Alcohol and the prior residence effect in male convict cichlids (Cichlasoma nigrofasciatum)

A series of experiments were undertaken to demonstrate the dominance advantage of a territorial resident over an intruder (the prior residence effect) in the adult male Convict Cichlid (Cichlasoma nigrofasciatum) and to evaluate the effects of alcohol (ethanol) on this phenomenon. After successfully demonstrating the prior residence effect, additional experiments were conducted in which the resident or intruder was given one of three different doses of ethanol (0.15%, 0.20%, 0.35%). These doses have previously been shown to reduce or increase intraspecific aggression in this species Over the entire dosage range, drugging the intruder produced no changes in the dominance advantage of the resident. However, drugging the resident at the higher doses (0.20%, 0.35%) resulted in loss of the resident's dominance advantage. These effects were not due to impaired locomotor behavior of the drugged fish, as no abnormal motor behaviors were noted. In general, as reported in previous studies, the resident fish, whether drugged or not, typically attacked first. Also as has been previously shown, the fish that attacked (bit) first typically showed eventual dominance. The present results show that the dominance advantage of the territorial resident is not altered by giving an intruder a hypo- or hyperaggression-producing dose of alcohol. Changes in the dominance advantage occur only as a function of drugging the resident, thereby providing evidence that the behavior of the territorial resident is either directly or indirectly (reaction of the intruder to these changes) responsible for the prior residence effect. The present studies also provide some pharmacological approaches for determining the variables important in a laboratory analog of a naturally occurring instance of intraspecific aggression.     

Male–male combats in a polymorphic lizard: residency and size,but not color,affect fighting rules and contest outcome

Theoretical models predict that the outcome of dyadic agonistic encounters between males is influenced by resource‐holding potential, resource value, and intrinsic aggressiveness of contestants. Moreover, in territorial disputes residents enjoy a further obvious competitive advantage from the residency itself, owing to the intimate familiarity with their territory. Costs of physical combats are, however, dramatically high in many instances. Thus, signals reliably reflecting fighting ability of the opponents could easily evolve in order to reduce these costs. For example, variation in color morph in polymorphic species has been associated with dominance in several case studies. In this study, we staged asymmetric resident‐intruder encounters in males of the common wall lizard Podarcis muralis, a species showing three discrete morphs (white, yellow, and red) to investigate the effects of asymmetries in color morph, residency, and size between contestants on the outcome of territorial contests. We collected aggression data by presenting each resident male with three intruders of different color morph, in three consecutive tests conducted in different days, and videotaping their interactions. The results showed that simple rules such as residency and body size differences could determine the outcome of agonistic interactions: residents were more aggressive than intruders, and larger males were competitively superior to smaller males. However, we did not find any effect of color on male aggression or fighting success, suggesting that color polymorphism in this species is not a signal of status or fighting ability in intermale conflicts. Aggr. Behav. 35:274–283, 2009. © 2009 Wiley‐Liss, Inc.     

Effect of residence and size asymmetries upon the agonistic interactions between juvenile white‐seabream (Diplodus sargus cadenati de la Paz,Bauchot and Daget, 1974)

The behavioural patterns of juvenile white‐seabream suggest that asymmetry in residence is an important factor governing the outcome of contest between individuals of this species. Asymmetries due to resident status had strong effects on agonistic behaviour, with asymmetries in body sizes producing weaker effects. Resident fish won all the combats against intruders of lower or similar length. However, when the intruder was larger than the residents (higher than 5% in length and 20% in weight), the percentage of combats won by the residents decreased to 85.7%. There was a significant correlation between fight intensity and size asymmetry in favour of the resident fish. The resident fish was more aggressive and persistent in attacks, and contests were more intense when the size of the intruder was greater. During agonistic interactions, the frontal attacks and lateral displays were more frequent when the intruders were similar or larger than residents. Attacks to the flanks and chases were more frequent in pairs where the intruders were smaller. Aggr. Behav. 25:297–303, 1999. © 1999 Wiley‐Liss, Inc.     

