A simple explanation for the evolution of complex song syntax in Bengalese finches

The songs of Bengalese finches (Lonchura striata var. domestica) have complex syntax and provide an opportunity to investigate how complex sequential behaviour emerges via the evolutionary process. In this study, we suggest that a simple mechanism, i.e. many-to-one mapping from internal states onto syllables, may underlie the emergence of apparent complex syllable sequences that have higher order history dependencies. We analysed the songs of Bengalese finches and of their wild ancestor, the white-rumped munia (L. striata), whose songs are more stereotypical and simpler compared with those of Bengalese finches. The many-to-one mapping mechanism sufficiently accounted for the differences in the complexity of song syllable sequences of these two strains.     

Age effect of deafening on stereotyped song maintenance in adult male bengalese finches Lonchura striata domestica

Birdsong is a complex learned vocal behavior that relies on auditory experience for development.However,it appears that among different species of close-ended songbirds,there are some variations in the necessity of auditory feedback for maintaining stereotyped adult song.In zebra finches,the deterioration of adult songs following deafness depends on the birds' age.It is unknown whether this age effect is a general rule in other avian species as well.Therefore,we chose Bengalese finches,whose songs show more...     

Uropygial gland volatiles may code for olfactory information about sex,individual,and species in Bengalese finches Lonchura striata

Over-shadowed by eye-catching vocal and visual signals,chemical communication has long been overlooked in birds.This study aimed at exploring whether volatile composition of the uropygial gland secretion(UGS)of birds was associated with the information about sex,individual and species.By using dichloromethane extraction and gas chromatography-mass spectrometry(GC-MS),we analyzed the UGS volatiles of domesticated Bengalese finches(Lonchura striata,Estrildiea)which is also known as white-rumped munias.We char...     

Uropygial gland volatiles may code for olfactory information about sex,individual, and species in Bengalese finches Lonchura striata

Over-shadowed by eye-catching vocal and visual signals, chemical communication has long been overlooked in birds. This study aimed at exploring whether volatile composition of the uropygial gland secretion （UGS） of birds was associated with the information about sex, individual and species. By using dichloromethane extraction and gas chromatography-mass spectrometry （GC-MS）, we analyzed the UGS volatiles of domesticated Bengalese finches （ Lonchura striata, Estrildiea） which is also known as white-rumped munias. We characterized 16 volatile molecules from the UGS, including eight n-alkanols, five diesters, an ester, an aldehyde and a fatty acid, and quantified them in terms of GC peak area percentages （relative abundances） . Among these compounds, hexadecanol and octadecanol were major components in both sexes. The former was richer in males than in females and the latter richer in females than in males, suggesting that they might be male and female pheromone candidates, respectively. The high inter-individual variations, in relative abundance, of the UGS volatiles implied that these compounds might carry information about individuality. The similarity between GC profiles of the UGS and wing feather from same individuals indicates that the birds might preen the secretion to their feathers to transmit chemical cues. Additionally, by comparing with three sympatric passerine species, i. e., zebra finches Taeniopygia guttata, yellow-bowed buntings Emberiza chrysophrys and rooks Corvus frugilegus, we found that the composition of C13 - C18 alkanols in the UGS might code for information about species. Our study also showed that quantitative differences （degree） of same UGS volatiles might be the key for the Bengalese finch to code for information about sex and individuality whereas both the kind and degree of UGS constituents could be utilized to code for information about species [ Current Zoology 55 （5）： 357-365, 2009].     

Migratory charr schools exhibit population and kin associations beyond juvenile stages

Few studies have critically investigated the genetic composition of wild fish schools. Yet, such investigations may have profound implications for the understanding of social organization and population differentiation in both fundamental and applied research. Using 20 microsatellite loci, we investigated the composition of 53 schools (total n = 211) of adult and subadult migratory brook charr (Salvelinus fontinalis) sampled from the known feeding areas of two populations inhabiting Mistassini Lake (Québec, Canada). We specifically tested whether (i) school members originated from the same population, (ii) individuals from the same population within schools were kin (half- or full-siblings), and (iii) kin schooling relationships differed between sexes. Randomization tests revealed a tendency for most schools to be population specific, although some schools were population mixtures. Significantly more kin were found within schools than expected at random for both populations (approximately 21-34% of the total number of school members). This result, combined with the observed size range of individuals, indicated that stable associations between kin may occur beyond juvenile stages for up to 4 years. Nevertheless, a high proportion of school members were non-kin (approximately 66-79%). No differences were detected between sexes in the propensity to school with kin. We discuss the hypothesis that the stable kin groups, rather than arising from kin selection, may instead be a by-product of familiarity based on individual selection for the maintenance of local adaptations related to migration (natal and feeding area philopatry). Our results are noteworthy because they suggest that there is some degree of permanence in the composition of wild fish schools. Additionally, they support the hypothesis that schools can be hierarchically structured (from population members down to family groups) and are thus nonrandom genetic entities.     

