The Effect of Hunger on Shoal Choice in Golden Shiners (Pisces: Cyprinidae,Notemigonus crysoleucas)

When an animal has a choice of joining one group over another, its decision may depend on its relative vulnerabilities to predation and starvation. For example, a well-fed animal may choose a large group of individuals with body size matching its own because this gives good protection against predators, but a hungry animal may prefer smaller groups made up of smaller individuals because this decreases food competition. To test this idea, a choice between various shoals was given to golden shiners, Notemigonus crysoleucas, that were either well fed or deprived of food for 48 h. In a choice of 10 vs. 3 shoalmates, both well-fed and hungry shiners spent more time near the shoal of 10. In a choice of 20 vs. 3 shoalmates, both well-fed and hungry shiners again preferred the larger shoal, but in one replicate this preference was significantly weaker in the hungry fish. This reduced preference did not appear to be an artefact of increased mobility by hungry fish searching for food. In a choice between shoals of small vs. large conspecifics, small well-fed shiners, small hungry shiners, and large well-fed shiners preferred shoalmates with body size matching their own, but large hungry shiners preferred smaller individuals. These results are consistent with the hypothesis that hungry fish sacrifice safety from predation in their shoaling behaviour (by avoiding larger groups to a certain extent and by risking the oddity effect) so as to decrease food competition.     

Can a minority of informed leaders determine the foraging movements of a fish shoal?

There is no information on whether the daily foraging movements of fish shoals are the result of chance, the collective will of all shoalmates, or the leadership of a few individuals. This study tested the latter possibility. Shoals of 12 golden shiners, Notemigonus crysoleucas, were trained to expect food around midday in one of the brightly lit corners of their tank. They displayed daily food-anticipatory activity by leaving the shady area of their tank and spending more and more time in the food corner up to the normal time of feeding. Past this normal time they remained in the shade, even on test days when no food was delivered. Most of these experienced individuals were then replaced by na?ve ones. The resulting ratio of experienced:na?ve fish could be 5:7, 3:9 or 1:11. On their own, na?ve individuals would normally spend the whole day in the shade, but in all tests the experienced individual(s) were able to entrain these more numerous na?ve fish out of the shade and into the brightly lit food corner at the right time of day. Entrainment was stronger in the 5:7 than in the 1:11 experiment. The test shoals never split up and were always led by the same fish, presumably the experienced individuals. These results indicate that in a strongly gregarious species, such as the golden shiner, a minority of informed individuals can lead a shoal to food, either through social facilitation of foraging movements or by eliciting following behaviour. Copyright 2000 The Association for the Study of Animal Behaviour.     

The influence of hunger on shoal size choice by three-spined sticklebacks, Gasterosteus aculeatus

Single sticklebacks ( Gasterosteus aculeatus ) were presented with a choice between (a) a shoal and an empty compartment, and (b) two different sized shoals of conspecifics. The choice of hungry test fish (food-deprived for 24 h and 72 h, respectively) was compared to that of fish which were fed to satiation once a day. It was hypothesized that due to the increased level of competition for food in large shoals, hungry fish would spend less time with the larger of two shoals than their well-fed conspecifics. The results showed that there was indeed a significant trend for test fish to spend a decreasing amount of time near the compartment with the larger number of fish with increasing duration of food deprivation.     

Shoal Choice in the Banded Killifish (Fundulus diaphanus,Teleostei, Cyprinodontidae): Effects of Predation Risk,Fish Size,Species Composition and Size of Shoals

Banded killifish (Fundulus diaphanus) were presented individually with a choice of shoaling with either of two stimulus shoals which differed in shoal size, species composition, and fish body size, before and after a simulated avian predator attack. When threatened, test fish preferred to shoal with the larger of two conspecific shoals, but only if members of both stimulus shoals were of the same size class as the test fish. Otherwise, they preferred to shoal with similarly sized fish irrespective of shoal size; threat of predation increased the magnitude of this preference. Furthermore, test fish preferred a shoal of similarly sized shiners (Notemigonus crysoleucas) over larger killifish, when shoal sizes were identical. This indicates that body size plays a key role in shoal choice, overriding the effects of shoal size and species preference. Notwithstanding the above, shoal choice was affected by predator threat only when differences between shoal size or body size of stimulus fish were large.     