The influence of chemical cues on the territorial prior residence effect in convict cichlids (cichlasoma nigrofasciatum Günther)

The importance of accumulated chemical cues on the occurrence of the territorial prior residence effect was investigated in two experiments using adult convict cichlids (Cichlasonul nigrofasciatum). In the first experiment, pairs of fish, matched on a number of criteria, were randomly assigned to three independent treatment conditions in which (1) the territorial marker (pot or clump of plants) was removed just prior to a direct encounter in the aquarium of the resident (pair member in whose aquarium the encounter took place) with accumulated chemical cues remaining intact; (2) the marker was removed, as well as the chemical cues; or (3) the marker was present, but the chemical cues were removed. Results showed that a significant prior residence effect occurred when the marker was present, but did not occur in either condition in which the marker was removed. A second experiment, in which the marker was removed and the gravel substrate altered, also did not reveal a signficant prior residence effect. Apparently, the territorial resident recognizes his territory by using certain visual cues, with accumulated chemical cues being neither necessary nor sufficient in this process.     

Intruding male red swamp crayfish,Procambarus clarkii,immediately dominate members of established communities of smaller,mixed-sex conspecifics

The effects of competing asymmetries (intruder size advantage vs. prior residence) on dominance relationships were investigated in a laboratory setting. Sexually mature (Form I) male red swamp crayfish, 25%-27% larger than the average member of several mixedsex communities of 20-25 sexually mature conspecifics, individually intruded upon these communities on successive days. Each community was invaded once a day, with each of these large intruders invading every community once during the experiment. Five days after the last large intruder invasion, novel intruder group males, approximately the same size as the average community member, individually invaded the same communities, all communities being invaded once during a single day of testing. These novel intruders were used to differentiate the effects of intruder size from those effects of being put into a novel environment. During each intrusion, the frequencies of dominance, submission, aggressive standoffs, and nonaggressive interactions between the intruder and members of the community were recorded. The large intruders on each day immediately and virtually completely dominated all encountered community members, and the large intruders were significantly more dominant than the novel, smaller-sized intruders. The size advantage of the large intruders overwhelmed prior residence in influencing dominance outcomes, even in these well established communities. © 1995 Wiley-Liss, Inc.     

Effects of temperature and prior residence on territorial aggression in the convict cichlid Cichlasoma nigrofasciatum

Aggression in the convict cichlid Cichlasoma nigrofasciatum was measured under two different environmental parameters after introduction of a conspecific intruder. Measurements were made after 24-hr and 48-hr teritorial residence times at both 26°C and 30°C. Eight fish established territories in individual tanks each containing one terra cotta pot. Intruders contained in a glass cylinder were introduced into the territories. An opaque divider separating the resident from the intruder was removed after a 15-min acclimation period. At 26°C, we observed a significant difference in the mean number of bites of the cichlids of 24-hr and 48-hr residency durations, whereas, at 30°C, such a difference did not exist. Statistical analysis showed that cichlids exhibited more aggression at 30°C than at 26°C. Increased aggression levels at the higher temperature may be related to the fact that cichlids establish territories and spawn at about 30°C.     

Habituation of aggressive behavior in mice: A parametric study

The decline of aggression as a result of repeated episodes of fighting may represent a habituation phenomenon. The frequency of biting and sideways threat by male mice toward conspecific intruders declined over ten 5-min confrontations. Aggression returned to about 50% of original levels when a new intruder was introduced. Variations in length or frequency of confrontation indicated that the rate of decline of attack bites and sideways threats depended on these confrontation parameters. The condition that provided the fewest opportunities to fight provided the least decline in the frequency of attack and threat. The frequency of attack bites and sideways threats elicited by the presentation of a new intruder after repeated confrontations was greater than the frequency of attack bite and sideways threat in the last confrontation with the original intruder, when the resident and intruder were relatively inexperienced. Spontaneous recovery of attack and threat occurred but rate of recovery did not depend on confrontation parameters. The results indicate that the decrement of aggression in mice in repeated confrontations may be the result of habituation but fatigue is an important cofactor.     

Ultrasounds produced by rats accompany decreases in intraspecific fighting

Male intruder rats were placed individually into the cage of an established resident on 2 occasions separated by a 7–8 day interval. Residents readily attacked intruders and both animals lost weight during the first encounter. In contrast, no serious fighting occurred on the second encounter, and both intruders and residents maintained their body weight during the 24-hr test. Observation of the intruder's behavior during the first 30 min of each encounter indicated that defensive-submissive postures represent a response to an attack that only temporarily inhibits aggression whereas the emission of 22 kHz calls by the intruder is associated with a relatively permanent decrease in the resident animal's aggressive response.     

