The development of shoaling behaviour in the European minnow, Phoxinus phoxinus

The effects of development and disturbance on schooling and shoaling were observed in laboratory-raised minnows. Both behaviours were present at the onset of free swimming. Shoaling increased to become the most important behaviour by 3 weeks after emergence from gravel, while the main increase in schooling behaviour occurred at 4 weeks. These changes were accompanied by rises in elective group size and in the number of following responses, and by a decrease in nearest neighbour distance. The increases in social behaviour were observed at an earlier stage in the same minnows threatened by a standard net chase. The main increase in schooling in disturbed minnows took place between 3 and 4 weeks after emergence; after this time schooling became the dominant behaviour. Disturbed fish were also more likely to follow another minnow during an encounter.     

The genetic structure and microdistribution of shoals of Phoxinus phoxinus, the European minnow

Efficient foraging and a reduction in predation risk have been proposed as reasons for shoal formation. Some behaviours in cyprinid shoals are at first sight altruistic (e.g. predator inspection behaviour, reactions to alarm substance), such that kin selection may have been involved in their evolution. If shoaling behaviour does evolve through kin selection, then genetic differentiation is expected to be greater between shoals than within shoals. Such a hypothesis was tested here by examining shoal integrity and the relatedness of individuals within and between shoals in the European minnow, Phoxinus phoxinus , using nuclear and mitochondrial DNA markers. The breeding structure of 13 minnow shoals collected from Dorset and North Wales, U.K., was examined using allozymes. Genetic affinity within and between shoals was tested using mitochondrial DNA and multi-locus DNA fingerprinting. Shoals consisted of a random assortment of allozyme genotypes, shoal members did not share the same maternal mtDNA lineages and DNA fingerprint profiles were as varied within shoals as between them. The data indicate that it is unlikely that kin selection occurs in P. phoxinus and there is no apparent relationship between shoaling behaviour and genotype distribution in this species.     

Flow velocity underpins microhabitat selection by gobies of the Australian Wet Tropics

1. Water flow is a critical driver of aquatic ecosystem health and function. Amid rising concerns over changing flow regimes, there is an urgent need to understand the functional mechanisms by which flow influences patterns of freshwater biodiversity. 2. We explored the functional link between flow velocity and microhabitat specialisation in a speciose group of freshwater gobies (comprising over half the total fish species richness) within insular streams of the Australian Wet Tropics under base flow conditions. We addressed two particular questions: (i) What is the relative selectivity of species towards streambed composition and water flow velocity? and (ii) Can patterns of microhabitat occupation be explained by differences in intrinsic flow performance among species? To answer these questions, we combined visual field observations of microhabitat use with flow tank assessments of flow speed performance. 3. Tropical freshwater gobies displayed strong specificity towards flow velocities, while being relatively non‐selective towards streambed composition. At opposite extremes of the spectrum, we found Sicyopterus lagocephalus occupying high‐flow (>1.0 ms^?1) microhabitats while Redigobius bikolanus selected slower‐flow (<0.05 ms^?1) areas. These patterns of microhabitat flow specificity were largely explained by the different abilities of species to swim and/or cling to the substratum under these different flow settings. 4. Our findings confirm suggestions that predictable base flows in tropical streams support habitat specialists, which include one species capable of occupying areas of extremely high flow that very few other fishes can withstand. 5. The functional link between flow and gobioid fish distribution patterns could occur throughout tropical streams of the Indo‐Pacific and Caribbean as a widespread phenomenon that may help inform stream flow management guidelines to maintain this substantial component of tropical freshwater biodiversity around the globe.     

The genetic and environmental basis of adaptive differences in shoaling behaviour among populations of Trinidadian guppies,Poecilia reticulata

The degree of plasticity an individual expresses when moving into a new environment is likely to influence the probability of colonization and potential for subsequent evolution. Yet few empirical examples exist where the ancestral and derived conditions suggest a role for plasticity in adaptive genetic divergence of populations. Here we explore the genetic and plastic components of shoaling behaviour in two pairs of populations of Poecilia reticulata (Trinidadian guppies). We contrast shoaling behaviour of guppies derived from high‐ and low‐predation populations from two separate drainages by measuring the shoaling response of second generation laboratory‐reared individuals in the presence and absence of predator induced alarm pheromones. We find persistent differences in mean shoaling cohesion that suggest a genetic basis; when measured under the same conditions high‐predation guppies form more cohesive shoals than low‐predation guppies. Both high and low‐predation guppies also exhibit plasticity in the response to alarm pheromones, by forming tighter, more cohesive shoals. These patterns suggest a conserved capacity for adaptive behavioural plasticity when moving between variable predation communities that are consistent with models of genetic accommodation.     

