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1.  The deterrent effect of bird song in territory defense 
Behavioral Ecology  2008;20(1):200-206.
Using the responses of territory owners to playback to infer the territorial function of acoustic signals is common practice. However, difficulties with interpreting the results of such experiments have obscured our understanding of territorial signalling. For instance, a stronger response to playback is often interpreted as more aggressive, but there is no consensus as to whether this should be in response to the least or most threatening simulated intruder. Rather than following a gradual increase or decrease, the relationship between signal intensity and response strength may instead describe a peaked curve. We manipulated banded wren (Thryophilus pleurostictus) songs to simulate low-, median-, and high-performance singers and used these songs as stimuli in playback experiments. Banded wrens were less likely to approach the high-performance stimulus compared with the low- and median-performance stimuli. However, the birds that did approach the high-performance stimulus sang more than those that approached the low-performance stimulus. In addition, birds were more likely to match the songs when exposed to the median- and high-performance stimuli compared with the low-performance stimuli, and song matching predicted approach behavior. These results are in accordance with theoretical models of aggressive encounters in which low-performance opponents are challenged without further assessment. Median- and high-performance opponents, however, may require further assessment, and the latter may be perceived as too intimidating for approach.
doi:10.1093/beheco/arn135
PMCID: PMC2662740  PMID: 19337589
assessment; playback; sexual selection; song; territory defense
2.  Courtship attention in sagebrush lizards varies with male identity and female reproductive state 
Behavioral Ecology  2008;19(6):1326-1332.
Previous experiments suggest that males spend more time with the more receptive of 2 novel females or the one with the higher fitness potential. However, males often court individual females repeatedly over a season; for example, male lizards sequentially visit familiar females as they patrol territorial boundaries. It may benefit males to vary display intensity as they move between multiple females. In this study, we explored the factors influencing amount of male courtship to familiar females in the sagebrush lizard, Sceloporus graciosus. We tested whether males vary the amount of courtship exhibited due to individual differences among males, female reproductive state, or female fitness potential. Each male was allowed to interact separately, but repeatedly, with 2 females until both females laid eggs. Male courtship behavior with each of the 2 females was assayed at an intermediate point, after 3 weeks of interaction. We found that individual differences among males were considerable. The number of male courtship displays was also positively correlated with female latency to lay eggs, with males displaying more often toward females with eggs that had not yet been fertilized. Courtship behavior was not well predicted by the number of eggs laid or by female width, both measures of female quality. Thus, male S. graciosus appear to alter courtship intensity more in response to signals of female reproductive state than in response to variation in potential female fitness.
doi:10.1093/beheco/arn072
PMCID: PMC2583109  PMID: 19458780
courtship; male choice; mate choice; reproductive state; Sceloporus graciosus; sexual selection
3.  The deterrent effect of bird song in territory defense 
Using the responses of territory owners to playback to infer the territorial function of acoustic signals is common practice. However, difficulties with interpreting the results of such experiments have obscured our understanding of territorial signaling. For instance, a stronger response to playback is often interpreted as more aggressive, but there is no consensus as to whether this should be in response to the least or most threatening simulated intruder. Rather than following a gradual increase or decrease, the relationship between signal intensity and response strength may instead describe a peaked curve. We manipulated banded wren (Thryophilus pleurostictus) songs to simulate low, median and high performance singers and used these songs as stimuli in playback experiments. Banded wrens were less likely to approach the high performance stimulus compared to the low and median performance stimuli. However, the birds that did approach the high performance stimulus sang more than those that approached the low performance stimulus. In addition, birds were more likely to match the songs when exposed to the median and high performance stimuli compared to the low performance stimuli and song matching predicted approach behavior. These results are in accordance with theoretical models of aggressive encounters in which low performance opponents are challenged without further assessment. Median and high performance opponents however, may require further assessment and the latter may be perceived as too intimidating for approach.
doi:10.1093/beheco/arn135
PMCID: PMC2662740  PMID: 19337589
playback; song; territory defense; sexual selection; assessment
4.  Courtship attention in sagebrush lizards varies with male identity and female reproductive state 
Previous experiments suggest that males spend more time with the more receptive of two novel females or the one with the higher fitness potential. However, males often court individual females repeatedly over a season; for example, male lizards sequentially visit familiar females as they patrol territorial boundaries. It may benefit males to vary display intensity as they move between multiple females. In this study, we explored the factors influencing amount of male courtship to familiar females in the Sagebrush lizard, Sceloporus graciosus. We tested whether males vary the amount of courtship exhibited due to individual differences among males, female reproductive state, or female fitness potential. Each male was allowed to interact separately, but repeatedly, with two females until both females laid eggs. Male courtship behavior with each of the two females was assayed at an intermediate point, after three weeks of interaction. We found that individual differences among males were considerable. The number of male courtship displays was also positively correlated with female latency to lay eggs, with males displaying more often towards females with eggs that had not yet been fertilized. Courtship behavior was not well predicted by the number of eggs laid or by female width, both measures of female quality. Thus, male S. graciosus appear to alter courtship intensity more in response to signals of female reproductive state than in response to variation in potential female fitness.