The effects of urine on aggressive responses by male golden hamsters

Twenty adult male golden hamsters were isolated into individual cages for a period of six weeks, at the end of which time they had introduced into their home cages, on three occasions, a castrated male intruder. On each occasion the castrated intruder was daubed on the anogenital region with urine from one of three sources: (1) intact females, (2) other castrated males, and (3) intact males. Urine from a different source was applied to the castrated intruder on each of the three tests. Resident males consistently showed more aggression, sniffing, and following and less defensive behavior than the intruders. However, aggression by the resident males showed a significant variance over the three urine treatments given to the intruder. It is concluded that like male mouse urine, male hamster urine contains attack-provoking cues, but that unlike that of the mouse, female urine does not appear to be attack-inhibiting in this species.     

The effects of negative reinforcement for irritable aggression on resident-intruder behavior

This experiment demonstrated that rats trained to display elevated levels of shock-induced aggression in a negative reinforcement paradigm displayed more boxing behavior than yoked control groups in a later test in which intruder rats were placed in the home cage of resident rats. Resident or intruder status did not affect the influence of the negative reinforcement procedure on the observed resident-intruder behavior of trained animals; however, naive intruders paired with trained residents displayed increased defensive behavior, suggesting that negative reinforcement for shock-induced aggression affected the behavior of these residents.     

The development of territorial-induced intermale agonistic behavior in albino laboratory mice

Male mice of the CF-1 strain (Mus musculus) were allowed to take up lone residence in a small territory consisting of a 60-cm enclosure attached by a tubular runway to a standard mouse cage with food, water, and bedding. A group of ten mice, each of which resided in its own separate enclosure for 24 hours, were more aggressive toward intruders than other groups of ten mice following six-hour residence periods, or no such residence. Aggression toward intruders increased in repeated weekly tests of the six-hour residents, but after four weeks of testing did not reach the maximum stable level displayed by the 24-hour residents over four weeks of testing. In another experiment, the 24-hour residence period of groups of 20 CF-1 male mice was disturbed by briefly removing the mouse from the enclosure, before introducing the intruder, at various intervals prior to testing. Removal of the resident five minutes before testing resulted in a marked decrease in aggression toward intruders. Although lesser decreases in aggression followed intervals of 30, 45, and 60 minutes, a 180 minute interval resulted in no appreciable effects compared to undisturbed controls. It is concluded that exposure to the stimuli provided by the enclosure results in an aggressive readiness in the resident mouse which reaches a high level within a 24-hour period.     

Maternal territoriality as an offspring defense strategy in red swamp crayfish (Procambarus clarkii,Girard)

The offspring‐defense hypothesis of maternal territoriality in solitary and semisocial mammals [Wolff JO and Peterson JA. 1998. Ethol Ecol Evol 10:227–239] was evaluated in another taxon using maternal and nonmaternal female red swamp crayfish (Procambarus clarkii), a freshwater decapod crustacean that shows pronounced shelter‐related maternal aggression. The maternal females were of three different categories: (1) ovigerous, (2) carrying primarily stage 1 offspring, or (3) tending offspring that were primarily stage 2. Each female was in residence in an individual aquarium containing only a gravel substrate for 24 hr prior to serial intrusions by male conspecifics. Virtually all of the residents, maternal and nonmaternal, excavated a depression (shelter) in the substrate prior to the first intrusion. The results showed that residents in each of the maternal categories won a significantly higher proportion of their encounters than did the nonmaternal residents, which showed only negligible defense and lost every encounter. However, there were no significant differences in encounter outcomes between the maternal resident categories. There was clear defense of the shelter by maternal females, with virtually all fighting being in or around it, thus demonstrating stable, heightened territorial defense throughout these maternal phases of reproduction. This support of the offspring‐defense hypothesis of maternal territoriality is the first such evidence for a freshwater crustacean and is very similar to that recently reported in a related marine decapod, the American lobster (H. americanus). Aggr. Behav. 27:391–403, 2001. © 2001 Wiley‐Liss, Inc.     