The effect of shoal size on patterns of body colour segregation in mollies

Individual white and black mollies Poecilia latipinna spent significantly more time near the larger group when given the choice between two shoals of similar colouration to themselves. When given the choice between a large and a small shoal of dissimilar colouration to themselves, black test fish spent significantly more time with the larger shoal while white test fish showed no preference for either group. Both white and black mollies chose the smaller of two shoals when given the choice between a large dissimilarly coloured shoal and a small similarly coloured shoal. The results indicate that mollies actively discriminate between shoals on the basis of both body colour and shoal size. However, body colour segregation appears to have a stronger influence on shoal choice.     

Background colouration influences body colour segregation in mollies

When given a choice of swimming against a black background near an empty black end compartment or swimming against a white background near an empty white compartment, black morph mollies Poecilia latipinna spent significantly more time against the black background. The presence of a shoal (either white or black mollies) in the black compartment increased the preference of the test fish for the black background. However, black molly test fish spent significantly less time against a black background if a black shoal was present in the white compartment than in assays in which a white shoal was in the white compartment. In assays where test fish were given the choice between two black shoals, test fish spent significantly more time near the black shoal on the black background. Similarly, when given the choice between two white shoals, test fish spent significantly more time against the black background. The results suggest that the presence of cryptic backgrounds alter shoalmate choices in fish that assort according to body colouration.     

Shoaling in juvenile guppies: the effects of body size and shoal size

While factors affecting shoal mate choice have been examined extensively in adult guppies (Poecilia reticulata), few studies have focused on the shoaling behavior of juveniles. In this study, juvenile guppies were tested for their ability to shoal as well as their response to shoal mates of different body size and to shoals with different numbers of individuals. In dichotomous choice tests, 10-day-old guppies (mean body length=8.83 mm), 30-day-old guppies (13.17 mm) and 50-day-old guppies (18.6mm) were given the opportunity to swim near shoals of five fish or an empty chamber. In most cases, the juvenile fish demonstrated shoaling behavior, swimming near a group of fish rather than an empty chamber, regardless of the age of the stimulus shoal. When presented with two shoals, one of similar age and body size and one of dissimilar age and body size, only the 50-day-old guppies showed a significant preference for the age-matched shoal. Similarly, when choosing between a large shoal and a small shoal, only the 50-day-old guppies spent significantly more time near the larger shoal. Thus, while juveniles at each age shoaled, only 50-day-old fish demonstrated the shoal mate discrimination seen in adult fish.     

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.     

Effects of fasting on collective movement and fission–fusion dynamics in both homogeneous and heterogeneous shoals of a group-living cyprinid fish species

The present study aimed to reveal the effect of fasting (21 days) on collective movement and interaction dynamics in both homogeneous (eight members fed a commercial diet or deprived of food) and heterogeneous (four fed + four starved members) shoals of juvenile qingbo (Spinibarbus sinensis). The authors of this study measured the shoaling behaviour in both a commonly used rectangular open arena with no spatial complexity and a radial arm maze. When measured in the open arena, the starved shoals had a faster swimming speed and acceleration rate and a longer interindividual distance than the fed shoals, possibly because of the elevated foraging motivation. Nonetheless, the values of the heterogeneous groups were similar to those of the fed groups. Furthermore, in contrast to the fish in homogeneous shoals, the starved fish in heterogeneous shoals showed a slower acceleration rate and speed than fed members in heterogeneous shoals. These results, combined with the relationships of variables at the among- and within-shoal levels, suggested that starved fish limited their motion in heterogeneous shoals to maintain group cohesion but that the fed fish contributed more to maintaining shoal structure, possibly because of the higher energy expenditure required for movement changes. When monitored in a radial arm maze, starved shoals showed more fission–fusion episodes without sacrificing group cohesion, as they adaptively adjusted the frequency and duration of each majority choice. The among-shoal variation revealed that the heterogeneous groups showed less variation in the open arena but more variation in the radius maze than did the homogeneous groups. This difference might arise because dominant members have opposite effects on shoal behaviour and consensus decisions. In conclusion, the present study showed opposite effects of feeding states on collective behaviour between homogeneous and heterogeneous shoals, possibly because of the complicated interactions among members with different energy storage levels and foraging motivations. Furthermore, the heterogeneous groups showed a difference between shoal behaviour in the open area and exploration in the radial arm maze. Future studies manipulating the personality composition of starved and fed members of heterogeneous groups might yield interesting results.     