Effects of kinship on aggression and RNA content in juvenile Arctic charr

The behaviour of juvenile Arctic charr Salvelinus alpinus was studied in groups of four siblings composed of familiar and unfamiliar individuals or in mixed groups of four where both siblings and unrelated individuals were present. The frequency of aggressive acts was significantly higher in the mixed groups compared to the pure sibling groups and the difference was present at all levels of the hierarchy rank order, based on a dominance index, except the lowest ranked individuals. The difference was significant after but not before feeding, implying that competition with non-kin for a food resource increased the aggression. No significant difference in weight gain was observed between sibling and mixed groups during the 6 days of the experiment, but the RNA contents of lateral musculature in dominant individuals from sibling groups were significantly higher than the corresponding fish in the mixed group, suggesting a difference in growth rate when the experiments ended. No significant difference in RNA content was observed between subordinate fish of the two treatments, i.e. siblings v . mixed.     

Forebrain circuits underlying the social modulation of vocal communication signals

Across vertebrate species, signalers alter the structure of their communication signals based on the social context. For example, male Bengalese finches produce faster and more stereotyped songs when directing song to females (female‐directed [FD] song) than when singing in isolation (undirected [UD] song), and such changes have been found to increase the attractiveness of a male's song. Despite the importance of such social influences, little is known about the mechanisms underlying the social modulation of communication signals. To this end, we analyzed differences in immediate early gene (EGR‐1) expression when Bengalese finches produced FD or UD song. Relative to silent birds, EGR‐1 expression was elevated in birds producing either FD or UD song throughout vocal control circuitry, including the interface nucleus of the nidopallium (NIf), HVC, the robust nucleus of the arcopallium (RA), Area X, and the lateral magnocellular nucleus of the anterior nidopallium (LMAN). Moreover, EGR‐1 expression was higher in HVC, RA, Area X, and LMAN in males producing UD song than in males producing FD song, indicating that social context modulated EGR‐1 expression in these areas. However, EGR‐1 expression was not significantly different between males producing FD or UD song in NIf, the primary vocal motor input into HVC, suggesting that context‐dependent changes could arise de novo in HVC. The pattern of context‐dependent differences in EGR‐1 expression in the Bengalese finch was highly similar to that in the zebra finch and suggests that social context affects song structure by modulating activity throughout vocal control nuclei. © 2015 Wiley Periodicals, Inc. Develop Neurobiol 76: 47–63, 2016     

Shoaling preference and evidence for maintenance of sibling groups by juvenile black perch Embiotoca jacksoni

Group sizes in free‐living juvenile black perch Embiotoca jacksoni were quantified and predictions of the hypothesis that such groups comprise sibling brood‐mates were tested. Group sizes in the field were within the range of female brood sizes and often occurred close to each other but did not merge. In captivity, juveniles formed groups immediately after birth. In laboratory experiments, they also associated significantly more with chambers containing familiar members of their own brood than empty chambers but did not associate more with chambers containing similar‐sized juveniles from a different brood. Juvenile E. jacksoni also associated significantly more with chambers containing familiar brood‐mates than with chambers containing unfamiliar members of a different brood. The strength of this preference increased with the number of days fish had been together since birth. When two broods were placed in a large outdoor tank, all individuals from both broods directed significantly more aggressive acts towards members of the other brood than towards members of their own brood. While the relative effects of familiarity and relatedness cannot be completely separated in this viviparous species, associating with familiar individuals would facilitate the maintenance of sibling groups in the field.     