A pinch or a lesion: a reconceptualization of biting consequences in mice

Intruder and resident male colony mice exhibit an array of distinct defensive and offensive behaviors. Intruders typically show more boxing, flight, defensive sideways position, on the back position and general locomotion, while residents exhibit higher levels of attack, olfactory investigation, aggressive grooming, and biting, with a preference for dorsal bite locations. Here, analysis of bite locations on the body of the intruder mice showed that the majority of bites produced few lesions (i.e. actual puncturing of the skin) when compared to scrapes or no markings. Most bites were directed to the back of the opponent animal with very few bites directed towards the opponents' vulnerable ventrum. In particular, bites directed at the relatively hairless ventrum produced no lesions. These findings, along with previous work on mice and rats, suggest that intraspecific offense with preferred target sites for biting, facilitates an effective but largely nonwounding interaction between resident and intruder mice. Furthermore, bruise and wound analyses suggest an association between bite targets and tissue damage. The preference for specific bite locations may be complimented by a differential intensity of attack, with the back attracting the strongest bites and the ventrum the weakest. This apparent nonwounding fighting pattern contradicts the current evaluation of rodent wounding severity in this paradigm and can therefore refine the usage of this model and of the protocols associated with it.     

Behaviour of Young Growing Pigs in a Resident‐Intruder Test Designed to Measure Aggressiveness

A total of 125 growing pigs (47 days old) were tested for aggressive responses on two occasions using a resident‐intruder (R‐I) design. Our aims were twofold: (1) to attempt to replicate earlier work on pigs showing that resident aggression is a consistent individual characteristic, unaffected by weight or sex of the resident or intruder and (2) to develop behavioural measures to characterise the wide range of aggressive responses in the test. Resident pigs, housed since birth with littermates, were placed individually in a divided‐off portion of their home pen, and a smaller, unfamiliar intruder (approximately 66% of the resident's weight) was introduced. The test ended 5 min after the first investigation of the intruder by the resident or when one of the pigs began to attack the other (by delivering a sudden, rapid series of bites). On days 1 and 2, 33.6% and 43.2% of tests, respectively, ended in an attack by a resident. Intruder attacks were rare. Pigs were consistent in whether they attacked or not over the two tests, although attack latencies for pigs attacking in both tests were not correlated. Females were more likely to attack and attacked more quickly than males on the first test day but not in the second test or overall. Intruder sex had some effect on the test outcome (males were attacked more rapidly in the second test only). Resident and intruder weight had no effect. Aggressive pigs (meaning pigs that attacked vs. pigs that did not and fast‐attacking pigs vs. slow‐attacking pigs) showed a number of differences in behaviour during the R‐I test: (1) they took longer to make initial contact with the intruder in their first test; (2) they showed a higher frequency of aggressive acts (single head knocks, bites, and shoves); (3) they spent a greater proportion of the test engaging in social contact with the intruder rather than non‐social behaviours; (4) their social behaviour involved more postures directed toward the head as opposed to flank‐ or rear‐directed postures or re‐establishing social contact; and (5) they showed closer physical contact with intruders during social encounters, as characterised by their lower head positions. Some of these behavioural measures could be used to improve the measuring power of the test in the future. Improved behavioural measures would enable aggressiveness scoring among pigs that did not attack instead of classifying them all together as “non‐attacking.” Aggr. Behav. 28:401–415, 2002. © 2002 Wiley‐Liss, Inc.     

Contradictory asymmetries in body and weapon size,and assessment in fighting male prawns,Macrobrachium rosenbergii

Asymmetries in weapon size and body size, both contributing to an animal's fighting ability, may be contradictory in two fighting Macrobrachium rosenbergii males: One male may be larger in body size but smaller in cheliped size than its rival. The aim of the study was to determine the relevance of these two asymmetries in the assessment of relative fighting ability, and to reveal the effect of the asymmetry level on the structure of contests. Thirty contests were staged between male prawns from the ‘blue-clawed’ morphotype and the ‘orange-clawed’ morphotype. The blue-clawed males were 0%, 10%, or 20% larger in cheliped length but 93%, 45%, or 25% smaller in weight, respectively (n = 10), as compared to their orange-clawed opponents. The blue-clawed males with 10% and 20% advantage in cheliped length almost invariably won the contests. When similar in cheliped length, the 93% smaller blue-clawed males still won half of the contests. This indicated that the prawns ignored large asymmetries in body size even when no asymmetry in cheliped size existed. The results suggest that only cheliped size is used by M. rosenbergii males in the assessment of fighting ability. The effect of asymmetry level on contest structure was apparent only with the increase from 10% to 20% difference in cheliped size: Both the frequency and the intensity of fighting were lower in contests between prawns with 20% difference, as compared to 0% and 10% difference in cheliped length. Aggr. Behav. 23:81–91, 1997.© 1997 Wiley-Liss, Inc.     

