, may haveoutweighed the potential energetic costs of taking this longerroute.     

Choice of feeding station in Atlantic salmon, Salmo salar, parr: effects of predation risk, season and life history strategy

The current speed at which underyearling salmon parr held feeding stations was examined from late summer to early winter in laboratory flume tanks that offered a choice between (a) areas with high water flow, high food availability but high predation risk and (b) areas with low flow, little food but shelter from predators. In August, those fish that would become smolts aged 1 + (and which by late winter formed the upper modal group, UMG, of the bimodal size distribution) adopted positions in faster currents than did the fish which would take a further year to reach the smolt stage (the lower modal group, LMG). However, the chosen current speed of UMG fish decreased through the period of study, so that by December all fish were found in areas of low flow, and hence little food. Both date and water temperature had independent effects on the chosen current speed of UMG fish.
The effect of predation risk was investigated using a model trout. A brief sight of this predator caused 47% of fish that had been in the main, exposed currents to move to slacker, sheltered areas; they took 1 h, on average, to return to their previous position. The fish that remained in position upon seeing the predator reduced their rate of tail beating, presumably increasing crypticity. Eventual UMG parr were less likely than were LMG fish to move away upon seeing a predator. The fish moved to faster currents than normal 2–3 h after seeing the predator, possibly compensating for the earlier reduction in feeding rate.     

Diet of post-smolt and one-sea-winter Atlantic salmon in the Bothnian Sea, Northern Baltic

Until July, post-smolt salmon Salmo salar ( n =337; 129–375 mm L _T, mean 225 mm) in the Bothnian Sea relied largely on surface fauna (mainly terrestrial insects). From August onwards, fish was the principal food type. The smallest piscivorous post-smolts were <200 mm, but the main shift to piscivory occurred at sizes of 240–320 mm. Piscivory was promoted by a large smolt size. Almost all one-sea-winter (1-SW) salmon ( n =316; 278–524 mm, mean 397 mm) were piscivorous. Over 70% of the post-smolt and 96% of the 1-SW salmon with identifiable fish species in their stomachs had preyed on herring Clupea harengus. Other fish prey included the ten-spined Pungitius pungitius and three-spined sticklebacks Gasterosteusaculeatus but no sprats Sprattus sprattus. The results support earlier observations of a close relationship between recruitment of herring and production of salmon in the Bothnian Sea, and of the crucial role of smolt size in determining the ability of feeding salmon for utilizing the food resources of the area.     

The Ecology of Temporary Streams III. Temporary Stream Fishes in Southern Ontario,Canada

The 12 fish species living in three temporary streams in Southern Ontario, Canada, during 1973–1974, are listed. Environmental parameters are recorded together with details of the abundance, distribution, age structure, growth, reproduction and food of the more common species. These species are thought to be either physiologically very tolerant, or to be capable of temporal or spatial migrations. Colonization of temporary streams bestows benefits in terms of plentiful invertebrate food, although resource sharing may occur, earlier spring breeding and reduced predation.     

Studies on the Genetics of Feeding Behaviour in the Cave Fish Astyanax mexicanus f. anoptichthys

The Mexican characid Astyanax mexicanus gave rise to a series of cavcrnicolous populations (“Anoptichthys”) not earlier than during the Pleistocene age. When searching for food the blind cave fish swims at an angle of about 45° to the ground. The river fish, however, when having no visual information, stands vertically on its head. With the aid of infrarcd-videorecording and on the basis of crossing experiments it could be demonstrated that this difference in feeding behaviour is controlled by polygenes, although frequency distributions of the crossings suggest an apparent monofactorial inheritance. The evolutionary processes of adaptive reduction of headstanding and of progressive development of the gustatory equipment are genetically independent and both traits have been achieved by small genetic steps.     