The relationship between foraging and shoal position in a mixed shoal of roach (Rutilus rutilus) and chub (Leuciscus cephalus): a field study

Feeding rates of mixed shoals of juvenile roach and chub were observed in a shallow stream near Cambridge (UK). Roach at the front of the shoal had significantly higher feeding rates than roach at the back and than chub in either front or back positions. Position in the shoal also had a significant effect on the kind of food consumed, with front roach feeding more on plankton and back roach more on bottom food. Altogether 36 fish from the stream were caught and marked. Half of these were deprived of food and the other half well-fed for 3 days in captivity. After release 36% of them joined their old shoal again. Individuals from the starved group occupied front positions significantly more often than well-fed fish, but after 2 days this difference disappeared.     

Juvenile convict cichlids shoaling decisions in relation to shoal size and age

Many animals form groups and socialize in response to evolutionary pressures such as predation, food availability, and mate acquisition. Evidence of social choice based on various phenotypic characters (Group Phenotypic Composition [GPC]) has been observed in several animal species. In addition to the physical characteristics of the social group, it is also interesting to consider how decisions of who to socialize with might be expected to change for an individual over time. Younger individuals with limited life experience may discriminate differently between social groups than older conspecifics who have had the opportunity to learn and who may be faced with different ecological or environmental pressures. Here, we used a traditional two‐choice design to explore the shoaling behavior of juvenile convict cichlids and determine whether the number of fish and/or the size/life stage of the individuals within a shoal influenced social choices. We found that juvenile convict cichlids spent more time shoaling with similarly sized juvenile individuals and also preferred to shoal with larger shoals, but not when shoals were comprised of adult fish. The size of the individuals in a shoal was a more influential factor than the size of the shoal itself. Size of individual juveniles was correlated with tendency to visit shoals, but was not correlated with overall time spent shoaling, regardless of shoal composition. As juveniles, convict cichlids can make discriminatory choices that are influenced by specific aspects of shoal composition.     

The influence of nutritional state on shoal choice in zebrafish, Danio rerio

We investigated the mechanisms and functions of shoal choice in relation to nutritional state in the zebrafish. Single fish that had been well fed or food deprived for 2 days were presented with a choice between two stimulus shoals. Food-deprived test fish showed a significant preference for well-fed stimulus fish over food-deprived ones whereas well-fed test fish did not exhibit any significant preference. Subsequent experiments showed that food-deprived test fish had a significantly higher foraging success in shoals consisting of well-fed individuals than in ones that comprised food-deprived fish. No difference in the locomotory behaviour of food-deprived and well-fed stimulus fish was found with respect to the proportion of time spent swimming (as opposed to being motionless), the proportion of time spent in the upper part of the test tank and the number of sharp turns. However, body weight, stomach width (measured directly behind the pectoral girdle) and ventro-dorsal height significantly decreased over a 48-h food-deprivation period. The potential use of the latter factors for the recognition of food-deprived individuals is discussed. Copyright 1999 The Association for the Study of Animal Behaviour.     

UV matters in shoaling decisions

Shoaling behaviour in fish is influenced by numerous factors, such as familiarity, kinship, group size and shoal composition. Grouping decisions are based on both olfactory and visual cues. The visual system of many vertebrates is extended into the ultraviolet (UV) wave range as in three-spined sticklebacks (Gasterosteus aculeatus, L.). We investigated whether the presence or absence of UV wavelengths has an influence on shoaling behaviour in this species. Reproductively non-active three-spined sticklebacks were given the choice between two shoals, equal in numbers of individuals, which could be seen either through a UV-transmitting [UV(+)] or a UV-blocking [UV(-)] filter. Test fish preferred to join the shoal seen under UV(+) conditions. Due to differences in quantal flux between the UV(+) and UV(-) filters used, control experiments with neutral-density optical filters were performed in order to clarify the role of luminance. Here, test fish spent significantly more time near shoals that were seen in a darker environment, suggesting a potential trade-off between UV radiation and lower brightness during shoal choice.To our knowledge, these results demonstrate for the first time that shoaling decisions are influenced by UV wavelengths.     