doi:10.1093/beheco/arn072
PMCID: PMC2583109  PMID: 19458780
Sceloporus graciosus; male choice; mate choice; sexual selection; reproductive state; courtship
5.  Genome-Wide Study of Response to Platinum, Taxane, and Combination Therapy in Ovarian Cancer: In vitro Phenotypes, Inherited Variation, and Disease Recurrence 
Background: The standard treatment for epithelial ovarian cancer (EOC) patients with advanced disease is carboplatin-paclitaxel combination therapy following initial debulking surgery, yet there is wide inter-patient variation in clinical response. We sought to identify pharmacogenomic markers related to carboplatin-paclitaxel therapy.
Methods: The lymphoblastoid cell lines, derived from 74 invasive EOC patients seen at the Mayo Clinic, were treated with increasing concentrations of carboplatin and/or paclitaxel and assessed for in vitro drug response using MTT viability and caspase3/7 apoptosis assays. Drug response phenotypes IC50 (effective dose at which 50% of cells are viable) and EC50 (dose resulting in 50% induction of caspase 3/7 activity) were estimated for each patient to paclitaxel and carboplatin (alone and in combination). For each of the six drug response phenotypes, a genome-wide association study was conducted.
Results: Statistical analysis found paclitaxel in vitro drug response phenotypes to be moderately associated with time to EOC recurrence (p = 0.008 IC50; p = 0.058 EC50). Although no pharmacogenomic associations were significant at p < 5 × 10−8, seven genomic loci were associated with drug response at p < 10−6, including at 4q21.21 for carboplatin, 4p16.1 and 5q23.2 for paclitaxel, and 3q24, 10q, 1q44, and 13q21 for combination therapy. Nearby genes of interest include FRAS1, MGC32805, SNCAIP, SLC9A9, TIAL1, ZNF731P, and PCDH20.
Conclusions: These results suggest the existence of genetic loci associated with response to platinum-taxane therapies. Further research is needed to understand the mechanism by which these loci may impact EOC clinical response to this commonly used regimen.
doi:10.3389/fgene.2016.00037
PMCID: PMC4801852  PMID: 27047539
pharmacogenomics; genome-wide association; carboplatin; paclitaxel; ovarian cancer; cell viability; apoptosis; lymphoblastoid cell lines
6.  A Prospective Study of Asymptomatic Intracranial Atherosclerotic Stenosis in Neurologically Normal Volunteers in a Japanese Cohort 
Atherosclerotic stenosis of major intracranial arteries is a leading cause of ischemic stroke in Asia. However, the long-term prognosis of asymptomatic intracranial atherosclerotic stenosis (ICAS) in healthy volunteers has not been fully examined. Here, we conducted a longitudinal study to examine the prognosis of healthy volunteers with asymptomatic ICAS and to determine the risk factors for ICAS, including asymptomatic brain parenchymal lesions. We studied 2,807 healthy Japanese volunteers with no history of stroke (mean age, 62.0 years). They were followed for a mean interval of 64.5 months. The degree of ICAS and the presence of asymptomatic brain lesions were assessed by using magnetic resonance imaging. Asymptomatic ICAS was detected in 166 volunteers (5.9%) at the initial examination. Moderate and mild stenoses were observed in 1.5 and 4.4% of patients, respectively. Significant risk factors for ICAS were older age and a history of hypertension and/or dyslipidemia. During follow-up, ischemic stroke developed in 32 volunteers. Seven strokes occurred in the ICAS group, whose stroke incidence rate was higher than that in the non-ICAS group (0.78 vs. 0.18% per year). According to a Cox regression analysis, asymptomatic ICAS was an independent risk factor for future ischemic stroke after adjustment for age. Furthermore, after asymptomatic brain lesions were taken into account, ICAS was still a significant risk factor for stroke onset. In conclusion, even mild to moderate asymptomatic ICAS was a significant risk factor for future stroke, independent of asymptomatic brain lesions, in a healthy Japanese population. Mild to moderate ICAS might be a therapeutic target for stroke prevention.