Song recognition memory in juvenile zebra finches: effects of varying the number of presentations of heterospecific and conspecific songs

Seven male and three female zebra finches were exposed to 14 zebra finch (CON) and 14 starling (HET) songs during their sensitive period for song learning and then tested for their recognition memory of both the CON and HET songs in two separate memory tests. Amount of song exposure was varied by presenting individual songs either 3, 9, 27, or 81 times per day for nine consecutive days. After song exposure the birds were trained to discriminate two of the exposed, familiar songs (FAM) from two novel songs (NOV) in a go/no-go operant discrimination procedure, with FAM songs as "go" stimuli. Following discrimination training, untrained FAM and NOV songs were presented as probe songs without reinforcement. Birds responded more to FAM than NOV songs at all levels of song exposure, indicating that the songs were recognized. There were no differences in recognition memory for CON and HET song at any level of song exposure. The results suggest that selective song learning does not result from selective memorization of conspecific song.     

Fine-tuned modulation of competitive behaviour according to kinship in barn swallow nestlings

Kin selection theory predicts that, in species where progeny members compete for limiting parental care, individual offspring should be more prone to monopolize parental resources as their genetic relatedness to brood competitors decreases. Mixed parentage among broodmates may arise as a consequence, for example, of extra-pair fertilization or brood parasitism events. In this experimental study of barn swallows (Hirundo rustica), we reciprocally partially cross-fostered hatchlings between broods and compared the behaviour of pairs of related and unrelated broodmates in a competitive context, both under normal food provisioning regime and after mild food deprivation. We found that scramble competition for food mediated by visual and vocal solicitation displays (begging) is inversely related to relatedness among competitors, independent of their level of satiation. Nestlings may modulate their competitive behaviour according to vocal cues that vary with their origin and allow kin recognition. We also uncover direct fitness costs to both parents and offspring arising from mixed parentage in a brood, in terms of increased parental workload and reduced survival of the nestlings. Such previously neglected costs may select for reduced frequency of extra-pair fertilizations and brood parasitism in species with extensive parental care.     

Ground squirrel sociality and the quest for the 'holy grail': does kinship influence behavioral discrimination by juvenile Columbian ground squirrels?

Research on social discrimination by ground-dwelling squirrelshas focused on die ability of squirrels to discriminate kinfrom nonkin. The predominant mechanism underlying that abilityis social familiarization. Although familiarity-based mechanismsmay result in kin discrimination where kin are associated reliablyin space and time, past investigations have largely ignoredadditional levels of social discrimination that may result fromongoing familiarization in the natural context Here we presentdata from a cross-fostering experiment that examines the relativecontributions of rearing association and relatedness to subsequentbehavioral discrimination among juvenile Columbian ground squirrels(Spermophilus columbianus). Rearing association significantlyinfluenced recognitive and agonistic behavior among juvenileswhile relatedness proved insignificant in affecting behavioralinteractions. Thus, direct familiarization in the natal burrowis both sufficient and necessary to account for the transitiveappearance of kin-differential behavior among newly emergedjuveniles documented in previous studies. Reanalyses of datafrom field experiments on S. columbianus social discriminationfail to detect any evidence of kin-biased behavior, and evidencefrom studies purporting to document kinship effects is equivocalat best Taken together with more recent findings, the scantdata suggestive of kin discrimination by Columbian ground squirrelsare best viewed as an artifact of selection promoting the discriminationof familiar from unfamiliar individuals at the level of localizedgroups. Fitness payoffs of such discrimination may accrue viareciprocal altruism, or ‘dear-enemy’ recognition,and would promote the evolution and maintenance of socialitywithout recourse to indirect components of inclusive fitnesscommonly invoked to explain ground squirrel sociality. Finally,our data call into doubt the notion that mechanisms allowingthe direct assessment of kinship should be common among themore social sciurids.     