Isolation and the development of shoaling in two populations of the guppy

Socially reared guppies Poecilia reticulata derived from two wild populations (Upper and Lower Aripo River, Trinidad) showed a significant relationship between body size and shoaling tendency, measured as the proportion of time spent in proximity to a bottle containing conspecifics. Larger females shoaled significantly more than smaller females. Fish from the high-shoaling population (Lower Aripo) showed significantly less shoaling behaviour when reared in isolation. In contrast, low-shoaling fish (Upper Aripo) demonstrated no significant change in their shoaling behaviour in response to social isolation.     

The adenohypophysis of the flounder,Pleuronectes flesus,and the minnow,Phoxinus phoxinus

Summary The pituitary gland of the flounder, Pleuronectes flesus, showed several unusual cytological features. Between the RPD and the PPD was a zone of cells that stained purple with Alcian blue—PAS—orange G. Many of these cells were apparently degenerating. In the PPD the strands and coils of presumptive STH cells showed a tremendous variation in both size and staining properties. In the PI there were two cell types, the PAS-positive one bordering the neurohypophysis. Around the periphery of the PI was a zone of chromophobic cells, and throughout the PI were numerous intracellular and extracellular acidophil spheres.No well defined ACTH cells were found in the RPD of the minnow (Phoxinus phoxinus). The Alcian blue—PAS—orange G technique distinguishes between blue TSH cells and purple GTH cells in the RPD and PPD. GTH cells from animals collected in the winter were vacuolated. The PI contained two cell types whose staining reactions and ultrastructure were extremely variable. Intra- and extra-cellular acidophil spheres were present.I should like to thank Dr. T. Kerr of the Zoology Department, Leeds University, for his help and encouragement, and Mr. J. Dingley of the Department of Zoology, Aberystwyth, for collecting the flounders.     

The effect of hunger and cestode parasitism on the shoaling decisions of small freshwater fish

Individual minnows Phoxinus phoxinus and three-spined sticklebacks Gasterosteus aculeatus were provided with a mutually exclusive choice between joining a shoal of conspecifics and foraging alone in a maze. The shoaling decisions and foraging behaviour of individual fish were studied when the fish were satiated and after 24- and 48-h periods of food deprivation. Hunger level was found to have a significant effect on shoaling behaviour. When satiated, fish of both species spent a greater proportion of time within one body length of the shoal and spent less time out of visual contact with the shoal than after periods of food deprivation. The effect of the cestode parasite Schistocephalus solidus on the shoaling behaviour of stickleback hosts was complex. When satiated, infected fish spent less time than uninfected fish within one body length of the shoal, preferring to remain outside the shoal, yet within visual contact, although when food deprived there was no difference in the proportion of time spent by infected and uninfected fish close to the shoal. The possible ecological significance of this change in behaviour is discussed with reference to the manipulation hypothesis of host-parasite interactions.     

Accumulation and effects of aluminium in the minnow (Phoxinus phoxinus L.) at different pH levels

The accumulation and effects of 150 pg All^-1 on minnows ( Phoxinus phoxinus L.) were studied in soft water at pH 7, 6 and 5. The fish were kept in a flow-through water system for up to 48 days. Addition of aluminium to the water resulted in poor appetite, passivity and a tendency for the fish to gather as far as possible from the aluminium inlet. The mortality was high at pH 5 and also after addition of aluminium at pH 6. Severe lesions were noted in the gills and olfactory organs after exposure to aluminium. The concentration of aluminium in the gills, liver and kidney was increased at low pH. A large proportion of the aluminium was located on the surface of the gill epithelium. The recovery capacity was pronounced when the aluminium exposure was terminated and the pH was increased from 5 to 7. No further mortality occurred and gill and olfactory organ structure returned to near normality in about 36 days.     

Effects of ration, temperature and body size on the body composition, energy content and condition of the minnow, Phoxinus phoxinus (L.)

The effects of ration and temperature on the body composition, energy content and indices of condition of the minnow. Phoxinus phoxinus (L.) (Cyprinidae). were studied experimentally. Five ration levels over the range starvation to ad libitum and five temperatures over the range 5–15°C were used. The fish weighed 1-5-4 g. At each temperature, the relative dry matter content, lipid content and energy content increased with increased ration. The response of the protein content to ration was temperature-dependent. Temperature had a significant effect on body composition and energy content, but this effect was irregular. The Fulton condition factor was less sensitive to ration than the wet or dry liver-somatic index. Temperature had a significant effect on the condition factor and liver indices. Significant, positive correlations were found between growth rate, lipid content and the indices of condition. The liver indices were better predictors of growth rate than the condition factor or lipid content and may provide useful information on the physiological condition offish in field studies.     