doi:10.3389/fneur.2016.00039
PMCID: PMC4801853  PMID: 27047445
asymptomatic brain lesions; asymptomatic intracranial atherosclerotic stenosis; risk factors; longitudinal study; magnetic resonance angiography; prognosis
7.  Giant Viruses of Amoebas: An Update 
During the 12 past years, five new or putative virus families encompassing several members, namely Mimiviridae, Marseilleviridae, pandoraviruses, faustoviruses, and virophages were described. In addition, Pithovirus sibericum and Mollivirus sibericum represent type strains of putative new giant virus families. All these viruses were isolated using amoebal coculture methods. These giant viruses were linked by phylogenomic analyses to other large DNA viruses. They were then proposed to be classified in a new viral order, the Megavirales, on the basis of their common origin, as shown by a set of ancestral genes encoding key viral functions, a common virion architecture, and shared major biological features including replication inside cytoplasmic factories. Megavirales is increasingly demonstrated to stand in the tree of life aside Bacteria, Archaea, and Eukarya, and the megavirus ancestor is suspected to be as ancient as cellular ancestors. In addition, giant amoebal viruses are visible under a light microscope and display many phenotypic and genomic features not found in other viruses, while they share other characteristics with parasitic microbes. Moreover, these organisms appear to be common inhabitants of our biosphere, and mimiviruses and marseilleviruses were isolated from human samples and associated to diseases. In the present review, we describe the main features and recent findings on these giant amoebal viruses and virophages.
doi:10.3389/fmicb.2016.00349
PMCID: PMC4801854  PMID: 27047465
giant virus; mimivirus; amoeba; Acanthamoeba; Megavirales; virophage; 4th TRUC
8.  Bone marrow-derived TNF-α causes diabetic neuropathy in mice 
Diabetologia  2014;58(2):402-410.
Aims/hypothesis
Dysregulation of biochemical pathways in response to hyperglycaemia in cells intrinsic to the nervous system (Schwann cells, neurons, vasa nervorum) are thought to underlie diabetic peripheral neuropathy (DPN). TNF-α is a known aetiological factor; Tnf-knockout mice are protected against DPN. We hypothesised that TNF-α produced by a small but specific bone marrow (BM) subpopulation marked by proinsulin production (proinsulin-producing BM-derived cells, PI-BMDCs) is essential for DPN development.
Methods
We produced mice deficient in TNF-α, globally in BM and selectively in PI-BMDCs only, by gene targeting and BM transplantation, and induced diabetes by streptozotocin. Motor and sensory nerve conduction velocities were used to gauge nerve dysfunction. Immunocytochemistry, fluorescent in-situ hybridisation (FISH) and PCR analysis of dorsal root ganglia (DRG) were employed to monitor outcome.
Results
We found that loss of TNF-α in BM only protected mice from DPN. We developed a strategy to delete TNF-α specifically in PI-BMDCs, and found that PI-BMDC-specific loss of TNF-α protected against DPN as robustly as loss of total BM TNF-α. Selective loss of PI-BMDC-derived TNF-α downregulated TUNEL-positive DRG neurons. FISH revealed PI-BMDC–neuron fusion cells in the DRG in mice with DPN; fusion cells were undetectable in non-diabetic mice or diabetic mice that had lost TNF-α expression selectively in the PI-BMDC subpopulation.
Conclusions/interpretation
BMDC-specific TNF-α is essential for DPN development; its selective removal from a small PI-BMDC subpopulation protects against DPN. The pathogenicity of PI-BMDC-derived TNF-α may have important therapeutic implications.
doi:10.1007/s00125-014-3440-4
PMCID: PMC4289451  PMID: 25399355
Apoptosis; Bone marrow; Cell fusion; Diabetic complications; Diabetic neuropathy; Dorsal root ganglia; Hyperglycaemia; Peripheral nervous system; Proinsulin-producing bone marrow-derived cells; TNF-α
9.  Comparison of Smoking Cessation Between Education Groups: Findings From 2 US National Surveys Over 2 Decades 
American journal of public health  2015;105(2):373-379.
Objectives
We examined smoking cessation rate by education and determined how much of the difference can be attributed to the rate of quit attempts and how much to the success of these attempts.
Methods
We analyzed data from the National Health Interview Survey (NHIS, 1991–2010) and the Tobacco Use Supplement to the Current Population Survey (TUS-CPS, 1992–2011). Smokers (≥ 25 years) were divided into lower- and higher-education groups (≤ 12 years and > 12 years).