Growth and feeding of juvenile pike <Emphasis Type="Italic">Esox lucius</Emphasis> (Esocidae) of different hatching dates during its first months of life

Groups of fingerlings of the pike Esox lucius, that hatched from the eggs of the same female at the beginning and end of hatching are compared. It was found that larvae of pike of the beginning of hatching passed to external feeding earlier. In the period of mixed and exogenous feeding, they not only grew, but also developed more rapidly than larvae from the same brood. It was shown that pike juveniles of the end of hatching were characterized by a retarded development and growth in the first weeks of life (until reaching a length of 100 mm and a weight of 7 g). In the second month of life, specimens of this group began to feed intensively, grow, and according to body length and weight, reached pike juveniles of the beginning of hatching. Pike fingerlings of the end of hatching were characterized by compensation growth that was manifested at the excess of fish food (at cultivation in separate aquariums when the inhibitory effect of large fish on the growth of small fish was excluded) and in ponds where fish food availability was considerably lower.     

Growling for food: acoustic emissions during competitive feeding of the streaked gurnard

The streaked gurnard Trigloporus lastoviza produced only one sound type, a growl, lasting up to 3 s and consisting of repeated groups of typically one to three pulses. The foraging fish followed two different strategies. In the first, the fish circled the feeding area, grasped a food item and fled, sometimes displaying aggressively to competitors. With this foraging strategy, fish usually made sounds as they circled, grasped and fled. Fish that growled while circling were more likely to grasp a food item subsequently than were silent fish. The second feeding strategy occurred when a fish had already ingested food or failed to get any. In this case, typically fish searched for food on the substratum or approached and touched other individuals that were feeding, sometimes grabbing food that was spat out during food handling by the other fish. Although payback experiments would be needed to draw firm conclusions on the communicative function of growling during competitive feeding in the streaked gurnard, the results suggest that sound production confers advantages to individuals competing for limited food resources.     

Phenotypic plasticity in Daphnia magna Straus: variable maturation instar as an adaptive response to predation pressure

Life history responses of four Daphnia magna clones at two food levels were studied to assess the importance of maturation instar on the plasticity of fitness responses under simulated mortality regimes. Females of the clones studied could vary offspring size with consequent effects on their maturation time. Significant genetic variability in life history and fitness responses, measured as the intrinsic rate of population increase, within and across food levels was observed, but most of this variation could be attributed to maturation instar differences among clones within and across environments. In the laboratory, without extrinsic mortality, females maturing earlier always had higher fitness than those maturing later, indicating a clear fitness cost of delaying maturity. Nevertheless using a model, we showed that the observed maturation instar effects on life history responses can lead to differences in fitness under different size-selective predation regimes, such that females with delayed maturity have higher fitness under invertebrate predation while females maturing earlier have higher fitness under fish predation regimes. These results suggest that intraclonal variation in offspring size and hence in the number of maturation instars can be an adaptation to living in habitats subject to temporal fluctuations in fish and invertebrate predation pressure.     

鱼体(去鳃)和鱼鳃对不同形态铜的积累特征

在实验室条件下研究实验鱼Ｐａｒａｃｈｅｉｒｏｄｏｎ对人工河水中不同形态的积累特征,对比了鱼体（去鳃）和鱼鳃对铜吸收量的差异,并探讨了鱼对铜的吸收机理。研究结果表明,实验鱼鳃部和体内铜积累量均随水相游离铜浓度增高,暴露时间增长而增加,但鳃部积累浓度较鱼体其余部分高一个数量级,其从水相富集铜的速率显著高于鱼体。     

Accumulation of plerocercoids of Triaenophorus crassus in the second intermediate host Coregonus lavaretus and their effect on growth of the host