Influence of Parasitism on Shoal Choice in the Banded Killifish (Fundulus diaphanus,Teleostei, Cyprinodontidae)

Banded killifish, Fundulus diaphanus, were presented individually with a choice of ‘shoaling’ with either of two conspecific stimulus shoals, one consisting of fish with externally visible black spots (indicating the presence of a parasitic trematode worm, Crassiphiala bulboglossa), the other consisting of fish without such spots. Both parasitized and unparasitized test fish significantly preferred to shoal with unparasitized stimulus shoals over parasitized ones. The relative preference of unparasitized killifish for unparasitized shoals increased with increasing degree of parasite load in parasitized stimulus fish. However, no significant preference by unparasitized fish for unparasitized shoals was observed if unparasitized shoals were presented versus mixed shoals consisting of parasitized and unparasitized fish. We conclude that killifish preferred unparasitized shoals over parasitized shoals only if all fish in the respective stimulus shoals were phenotypically uniform (either with or without black spots). A final experiment showed that killifish used black spots as an indicator of parasite presence to make their shoal choice. The significance of these results for the formation of parasite-assorted shoals is discussed.     

Fish shoal composition: mechanisms and constraints

Observations were made on three fish species (banded killifish (Fundulus diaphanus), golden shiner (Notemigonus crysoleucas) and white sucker (Catostomus commersoni)) in a temperate lake (New Brunswick, Canada) in order to investigate the relationship between shoal choice behaviour of individual fishes and shoal composition. Encounters between shoals were observed to take place every 1.1 min per shoal and an encounter lasted 3.7 s on average. The duration of shoal encounters was influenced by shoal size but not by differences between shoals in either body length or species. Conversely, the outcome of shoal encounters (i.e. ences between shoals in either body length or species. Conversely, the outcome of shoal encounters (i.e. whether or not an individual changes shoal) was influenced by body length and species differences but not by shoal size. Together, these results suggest that encounter duration itself is unlikely to have an important influence on encounter outcome. The collection of ten entire fish shoals showed that they were assorted by species and body length. A simulation model demonstrated that individual shoal choice behaviour alone could account for the generation and maintenance of the observed levels of size assortedness of shoals without invoking the existence of other sorting mechanisms such as differential swimming speeds. However, the generation of species assortedness was not predicted by the model. Furthermore, our data suggest that fish density acts as a constraint on shoal choice, influencing both shoal size and composition. This work has implications for studies on information transfer and reciprocal altruism within populations.     

The effect of sex and shoal size on shoaling behaviour in Danio rerio

Male and female zebra fish Danio rerio were given choices of shoals that differed in sex and size. Male zebra fish preferred to associate with female shoals over male shoals, but had no preference when given a choice between a mixed‐sex shoal and either a male or female shoal. Female zebra fish showed no significant preference when given a choice between male and females shoals, nor between mixed‐sex shoals and either male or female shoals. When given choices between shoals of differing size, females preferred to associate with the larger shoal, whether or not they were composed of males or females. Males, however, had no preference for larger shoals over smaller shoals, whether or not they were composed of males or females. These results showed that male zebra fish were capable of distinguishing between males and females solely on the basis of visual cues. Furthermore, these results demonstrated a significant difference between the shoaling choices of male and female zebra fish, which may indicate a difference in the function of shoaling for the two sexes.     