Results
A significant difference in cessation rate between the lower- and the higher-education groups persisted over the last 2 decades. On average, the annual cessation rate for the former was about two thirds that of the latter (3.5% vs 5.2%; P < .001, for both NHIS and TUS-CPS). About half the difference in cessation rate can be attributed to the difference in quit attempt rate and half to the difference in success rate.
Conclusions
Smokers in the lower-education group have consistently lagged behind their higher-education counterparts in quitting. In addition to the usual concern about improving their success in quitting, tobacco control programs need to find ways to increase quit attempts in this group.
doi:10.2105/AJPH.2014.302222
PMCID: PMC4289455  PMID: 25521868
10.  ILC1s in Tissue Inflammation and Infection 
Innate lymphoid cells (ILCs) are innate immune cells that provide an early source of cytokines to initiate and tailor the immune response to the type of the encountered pathogen or insult. The group 1 ILCs are comprised of conventional natural killer (cNK) cells and subsets of “unconventional NK cells,” termed ILC1s. Although cNK cells and ILC1s share many features, such as certain phenotypic markers and the ability to produce IFN-γ upon activation, it is now becoming apparent that these two subsets develop from different progenitors and show unique tissue distribution and functional characteristics. Recent studies have aimed at elucidating the individual contributions of cNK cells and ILC1s during protective host responses as well as during chronic inflammation. This review provides an overview of the current knowledge of the developmental origins as well as of the phenotypic and functional characteristics of ILC1s.
doi:10.3389/fimmu.2016.00104
PMCID: PMC4801855  PMID: 27047491
ILC1; innate lymphoid cells; inflammation; host response; ILC development; antimicrobial defense
11.  Working Memory Training and Speech in Noise Comprehension in Older Adults 
Understanding speech in the presence of background sound can be challenging for older adults. Speech comprehension in noise appears to depend on working memory and executive-control processes (e.g., Heald and Nusbaum, 2014), and their augmentation through training may have rehabilitative potential for age-related hearing loss. We examined the efficacy of adaptive working-memory training (Cogmed; Klingberg et al., 2002) in 24 older adults, assessing generalization to other working-memory tasks (near-transfer) and to other cognitive domains (far-transfer) using a cognitive test battery, including the Reading Span test, sensitive to working memory (e.g., Daneman and Carpenter, 1980). We also assessed far transfer to speech-in-noise performance, including a closed-set sentence task (Kidd et al., 2008). To examine the effect of cognitive training on benefit obtained from semantic context, we also assessed transfer to open-set sentences; half were semantically coherent (high-context) and half were semantically anomalous (low-context). Subjects completed 25 sessions (0.5–1 h each; 5 sessions/week) of both adaptive working memory training and placebo training over 10 weeks in a crossover design. Subjects' scores on the adaptive working-memory training tasks improved as a result of training. However, training did not transfer to other working memory tasks, nor to tasks recruiting other cognitive domains. We did not observe any training-related improvement in speech-in-noise performance. Measures of working memory correlated with the intelligibility of low-context, but not high-context, sentences, suggesting that sentence context may reduce the load on working memory. The Reading Span test significantly correlated only with a test of visual episodic memory, suggesting that the Reading Span test is not a pure-test of working memory, as is commonly assumed.
doi:10.3389/fnagi.2016.00049
PMCID: PMC4801856  PMID: 27047370
cognitive training; Cogmed; working memory training; speech-in-noise; speech perception; reading span
12.  Editorial: Combination Strategies in the Treatment of Melanoma 
doi:10.3389/fonc.2016.00067
PMCID: PMC4801857  PMID: 27047798
melanoma; immunotherapy; vasculopathy; inhibitors; angiogenesis inhibitors
13.  Advancing regionalism and a social policy agenda for positive change: From rhetoric to action 
Global Social Policy  2015;15(3):329-335.
doi:10.1177/1468018115600123
PMCID: PMC4639825  PMID: 26635501
14.  Issue Information 
British Journal of Pharmacology  2014;171(21):i-ii.
doi:10.1111/bph.12390
PMCID: PMC4294104
15.  Variance in male lifetime reproductive success and estimation of the degree of polygyny in a primate 
Behavioral Ecology  2014;25(4):878-889.