Dominance-subordinance in cohabiting pairs of adult rats: Effects on Aggressive behavior

Following an initial intruder aggression test, 10 pairs of adult male rats matched on aggressiveness were formed. The first 20 min of paired cohabitation were used to determine dominance and subordinance between pau members. Residents judged to be dominant from this observation session gained significantly more weight during cohabitation and exhibited significantly more aggression on the second aggression test than their subordinate counterparts. Significant correlations among various measures of aggression were found, but open field performance did not correlate, with the measures of aggression nor did changes in open field scores reflect changes in aggression.     

Sensory control of maternal aggression in Rattus norvegicus

Female rats become aggressive toward adult conspecifics during lactation. This change in social affect is dependent on the presence of the offspring, because maternal aggression disappears when the pups are removed. It was found that a similar decline occurs when the litter is placed in a glass flask while remaining in the home cage. In contrast, maternal aggression persists following placement of the pups in a nylon mesh bag. The pups did not vocalize while being in the mesh bags, so it appears that olfactory cues from the offspring constitute a critical element in the maintenance of maternal aggression in the rat. It has been suggested that the odor not only of the pups but also of the intruder may contribute to eliciting aggressive behavior in lactating rodents. In line with this proposal, it was found that mother rats spend about one third of the time preceding the first attack sniffing the body of the intruder. In contrast to findings in mice, housing of the prospective intruder behind a double wire mesh partition in the lactating female's home cage failed to reduce her aggressiveness toward him. Rats, then, may require more intimate contact with an individual than do mice for the aggression-reducing effect of familiarization to be observed.     

Role of female aggression in prevention of infanticidal behavior in male common voles,Microtus arvalis (Pallas, 1779)

Infanticidal behavior of male common voles (Microtus arvalis) was investigated in relation to the age of unfamiliar pups. Sires from 18 pairs were removed after parturition of their offspring and replaced by unfamiliar males at different ages of the neonates. In group 1, containing six females with their offspring, the new male was introduced into the females' cages on day 1 after the parturition. In group 2 the new male was introduced on day 5 after parturition and in group 3 on day 9 after parturition. The male was removed after 3 days and the neonates recounted. Male common voles killed up to 100% of the neonates in litters containing neonates younger than 9 days by rapid bites to the head of the neonates (groups 1 and 2, mortality rates of 0.86 and 0.38). However, with older pups, a mortality rate of 0.06 was observed while 33% of the intruder males showed signs of injuries (group 3). These findings show that infanticidal behavior of male common voles represents one factor of pup mortality. The results are discussed in relation to changes in the degree of maternal aggression and in the context of the social system of the common vole for solving the conflict between postpartum estrous mating and infanticidal behavior of male common voles. Aggr. Behav. 23:293–298, 1997. © 1997 Wiley-Liss, Inc.     

A general theory concerning the prenatal origins of cerebral lateralization in humans

The origins of cerebral lateralization in humans are traced to the asymmetric prenatal development of the ear and labyrinth. Aural lateralization is hypothesized to result from an asymmetry in craniofacial development, whereas vestibular dominance is traced to the position of the fetus during the final trimester. A right-ear sensitivity advantage may contribute to a left-hemispheric advantage in speech perception and language functions, whereas left-otolithic dominance may independently promote right-sided motoric dominance and a right-hemispheric superiority in most visuospatial functions. The emergence of handedness is linked to the assumption of an upright posture in the early hominids, whereas the failure to develop clear vestibular asymmetry may underlie the poor motoric lateralization found in several neurodevelopmental disorders.     

Spectral versus temporal features in dichotic listening

Ear advantage for the processing of dichotic speech sounds can be separated into two components. One of these components is an ear advantage for those phonetic features that are based on spectral acoustic cues. This ear advantage follows the direction of a given individual's ear dominance for the processing of spectral information in dichotic sounds, whether speech or nonspeech. The other factor represents a right-ear advantage for the processing of temporal information in dichotic sounds, whether speech or nonspeech. The present experiments were successful in dissociating these two factors. Since the results clearly show that ear advantage for speech is influenced by ear dominance for spectral information, a full understanding of the asymmetry in the perceptual salience of speech sounds in any individual will not be possible without knowing his ear dominance.