Shoaling preferences of two common killifish (Fundulus heteroclitus and F. diaphanus) in the laboratory and in the field: A new analysis of heterospecific shoaling

Heterospecific grouping behavior of mummichogs (Fundulus heteroclitus) and banded killifish (Fundulus diaphanus) was analyzed in the laboratory and in a freshwater tidal marsh in Cremona, Maryland. Several parameters of wild, intact shoals were measured, including species composition, body length, parasite load, gender, and any physical abnormalities. Fish collected were used for laboratory analysis of shoaling preferences. When size was equal, banded killifish and mummichogs preferred conspecific shoals to heterospecific shoals, consisting of mummichogs, banded killifish, and sheepshead minnows (Cyprinodon variegates). Shoal collection from the field resulted in mixed species shoals with individuals predominantly unaffected by parasites or other physical abnormalities. Size appeared to be a sorting mechanism. A temporal shift in lengths was evident. Initial shoals caught contained significantly smaller fish compared to the final shoals caught. Results are compared with previous studies on heterospecific shoaling in killifish and new characteristics of heterospecific shoals inhabiting freshwater tidal marshes are discussed.     

Grouping fails to affect the growth and energy budget of a cyprinid, Phoxinus phoxinus (L.)

The effects of grouping on the growth and some components of the energy budget of the minnow, Phoxinus phoxinus (L.), were tested in the laboratory. No effects of grouping were detected on absorption efficiency, the proportion of food energy lost in nitrogenous excretion, maximum rate of food consumption and conversion efficiency. Grouping had no significant effect on either specific growth rate or the dry matter content of individual fish.     

Effects of two oil dispersants on phototaxis and swimming behaviour of barnacle larvae

The effects of two oil dispersants (Vecom B-1425 GL and Norchem OSD-570) mixed with diesel oil on the survival and behaviour of the stage II nauplii of the barnacle Balanus amphitrite were investigated. The24 and 48-hour LC₅₀ values for Vecom B-1425 GL:diesel mixture were 514 and 48 mg l₋₁ respectively, while respective values for Norchem OSD-570:diesel mixture were 505 and 71 mg l₋₁. Under sublethal concentrations, increased levels of the dispersant:diesel mixtures caused a reduction in phototactic responses. Balanus amphitrite nauplii failed to exhibit phototactic responses when exposed to Vecom B-1425GL:diesel mixtures of 400 mg l₋₁ and higher for 24 hours. A longer exposure time of 48 hours further reduced the Lowest Observable Effect Concentrations (LOECs) to 60 mg l₋₁. The LOECs for Norchem OSD-570:diesel mixtures under exposure periods of 24 and 48 hours were 400and 80 mgl₋₁ respectively. The curvilinear velocities (VCL) and straight-line velocities (VSL) of the stage II nauplii ranged from 0.7–1.1and 0.2-0.4 mms₋₁ respectively. Increased concentrations of dispersant:diesel mixtures caused a significant change in the curvilinear and straight-line velocities. Both oil dispersants, dispersant:diesel mixtures of 20 to 40 mgl₋₁ caused significant increases in VCL, but no significant change in VSL. Dispersant:diesel mixtures of 100 mg l₋₁ and higher resulted in a reduction in VSL for both dispersants. This revised version was published online in August 2006 with corrections to the Cover Date.     

The effects of spinal nerve section on responses of melanophores in the minnow, Phoxinus phoxinus(L.)

A continuous observation apparatus was used to study the responses of Phoxinus phoxinus melanophores to illuminated black/white backgrounds and their reversal. The fish. Although confined, showed maximum melanosome dispersion (MI 5) and maximum melanosome aggregation (MI 1) when exposed to illuminated black and white backgrounds respectively. Melanophores affected by spinal nerve section showed full melanosome dispersion and the affected area appeared as a black band. The affected melanophores marginally and gradually aggregated their melanosomes if the fish was exposed to an illuminated white background for about a week. The responses of these melanophores to illuminated black and white backgrounds and their reversal indicates that the dispersal of their melanosomes in response to a black background is much faster than their aggregation in response to a white background. It is concluded that an active mechanism is involved and possible factors controlling it are discussed.     