Host-parasite relatedness in wood ducks: patterns of kinship and parasite success

We investigated the role of kinship in intraspecific nest parasitismof wood ducks (Aix sponsa). Among waterfowl, female philopatrycreates the potential for female relatives to nest in proximity.Costs of intraspecific nest parasitism to host females may bereduced if parasites lay eggs with kin. However, previous observationsof marked wood ducks indicated that females avoided parasitizingclutch mates or the female that incubated them. To further examinethe role of kinship, we determined the genotypes of 27 host-parasitepairs at five microsatellite loci. Average relatedness betweenhosts and all females laying parasitic eggs was only 0.04 ±0.03. Parasites appeared to choose hosts randomly with respectto kinship from among females with nests in the neighborhoodand those within the entire study area. However, host relatednessto the parasite with the greatest number of young leaving thenest was 0.11 ± 0.03, which was greater than expectedif eggs were accepted randomly from neighboring females or fromfemales present on the entire study area (p = .03 and p = .02,respectively). These patterns may reflect parasitism of randomlyselected nests followed by differential acceptance by hosts,differential hatching success of related parasites (e.g., dueto greater laying synchrony), or a mixture of parasitic strategies,one with a focus on related hosts and the other on unrelatedhosts. Genetic data revealed that social relationships did notalways reflect true relatedness and that success of primaryparasites was associated with kinship to hosts.     

The cost of aggregation: juvenile salmon avoid sharing winter refuges with siblings

Kin selection advantages are usually accrued by individualsthat associate with close relatives. But aggregation may alsobe costly, by increasing the risk of predation or resource competition,for example. As a result, individuals should increase theirinclusive fitness by trading the costs and benefits of kin associationand aggregation. Studies of kin selection to date have focusedon situations where there is ample opportunity for kin-biasedbehavior and therefore for the formation of kin groups. Herewe used juvenile Atlantic salmon to test an alternative strategy:that under conditions where the potential for kin-biased behavioris negligible, individuals should, when aggregating, avoid ratherthan associate with kin to avoid imposing the costs of aggregationupon close relatives. By testing salmon during winter, whenjuveniles shelter inactively in streambed refuges, we testedwhether individuals associate with or avoid their siblings ata time when the opportunity for kin-directed behaviors is restricted.Our results provide the first evidence of kin avoidance in nonreproductiveanimals studied under semi-natural conditions.     

Brood parasitism,relatedness and sociality: a kinship role in female reproductive tactics

Conspecific brood parasitism (CBP) is a reproductive tactic in which parasitic females lay eggs in nests of other females of the same species that then raise the joint brood. Parasites benefit by increased reproduction, without costs of parental care for the parasitic eggs. CBP occurs in many egg‐laying animals, among birds most often in species with large clutches and self‐feeding young: two major factors facilitating successful parasitism. CBP is particularly common in waterfowl (Anatidae), a group with female‐biased natal philopatry and locally related females. Theory suggests that relatedness between host and parasite can lead to inclusive fitness benefits for both, but if host costs are high, parasites should instead target unrelated females. Pairwise relatedness (r) in host–parasite (h‐p) pairs of females has been estimated using molecular genetic methods in seven waterfowl (10 studies). In many h‐p pairs, the two females were unrelated (with low r, near the local population mean). However, close relatives (r = 0.5) were over‐represented in h‐p pairs, which in all 10 studies had higher mean relatedness than other females. In one species where this was studied, h‐p relatedness was higher than between nesting close neighbours, and hosts parasitized by non‐relatives aggressively rejected other females. In another species, birth nest‐mates (mother–daughters, sisters) associated in the breeding area as adults, and became h‐p pairs more often than expected by chance. These and other results point to recognition of birth nest‐mates and perhaps other close relatives. For small to medium host clutch sizes, addition of a few parasitic eggs need not reduce host offspring success. Estimates in two species suggest that hosts can then gain inclusive fitness if parasitized by relatives. Other evidence of female cooperation is incubation by old eider Somateria mollissima females of clutches laid by their relatives, and merging and joint care of broods of young. Merging females tended to be more closely related. Eiders associate with kin in many situations, and in some geese and swans, related females may associate over many years. Recent genetic evidence shows that also New World quails (Odontophoridae) have female‐biased natal philopatry, CBP and brood merging, inviting further study and comparison with waterfowl. Kin‐related parasitism also occurs in some insects, with revealing parallels and differences compared to birds. In hemipteran bugs, receiving extra eggs is beneficial for hosts by diluting offspring predation. In eggplant lace bugs Gargaphia solani, host and parasite are closely related, and kin selection favours egg donation to related females. Further studies of kinship in CBP, brood merging and other contexts can test if some of these species are socially more advanced than presently known.