Patterns of accumulation of Triaenophorus crassus in its second intermediate host whitefish Coregonus lavaretus s.l. were studied between 1991 and 1996 from two host populations in two separate areas of Lake Saimaa, Finland. Whitefish were infected commonly with several T. crassus plerocercoids and the parasites were aggregated into the oldest hosts. In one host population the annual parasite accumulation was 0·9 parasites in all host age groups between 3 and 8 years. In the other host population the annual accumulation was 1·6 parasites in 3–5-year-old fish, but increased up to 3 to 4 parasites per year in fish over 5 years old. The increase did not coincide with the period of maturation or any increase in whitefish growth, both of which could alter the food intake of the fish. The sharp increase in the annual accumulation suggests a threshold intensity above which the probability of acquiring further parasites increases. In spite of a heavy aggregation of parasites there was no evidence of parasite-induced host mortality. The annual increase in mean abundance was not correlated with the mean annual weight increase in 2–4-year-old fish within cohorts. However, evidence of a negative effect of parasites on whitefish growth was revealed by back-calculation of lengths of uninfected and infected whitefish and correlations between length or weight of fish and intensity of infection with fish age. Both analyses suggested that larger young fish harboured more parasites than the smaller ones while in older fish the reverse was true, a pattern that has not been shown earlier for parasitized fish.     

Juvenile migration in brown trout: a consequence of energetic state

1. We explored the mechanisms determining age and size at juvenile migration in brown trout Salmo trutta L. A ¹³³Cs tracer methodology was used to estimate food consumption of juvenile brown trout in a Norwegian stream, and the energy budgets of early migrants and stream residents were compared.
2. Fast-growing brown trout migrated to the lake earlier and at a smaller body size than slower-growing individuals. The 2+ migrants were significantly larger than those that remained 1 or more years longer in the stream. The 3+ migrants were significantly larger than the 2+ migrants. Some fast-growing males matured in the stream, whereas all females left the stream before maturing sexually.
3. The food consumption and the energy budgets for 2+ migrants were more than four times higher than those of the resident 2+ fish. Total energy allocated to growth was also higher among migrants, and the total metabolic costs were five times higher among migrants than among resident fish.
4. The proportional energy allocation to growth among the 2+ migrants was much lower (about half) than that of those remaining longer in the stream. The reduction in the proportion of energy available for growth from age 1+ to 2+ was larger among migrants (88%) than among resident fish (68%). Reduction in the proportion of energy available for growth is a probable explanation for why migrations are initiated at age 2.
5. Our study supports the hypothesis that fast-growing individuals shift their niche earlier and at a smaller body size than slower-growing individuals because they maintain higher metabolic rates and are energetically constrained at a younger age by limited food resources than slow growers.     

Behavioural versus physiological mediation of life history under predation risk

Predator-generated variation in prey energy intake remains the dominant explanation of adaptive response to predation risk in prey life history, morphology and physiology across a wide range of taxa. This "behavioural hypothesis" suggest that chemical or visual signals of predation risk reduce prey energy intake leading to a life history characterized by a small size and late age at maturity. However, size-selective predation can induce either smaller size-early age or large size-late age life history. The alternative "physiological hypothesis" suggests that size-selective cues decouple the relationship between energy and life history, acting instead directly on development. Here we use a series of experiments in a fish-daphnid predator-prey system to ask whether size-selective predator cues induce a physiological mediation of development, overshadowing behaviourally based changes in food intake. We found fish chemical cues reduce the net energy intake in Daphnia magna, suggesting a behaviourally mediated reduction in energy. Experimental manipulation of food levels show further that reductions in food lead to later but smaller size at maturity. However, in line with the physiological hypothesis, we show that D. magna matures earlier and at a smaller size when exposed to fish predation cues. Furthermore, our data shows that they do this by increasing their development rate (earlier maturity) for a given growth rate, resulting in a smaller size at maturity. Our data, from a classic size-selective predation system, indicate that predator-induced changes in this system are driven by physiological mediation of development rather than behavioural mediation of energy intake.     