The degree of polygyny is predicted to influence the strength of direct male–male competition, leading to a high variance in male lifetime reproductive success and to reproduction limited to the prime period of adulthood. Here, we explore the variance in male lifetime reproductive success and reproductive time in an anthropoid primate forming multimale–multifemale groups. Males of this species form dominance hierarchies, which are expected to skew reproduction toward few high-ranking males. At the same time, however, females mate with multiple males (polygynandry), which should limit the degree of polygyny. Using 20 years of genetic and demographic data, we calculated lifetime reproductive success for the free-ranging rhesus macaque (Macaca mulatta) population of Cayo Santiago for subjects that died naturally or reached senescence. Our results show that 1) male lifetime reproductive success was significantly skewed (range: 0–47 offspring; males reproducing below average: 62.8%; nonbreeders: 17.4%), 2) variance in male lifetime reproductive success was 5 times larger than in females, and 3) male lifetime reproductive success was more influenced by variation in fecundity (60%) than longevity (25%), suggesting that some direct male–male competition takes place. However, the opportunity for selection (i.e., standardized variance in male lifetime reproductive success) is low compared with that in other large mammal species characterized by a high degree of polygyny. Moreover, male reproductive life extended much beyond the prime period, showing that physical strength was not required to acquire mates. We conclude that rhesus macaques exhibit a moderate degree of polygyny and, therefore, low levels of direct male–male competition for fertile females, despite the fact that males form linear dominance hierarchies.
Lay summary
Male rhesus macaques face moderate strength of direct competition for fertile mates, creating opportunity for indirect forms of competition. Using 20 years of genetic and demographic data, we show that variance in lifetime reproductive success is more pronounced in males than in females. Yet, most males reproduce in their lifetime, that surviving as long as possible increases males’ success, and that males reach high fecundity before the full development of secondary sex characteristics.
doi:10.1093/beheco/aru052
PMCID: PMC4095946  PMID: 25024637
lifetime reproductive success; male–male competition; mammals; opportunity for selection; reproductive skew; reproductive timing; rhesus macaques.
16.  Vocal fundamental and formant frequencies are honest signals of threat potential in peripubertal males 
Behavioral Ecology  2014;25(4):984-988.
Lay Summary
Men with lower voices are perceived as more physically dominant; however, it is unclear why, and whether this perception is accurate. We examine whether vocal frequencies are more strongly related to measures of threat potential (size or strength) among adolescents. Results show that vocal frequencies are more informative about strength than about size. These findings suggest that listeners ascribe greater dominance to low male voices because low voices are reliably associated with strength.
Fundamental and formant frequencies influence perceived pitch and are sexually dimorphic in humans. The information content of these acoustic parameters can illuminate the forces of sexual selection shaping vocal sex differences as well as the mechanisms that ensure signal reliability. We use multiple regression to examine the relationships between somatic (height, adiposity, and strength) and acoustic (fundamental frequency [F 0], formant position [P f], and fundamental frequency variation [F 0-SD]) characteristics in a sample of peripubertal Bolivian Tsimane. Results indicate that among males—but not females—strength is the strongest predictor of F 0 and P f and that F 0 and P f are independent predictors of strength when height and adiposity are controlled. These findings suggest that listeners may attend to vocal frequencies because they signal honest, nonredundant information about male strength and threat potential, which are strongly related to physical maturity and which cannot be ascertained from visual or other indicators of height or adiposity alone.
doi:10.1093/beheco/aru081
PMCID: PMC4095947  PMID: 25024638
costly signaling; formants; fundamental frequency; pitch; voice.
17.  Flexible compensation of uniparental care: female poison frogs take over when males disappear 
Behavioral Ecology  2015;26(4):1219-1225.
Lay Summary
Caring mothers step in for deadbeat dads. Flexible compensation has evolved as a countermeasure against reduced or lost parental care and is commonly found in biparental species. In the poison frog Allobates femoralis with obligatory male-only care, we show that females flexibly perform tadpole transport when males disappear. This demonstrates that compensatory flexibility also evolved in species with unisexual care, suggesting that parental care systems are more flexible than previously thought.
Parental care systems are shaped by costs and benefits to each sex of investing into current versus future progeny. Flexible compensatory parental care is mainly known in biparental species, particularly where parental desertion or reduction of care by 1 parent is common. The other parent can then compensate this loss by either switching parental roles and/or by increasing its own parental effort. In uniparental species, desertion of the caregiver usually leads to total brood loss. In the poison frog, Allobates femoralis, obligatory tadpole transport (TT) is generally performed by males, whereas females abandon their clutches after oviposition. Nevertheless, in a natural population we previously observed 7.8% of TT performed by females, which we could link to the absence of the respective fathers. In the following experiment, under laboratory conditions, all tested A. femoralis females flexibly took over parental duties, but only when their mates were removed. Our findings provide clear evidence for compensatory flexibility in a species with unisexual parental care. Contrary to the view of amphibian parental care as being stereotypical and fixed, these results demonstrate behavioral flexibility as an adaptive response to environmental and social uncertainty. Behavioral flexibility might actually represent a crucial step in the evolutionary transition from uniparental to biparental care in poison frogs. We suspect that across animal species flexible parental roles are much more common than previously thought and suggest the idea of a 3-dimensional continuum regarding flexibility, parental involvement, and timing, when thinking about the evolution of parental care.
doi:10.1093/beheco/arv069
PMCID: PMC4495760  PMID: 26167099
behavioral flexibility; compensation; partner removal; poison frog; tadpole transport; uniparental care.