Effects of impoundment on populations of bullhead Cottus gobio L. and minnow, Phoxinus phoxinus (L.), in the basin of Cow Green Reservoir

The distribution, growth and reproduction of bullhead, Cottus gobio , L., and minnow, Phoxinus phoxinus (L.), were studied at Cow Green Reservoir, Upper Teesdale, for 10 years following impoundment in 1970. Comparisons were made with pre-impoundment results. The length-for-age of bullheads increased after impoundment. The length-for-age of minnows decreased. In winter both species were confined mainly to deep water or to the stone facing of the earth dam. In summer they were found throughout the reservoir though bullheads were scarce in the littoral zone. Female bullheads became sexually mature at an earlier age (84% at age-group I) than before impoundment (33% at age-group I) but no change was apparent amongst males (67–62% in age-group I). The post-impoundment sex ratio did not differ from 1:1 and individual fecundity did not change after impoundment. There was a significant reduction in individual fecundity of the minnow. The mean instantaneous rate of mortality ( Z ) for bullhead of age-groups II to IV was 0.96 ± 0.34 year⁻¹, compared with 0.85 ± 0.29 year in the Tees before impoundment.     

Effects of discharge and local density on the growth of juvenile Atlantic salmon Salmo salar

The study explored the combined effects of density, physical habitat and different discharge levels on the growth of juvenile Atlantic salmon Salmo salar in artificial streams, by manipulating flow during both summer and winter conditions. Growth was high during all four summer trials and increased linearly with discharge and mean velocity. Differences in fish densities (fish m^?3) due to differences in stream volume explained a similar proportion of the variation in mean growth among discharge treatments. Within streams, the fish aggregated in areas of larger sediment size, where shelters were probably abundant, while growth decreased with increasing densities. Fish appeared to favour the availability of shelter over maximization of growth. Mean growth was negative during all winter trials and did not vary among discharge treatments. These results suggest that increased fish densities are a major cause of reduced summer growth at low discharge, and that habitat‐mediated density differences explain the majority of the growth variation across habitat conditions both during summer and winter.     

Effects of parasites on fish behaviour: a review and evolutionary perspective

Fish serve as hosts to a range of parasites that are taxonomically diverse and that exhibit a wide variety of life cycle strategies. Whereas many of these parasites are passed directly between ultimate hosts, others need to navigate through a series of intermediate hosts before reaching a host in (or on) which they can attain sexual maturity. The realisation that parasites need not have evolved to minimise their impact on hosts to be successful, and in many cases may even have a requirement for their hosts to be eaten by specific predators to ensure transmission, has renewed interest in the evolutionary basis of infection-associated host behaviour. Fishes have proved popular models for the experimental examination of such hypotheses, and parasitic infections have been demonstrated to have consequences for almost every aspect of fish behaviour. Despite a scarcity of knowledge regarding the mechanistic basis of such behaviour changes in most cases, and an even lower understanding of their ecological consequences, there can be little doubt that infection-associated behaviour changes have the potential to impact severely on the ecology of infected fishes. Changes in foraging efficiency, time budget, habitat selection, competitive ability, predator-prey relationships, swimming performance and sexual behaviour and mate choice have all been associated with – and in some cases been shown to be a result of – parasite infections, and are reviewed here in some detail. Since the behavioural consequences of infections are exposed to evolutionary selection pressures in the same way as are other phenotypic traits, few behavioural changes will be evolutionarily neutral and host behaviour changes that facilitate transmission should be expected. Despite this expectation, we have found little conclusive evidence for the Parasite Increased Trophic Transmission (PITT) hypothesis in fishes, though recent studies suggest it is likely to be an important mechanism. Additionally, since the fitness consequences of the many behavioural changes described have rarely been quantified, their evolutionary and ecological significance is effectively unknown.Potential hosts may also change their behaviour in the presence of infective parasite stages, if they adopt tactics to reduce exposure risk. Such `behavioural resistance', which may take the form of habitat avoidance, prey selectivity or avoidance of infected individuals, can be viewed as behavioural change associated with the threat of being parasitised, and so is included here. Actually harbouring infections may also stimulate fishes to perform certain types of simple or complex behaviours aimed at removing parasites, such as substrate scraping or the visitation of cleaning stations, although the efficacy of the latter as a parasite removal strategy is currently subject to a good deal of debate.The effects parasites have on shoaling behaviour of host fish have attracted a good deal of attention from researchers, and we have provided a case study to summarise the current state of knowledge. Parasites have been shown to affect most of the antipredator effects of shoaling (such as vigilance, co-ordinated evasion and predator confusion) and can also impair an individual's foraging ability. It therefore seems unsurprising that, in a number of species avoidance of parasitised individuals has evolved which may explain the occurrence of parasite-assorted shoals in the field. Parasitised fish are found more often in peripheral shoal positions and show a reduced tendency for shoaling in some fish species. Given the array of host behaviours that may be changed, the fitness consequences of shoal membership for parasitised hosts and their parasites are not always easy to predict, yet an understanding of these is important before we can make predictions regarding the ecological impact of infections on host fish populations.Clearly, there remain many gaps in our knowledge regarding the effects of parasites on the behaviour of host fish. We believe that a much greater understanding of the importance of infection-associated behaviour changes in fish could be gained from high quality research in comparatively few areas. We have completed our review by highlighting the key research topics that we believe should attract new research in this field.     