Size selective feeding and its limitations for the black rockfish, Sebastes inermis, in a demersal fish assemblage of Onagawa Bay, northeastern Japan

The size selective feeding of the black rockfish, Sebastes inermis, in a demersal fish assemblage of Onagawa Bay, located in northeastern Japan, was studied using laboratory experiments and stomach content analysis of the dominant demersal fish coexisting with the black rockfish in Onagawa Bay. The preferred food size of the black rockfish was determined in the laboratory experiments. The average food size taken by the black rockfish (150–227 mm long) under natural conditions was much smaller than that taken by fish in laboratory experiments (148–158 mm sized). The most preferred size of food taken by fish in laboratory experiments was rarely found under natural conditions. Analysis of the competition coefficients of fish species in the fish assemblage based on stomach content analysis suggested that the degrees to which the preferred food size was found depended upon the foods availability, by competition with other fish species and by the feeding ability of the fish. Intra- and inter-specific competition in a fish assemblage might be limiting factors on the food size selection of fish under natural conditions.     

River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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The influence of food distribution upon the development of aggressive and competitive behaviour in juvenile chum salmon, Oncorhynchus keta

Groups of juvenile chum salmon were reared on food that was either dispersed throughout, or localized in one area of, the rearing tank. Groups receiving localized food displayed more aggression than those receiving dispersed food. This led to differences in growth, with fish reared on localized food having greater individual growth variability, i.e. growth depensation. However, after several months of rearing in these different feeding/social environments, fish reared on dispersed food were just as aggressive when first exposed to localized food as were fish reared on localized food. Furthermore, in competitive contests between fish of the two rearing histories, those reared on dispersed food were just as likely to become dominant as those reared on localized food. These results suggest that the behavioural development of aggressiveness is not amenable to alteration by manipulation of food distribution.     

Social dominance, prior residence and the acquisition of profitable feeding sites in juvenile Atlantic salmon

Twelve groups of 10 Atlantic salmon Salmo salar parr were screened for dominance in small tanks with a single localized food source using a serial removal method and the top, middle and bottom two fish in each group were classified as high, medium and low social status, respectively. These 72 ranked fish were weighed and formed into four groups or waves, each consisting of six fish of each status category. The fish were given dye marks according to status and wave and were then introduced into an artificial stream in four waves over a 12-day period. Patterns of settlement/emigration were recorded over a 2-month period, at the end of which the fish were removed from the stream, identified and reweighed, and daily specific growth rates calculated. Fifty-one fish established feeding stations in the flume, the remainder emigrating via a downstream trap. The average growth rate over the 2-month period was 0·87% day⁻¹. Probability of settlement and growth rates following settlement were significantly higher in fish from the earlier waves. In contrast, salmon that were dominant in the initial trial were less likely than subordinate fish to settle in the experimental stream and showed lower postsettlement growth rates. The possibility that fish have individually consistent behavioural profiles that promote success in different competitive environments is discussed with reference to published literature for other groups. A small-scale trial at the end of the main experiment confirmed the prior residence effect and demonstrated that 1 day was sufficient for such an effect to be evident.     

A test of the model of the economic defendability of a resource and territoriality using youngEtroplus maculatus andPelmatochromis subocellatus kribensis

YoungEtroplus maculatus andPelmatochromis subocellatus kribensis were reared on diets of low, medium, or high quantities of food and the ontogeny of behavior documented until fish matured. Data were categorized into: (1) from the beginning of the experiment to the midpoint (100 days forE. maculatus and 118 days forP. s. kribensis); and (2) from the midpoint to maturity. For both species the tendency to be territorial, as deduced by the numbers of territories defended, or number of charges or rams performed, was explained by a unimodal relationship between territorial tendency and food available per gram of fish. Young fish receiving low quantities of food were highly territorial, but as they grew and food available per gram of fish decreased, they gave up territorial behavior. Young fish receiving high quantities of food were not territorial initially, but as they grew became increasingly territorial. Fish receiving intermediate quantities of food always defended territories. Lateral displays were performed most often by fish receiving intermediate quantities of food apparently because they were ambivalent being close to either the lower or upper thresholds for territoriality.