18.  An arms race between producers and scroungers can drive the evolution of social cognition 
Behavioral Ecology  2014;25(3):487-495.
Lay Summary
Many cognitively advanced species live in groups, suggesting that this could have been important in the evolution of their high intelligence. We use theoretical analyses to show how the widespread behavior of foraging for food in groups can fuel an evolutionary arms race of intelligence. Strategies to outsmart others, stealing their food, and strategies to avoid being stolen from will continue to evolve so long as both sides use the same cognitive capacity in implementing these.
The “social intelligence hypothesis” states that the need to cope with complexities of social life has driven the evolution of advanced cognitive abilities. It is usually invoked in the context of challenges arising from complex intragroup structures, hierarchies, and alliances. However, a fundamental aspect of group living remains largely unexplored as a driving force in cognitive evolution: the competition between individuals searching for resources (producers) and conspecifics that parasitize their findings (scroungers). In populations of social foragers, abilities that enable scroungers to steal by outsmarting producers, and those allowing producers to prevent theft by outsmarting scroungers, are likely to be beneficial and may fuel a cognitive arms race. Using analytical theory and agent-based simulations, we present a general model for such a race that is driven by the producer–scrounger game and show that the race’s plausibility is dramatically affected by the nature of the evolving abilities. If scrounging and scrounging avoidance rely on separate, strategy-specific cognitive abilities, arms races are short-lived and have a limited effect on cognition. However, general cognitive abilities that facilitate both scrounging and scrounging avoidance undergo stable, long-lasting arms races. Thus, ubiquitous foraging interactions may lead to the evolution of general cognitive abilities in social animals, without the requirement of complex intragroup structures.
doi:10.1093/beheco/aru002
PMCID: PMC4014306  PMID: 24822021
game theory; intraspecific arms race; social foraging; social intelligence hypothesis.
19.  Rival male chemical cues evoke changes in male pre- and post-copulatory investment in a flour beetle 
Behavioral Ecology  2015;26(4):1021-1029.
Lay Summary
Males adjust both courtship effort and ejaculate expenditure when mating with females that are coated in the chemical cues of other males. Using a manipulative approach, we show that male flour beetles use the chemical cues of rival males left behind on virgin female cuticles to assess sperm competition risk. These cues do not make virgin females more chemically similar to mated females but appear to allow males to indirectly assess competition within the population.
Males can gather information on the risk and intensity of sperm competition from their social environment. Recent studies have implicated chemosensory cues, for instance cuticular hydrocarbons (CHCs) in insects, as a key source of this information. Here, using the broad-horned flour beetle (Gnatocerus cornutus), we investigated the importance of contact-derived rival male CHCs in informing male perception of sperm competition risk and intensity. We experimentally perfumed virgin females with male CHCs via direct intersexual contact and measured male pre- and post-copulatory investment in response to this manipulation. Using chemical analysis, we verified that this treatment engendered changes to perfumed female CHC profiles, but did not make perfumed females “smell” mated. Despite this, males responded to these chemical changes. Males increased courtship effort under low levels of perceived competition (from 1–3 rivals), but significantly decreased courtship effort as perceived competition rose (from 3–5 rivals). Furthermore, our measurement of ejaculate investment showed that males allocated significantly more sperm to perfumed females than to control females. Together, these results suggest that changes in female chemical profile elicited by contact with rival males do not provide males with information on female mating status, but rather inform males of the presence of rivals within the population and thus provide a means for males to indirectly assess the risk of sperm competition.
doi:10.1093/beheco/arv047
PMCID: PMC4495758  PMID: 26167098
chemical cues; cuticular hydrocarbons; ejaculate expenditure; Gnatocerus cornutus; sperm competition risk.
20.  Dynamics of among-individual behavioral variation over adult lifespan in a wild insect 
Behavioral Ecology  2015;26(4):975-985.
Lay Summary
Wild crickets show consistent patterns of behavior over their adult lifetimes and as they get older they become increasingly predictable. We tagged crickets and then periodically recaptured them and measured their behavior in the lab. This revealed that rather than variation in how age affects behavior, there were consistent patterns across the whole population. We do not have a situation where some crickets live fast and die young while others take it easy and slow.