CSF-contacting and other somatostatin-immunoreactive neurons in the brains of Anguilla anguilla,Phoxinus phoxinus,and Salmo gairdneri (Teleostei)

Summary A system of somatostatin-immunoreactive neurons was demonstrated in the brains of the eel, Anguilla anguilla, the European minnow, Phoxinus phoxinus, and the rainbow trout, Salmo gairdneri, by means of the light-microscopic indirect immunoperoxidase technique.In the anterior periventricular nucleus, somatostatin-immunoreactive cerebrospinal fluid (CSF)-contacting neurons display intensely stained intraventricular dendritic protrusions, perikarya, and axonal processes. The latter taper into a somatostatin-immunoreactive fiber plexus extending to the infundibulum, the proximal neurohypophysis, and the lateral and mammillary recesses.In addition, somatostatin-immunoreactive neurons were demonstrated in the magnocellular preoptic, entopeduncular and dorsolateral thalamic nuclei, further in the pretectal area and the ventrolateral tegmentum. Somatostatin-immunoreactive fiber bundles project via the stria medullaris toward the habenular nucleus; they also course in the dorsomedial-ventrolateral direction at the level of the pretectal-tegmental area, and within the ventral and dorsal tegmentum.The presence of somatostatin in a variety of different neurons of the teleost brain is discussed in connection with their tentative inhibitory function. The CSF-contacting neurons of the anterior periventricular nucleus are supposed to function as sensors that pass information from the CSF to the somatostatin system of the hypothalamus and/or other components of the neuroendocrine apparatus.Dedicated to Professor Helmut Leonhardt, Kiel, on the occasion of his 65^th birthdayThis investigation was supported by grants from the Deutsche Forschungsgemeinschaft to A.O. (Ok 1/25-3) and H.W.K. (Ko 758/2-2)On leave of absence from the second Department of Anatomy, Semmelweis OTE, Budapest, Hungary     

Bioenergetics of growth of a cyprinid, Phoxinus phoxinus (L.): the effect of ration and temperature on growth rate and efficiency

Growth of minnows, Phoxinus phoxinus , weighing 1-5.5 g was studied experimentally at five ration levels from starvation to ad libitum and four temperatures ranging from 5 to 15°C. The relationship between specific growth rate (SGR) and ration was a decelerating curve. SGR at maximum rations increased with increased temperature, but at restricted rations it decreased with increased temperature. Predictive models for the specific growth rates were developed using multiple regression. Maintenance rations and optimum rations both increased with increased temperature. Maintenance rations were less sensitive to temperature than optimum rations and mostly lay between 1 and 2% of body weight per day. Conversion efficiencies increased with increased ration from zero value at the maintenance ration to a peak at the optimum ration, then decreased with further increases in ration. At a given restricted ration level, conversion efficiencies generally decreased with increased temperature. At maximum rations, conversion efficiencies were relatively insensitive to temperature. Growth in wet weight, dry weight and energy content showed similar responses to ration, temperature and body weight.     

Appetite and growth compensation in the European minnow,Phoxinus phoxinus (Cyprinidae), following short periods of food restriction

Synopsis Changes in the daily appetite and weekly growth rates of individual adult minnows,Phoxinus phoxinus, on ad libitum rations were recorded before and after they had experienced 4 or 16 days of food restriction. Feeding levels during the restriction periods were either starvation or a maintenance ration. The latter was estimated from a previously determined regression model. Water temperature was 15°C and the photoperiod 9L15D in all experiments. The mean weight of fish used ranged from 1.06 to 2.15 g. The 4 day restriction had no detectable effects on appetite or growth. After the 16 day restriction, the minnows showed hyperphagia and had increased specific growth rates and growth efficiencies compared with control fish. The compensatory increases in appetite and growth were not sustained and within three weeks had declined to levels not significantly different from those of the control fish. At the end of the experiments, there were no significant differences between the mean weights or cumulative food consumption of the restricted and control groups. The results suggest that adult minnows regulate their appetite and growth rate in relation to their previous nutritional history.