Effects of nutritional state on the shoaling tendency of banded killifish, Fundulus diaphanus, in the field

Although group living can confer benefits to individuals in terms of reduced predation risk and enhanced foraging success, it may also be associated with costs, such as increased competition for food. The nutritional state of an individual could therefore affect its readiness to join or remain in groups. We investigated the shoaling behaviour of banded killifish by following marked individuals, whose nutritional state had been experimentally manipulated, and recording their shoaling behaviour in a field enclosure containing unmarked conspecifics. Overall, food-deprived fish spent more time alone, and therefore less time shoaling, than well-fed individuals. When shoaling, however, food-deprived fish were not found in smaller shoals than well-fed conspecifics. Furthermore, they did not show a greater latency to shoal initially. Having joined a shoal, however, food-deprived fish left shoals more frequently to be alone than fed fish. Rates of change in the membership size of shoals occupied by either well-fed or food-deprived fish did not differ. We conclude that nutritional state seems to affect an individual's decision to continue shoaling once an association has been made. This study is the first to investigate experimentally state-dependent changes in the size of social groups in fish under field conditions. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

How does hunger affect convergence on prey patches in a social forager?

Internal state, in this case hunger, is known to influence both the organisation of animal groups and the social foraging interactions that occur within them. In this study, we investigated the effects of hunger upon the time taken to locate and converge upon hidden simulated prey patches in a socially foraging fish, the threespine stickleback (Gasterosteus aculeatus). We predicted that groups of food‐deprived fish would find and recruit to prey patches faster than recently fed groups, reasoning that they might search more rapidly and be more attentive to inadvertent social information produced by other foragers. Instead we saw no difference between the two groups in the time taken to find the patches and found that in fact, once prey patches had been discovered, it was the recently fed fish that converged on them most rapidly. This finding is likely due to the fact that recently fed fish tend to organise themselves into fewer but larger subgroups, which arrived at the food patch together. Hunger has a significant impact upon the social organisation of the fish shoals, and it appears that this has a stronger effect upon the rate at which they converged upon the food patches than does internal state itself.     

Shoal composition determines foraging success in the guppy

The composition of an animal group can impact greatly on thesurvival and success of its individual members. Much recentwork has concentrated on behavioral variation within animalpopulations along the bold/shy continuum. Here, we screenedindividual guppies, Poecilia reticulata, for boldness usingan overhead fright stimulus. We created groups consisting of4 bold individuals (bold shoals), 4 shy individuals (shy shoals),or 2 bold and 2 shy individuals (mixed shoals). The performanceof these different shoal types was then tested in a novel foragingscenario. We found that both bold and mixed shoals approacheda novel feeder in less time than shy shoals. Interestingly,we found that more fish from mixed shoals fed than in eitherbold or shy shoals. We suggest that this can be explained bythe fact that nearly all the cases where one fish was followedinto the feeder by another occurred within mixed shoals andthat it was almost always a shy fish following a bold one. Theseresults suggest clear foraging benefits to shy individuals throughassociating with bold ones. Surprisingly, our results also suggestpotential foraging benefits to bold individuals through associatingwith shy individuals. This study highlights a possible mechanismby which interindividual variation in behavioral types is maintainedin a population.     

The Influence of Food Competition and Prédation Risk on Size-assortative Shoaling in Juvenile Chub (Leuciscus cephalus)

Size-assortative shoaling is a widespread phenomenon potentially linked to competition for resources among shoal members and anti-predator benefits gained from living in shoals. This paper investigates the influence of both factors on size-assortative shoaling in chub (Leuciscus cephalus). In a first experiment food competition was studied by manipulating the proportions of small and large chub in shoals of 20 fish. Large fish were generally more competitive than small ones. However, the competitive effect of large fish was strongly dependent on their number in the shoal. This has important consequences for the theory of competition because it makes it difficult to predict stable combinations of different phenotypes when foragers divide between food patches. Despite the strong effects of food competition no significant trend for size-assortativeness was observed in free-swimming shoals when undisturbed. This changed after the shoal had been frightened by introducing Schreckstoff (an alarm substance) to the test tank. Large and small fish still remained in the same shoal but showed a strong preference to be close to neighbours of their own size class. Large fish were observed mainly in the centre of the shoal and small fish were found more in the periphery. It is unclear whether these position differences are due to aggressive behaviour of large fish towards small fish which excludes them from the potentially safe centre of the shoal. These results indicate that prédation plays a more important role than food competition for size-assortative shoaling in chub.