Investigating patterns of among and within-individual trait variation in populations is essential to understanding how selection shapes phenotypes. Behavior is often the most flexible aspect of the phenotype, and to understand how it is affected by selection, we need to examine how consistent individuals are. However, it is not well understood whether among-individual differences tend to remain consistent over lifetimes, or whether the behavior of individuals relative to one another varies over time. We examined the dynamics of 4 behavioral traits (tendency to leave a refuge, shyness, activity, and exploration) in a wild population of field crickets (Gryllus campestris). We tagged individuals and then temporarily removed them from their natural environment and tested them under laboratory conditions. All 4 traits showed among-individual variance in mean levels of expression across the adult lifespan, but no significant differences in how rapidly expression changed with age. For all traits, among-individual variance increased as individuals got older. Our findings reveal seldom examined changes in variance components over the adult lifetime of wild individuals. Such changes will have important implications for the relationship between behavioral traits, life-histories, and fitness and the consequences of selection on wild individuals.
doi:10.1093/beheco/arv048
PMCID: PMC4495759  PMID: 26167097
animal personality; behavioral reaction norms; Gryllus; plasticity; wild crickets.
21.  Cicadas impact bird communication in a noisy tropical rainforest 
Behavioral Ecology  2015;26(3):839-842.
Lay Summary
When the cicada buzz starts up, bird song shuts down in a neotropical rainforest. When birds do sing at the same time as cicadas, their song bandwidths do not overlap.
Many animals communicate through acoustic signaling, and “acoustic space” may be viewed as a limited resource that organisms compete for. If acoustic signals overlap, the information in them is masked, so there should be selection toward strategies that reduce signal overlap. The extent to which animals are able to partition acoustic space in acoustically diverse habitats such as tropical forests is poorly known. Here, we demonstrate that a single cicada species plays a major role in the frequency and timing of acoustic communication in a neotropical wet forest bird community. Using an automated acoustic monitor, we found that cicadas vary the timing of their signals throughout the day and that the frequency range and timing of bird vocalizations closely track these signals. Birds significantly avoid temporal overlap with cicadas by reducing and often shutting down vocalizations at the onset of cicada signals that utilize the same frequency range. When birds do vocalize at the same time as cicadas, the vocalizations primarily occur at nonoverlapping frequencies with cicada signals. Our results greatly improve our understanding of the community dynamics of acoustic signaling and reveal how patterns in biotic noise shape the frequency and timing of bird vocalizations in tropical forests.
doi:10.1093/beheco/arv018
PMCID: PMC4433330  PMID: 26023277
acoustic partitioning; bird vocalizations.
22.  Mate choice and genetic monogamy in a biparental, colonial fish 
Behavioral Ecology  2015;26(3):782-788.
Lay Summary
We report a rare case of genetic monogamy in a biparental fish species. Males and females paired assortatively by size, which is compatible with mutual mate choice. Mate choice in monogamous species is interesting because both sexes provide essential parental care, making males, as well as females, choosy. Social monogamy in the form of biparental care is well known from a variety of species, but uncommon in fish.
In socially monogamous species, in which both sexes provide essential parental care, males as well as females are expected to be choosy. Whereas hundreds of studies have examined monogamy in biparental birds, only several such studies exist in fish. We examined mate choice in the biparental, colonial cichlid fish Neolamprologus caudopunctatus in Lake Tanganyika, Zambia. We genotyped more than 350 individuals at 11 microsatellite loci to investigate their mating system. We found no extrapair paternity, identifying this biparental fish as genetically monogamous. Breeders paired randomly according to their genetic similarity, suggesting a lack of selection against inbreeding avoidance. We further found that breeders paired assortatively by body size, a criterion of quality in fish, suggesting mutual mate choice. In a subsequent mate preference test in an aquarium setup, females showed a strong preference for male size by laying eggs near the larger of 2 males in 13 of 14 trials.
doi:10.1093/beheco/arv011
PMCID: PMC4433329  PMID: 26023276
assortative mating; cichlid; colony; extrapair paternity.
23.  Populations, pools, and peccaries: simulating the impact of ecosystem engineers on rainforest frogs 
Behavioral Ecology  2015;26(2):340-349.
Lay Summary
Peccary wallows and footprints are important breeding pools for rainforest frogs. We performed a resource supplementation experiment with artificial pools, simulating peccary actions, in a population of the poison frog Allobates femoralis. The population almost doubled resulting from increased local reproduction, but not from immigration. These findings demonstrate the importance of “ecosystem engineers,” such as peccaries, for other species, the frogs. Our results also indicate that human engineering may help to protect amphibian populations.
“Ecosystem engineering” describes habitat alteration by an organism that affects another organism; such nontrophic interactions between organisms are a current focus in ecological research. Our study quantifies the actual impact an ecosystem engineer can have on another species by using a previously identified model system—peccaries and rainforest frogs. In a 4-year experiment, we simulated the impact of peccaries on a population of Allobates femoralis (Dendrobatidae) by installing an array of artificial pools to mimic a forest patch modified by peccaries. The data were analyzed using a gradual before-after control-impact (gBACI) model. Following the supplementation, population size almost doubled as a result of increased autochthonous recruitment driven by a higher per-capita reproduction of males and a higher proportion of reproducing females. The effect was evenly distributed across the population. The differential response of males and females reflects the reproductive behavior of A. femoralis, as only the males use the aquatic sites for tadpole deposition. Our study shows that management and conservation must consider nontrophic relationships and that human “ecosystem engineering” can play a vital role in efforts against the “global amphibian decline.”
doi:10.1093/beheco/aru243
PMCID: PMC4374131  PMID: 25825586
Allobates femoralis; Dendrobatidae; ecosystem engineering; nontrophic interaction; population manipulation experiment; reproductive resource supplementation.
24.  A larger brain confers a benefit in a spatial mate search learning task in male guppies 
Behavioral Ecology  2014;26(2):527-532.
Lay Summary
Does a large brain make you smarter? If you are a guppy male searching for a female in a maze, it does. The association between brain size and smartness is a debated issue, largely due to the lack of experimental data. We compared guppies artificially bred for large and small brains and found that large-brained males learned the route through a spatial maze faster. These results thus support a link between brain size and smartness.
Brain size varies dramatically among vertebrates, and selection for increased cognitive abilities is thought to be the key force underlying the evolution of a large brain. Indeed, numerous comparative studies suggest positive relationships between cognitively demanding aspects of behavior and brain size controlled for body size. However, experimental evidence for the link between relative brain size and cognitive ability is surprisingly scarce and to date stems from a single study on brain size selected guppies (Poecilia reticulata), where large-brained females were shown to outperform small-brained females in a numerical learning assay. Because the results were inconclusive for males in that study, we here use a more ecologically relevant test of male cognitive ability to investigate whether or not a relatively larger brain increases cognitive ability also in males. We compared mate search ability of these artificially selected large- and small-brained males in a maze and found that large-brained males were faster at learning to find a female in a maze. Large-brained males decreased the time spent navigating the maze faster than small-brained males and were nearly twice as fast through the maze after 2 weeks of training. Our results support that relatively larger brains are better also for males in some contexts, which further substantiates that variation in vertebrate brain size is generated through the balance between energetic costs and cognitive benefits.
doi:10.1093/beheco/aru227
PMCID: PMC4374130  PMID: 25825587
brain size; cognition; guppy; maze.
25.  Flight calls signal group and individual identity but not kinship in a cooperatively breeding bird 
Behavioral Ecology  2013;24(6):1279-1285.
Lay Summary
Ants are models of conflict, generally working together but at the same time competing over individual fitness. We show that ant larvae compete by cannibalizing eggs, which increases their survival. Male larvae are particularly selfish, and eat eggs three times more often than females. Larvae also discriminate between sibling and alien eggs, which suggests that they can react to chemical recognition cues. Remarkably, ant larvae thus possess the power to act in social conflict.
In many complex societies, intricate communication and recognition systems may evolve to help support both direct and indirect benefits of group membership. In cooperatively breeding species where groups typically comprise relatives, both learned and innate vocal signals may serve as reliable cues for kin recognition. Here, we investigated vocal communication in the plural cooperatively breeding superb starling, Lamprotornis superbus, where flight calls—short, stereotyped vocalizations used when approaching conspecifics—may communicate kin relationships, group membership, and/or individual identity. We found that flight calls were most similar within individual repertoires but were also more similar within groups than within the larger population. Although starlings responded differently to playback of calls from their own versus other neighboring and distant social groups, call similarity was uncorrelated with genetic relatedness. Additionally, immigrant females showed similar patterns to birds born in the study population. Together, these results suggest that flight calls are learned signals that reflect social association but may also carry a signal of individuality. Flight calls, therefore, provide a reliable recognition mechanism for groups and may also be used to recognize individuals. In complex societies comprising related and unrelated individuals, signaling individuality and group association, rather than kinship, may be a route to cooperation.
doi:10.1093/beheco/art062
PMCID: PMC3796708  PMID: 24137044
cooperative breeding; flight call; individual recognition; kin recognition; Lamprotornis superbus; vocal communication.

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