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The expression of sexually selected traits, such as ornaments or body coloration, is often influenced by environmental conditions. While such phenotypic plasticity is often thought to precede evolutionary change, plasticity itself can also be a target of selection. However, the selective forces supporting the evolution and persistence of plasticity in sexual traits are often unclear. Using the cichlid fish Astatotilapia burtoni, we show that variation in the level of mate competition may promote plasticity in body coloration. In this species, males can change between yellow and blue colour. We found that experimentally increased competition over mating territories led to a higher proportion of males expressing the yellow phenotype. The expression of yellow coloration was found to be beneficial because yellow males won more staged dyadic contests and exhibited a lower level of oxidative stress than blue males. However, females were more likely to spawn with blue males in mate choice experiments, suggesting that expression of blue coloration is sexually more attractive. The ability to adjust colour phenotype according to the local competitive environment could therefore promote the persistence of plasticity in coloration.
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Ciclídeos , Fenótipo , Pigmentação , Animais , Ciclídeos/fisiologia , Masculino , Feminino , Preferência de Acasalamento Animal , Seleção Sexual , Evolução BiológicaRESUMO
An individual's social environment can have widespread effects on their physiology, including effects on oxidative stress and hormone levels. Many studies have suggested that variation in oxidative stress experienced by individuals of different social statuses might be due to endocrine differences, however, few studies have evaluated this hypothesis. Here, we assessed whether a suite of markers associated with oxidative stress in different tissues (blood/plasma, liver, and gonads) had social status-specific relationships with circulating testosterone or cortisol levels in males of a cichlid fish, Astatotilapia burtoni. Across all fish, blood DNA damage (a global marker of oxidative stress) and gonadal synthesis of reactive oxygen species [as indicated by NADPH-oxidase (NOX) activity] were lower when testosterone was high. However, high DNA damage in both the blood and gonads was associated with high cortisol in subordinates, but low cortisol in dominants. Additionally, high cortisol was associated with greater production of reactive oxygen species (greater NOX activity) in both the gonads (dominants only) and liver (dominants and subordinates). In general, high testosterone was associated with lower oxidative stress across both social statuses, whereas high cortisol was associated with lower oxidative stress in dominants and higher oxidative stress in subordinates. Taken together, our results show that differences in the social environment can lead to contrasting relationships between hormones and oxidative stress.
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Ciclídeos , Hidrocortisona , Animais , Masculino , Ciclídeos/fisiologia , Status Social , Espécies Reativas de Oxigênio , Estresse Oxidativo , TestosteronaRESUMO
3D bioprinting is a powerful fabrication technique in biomedical engineering, which is currently limited by the number of available materials that meet all physicochemical and cytocompatibility requirements for biomaterial inks. Inspired by the key role of coacervation in the extrusion and spinning of many natural materials, hyaluronic acid-chitosan complex coacervates are proposed here as tunable biomaterial inks. Complex coacervates are obtained through an associative liquid-liquid phase separation driven by electrostatic attraction between oppositely charged macromolecules. They offer bioactive properties and facile modulation of their mechanical properties through mild physicochemical changes in the environment, making them attractive for 3D bioprinting. Fine-tuning the salt concentration, pH, and molecular weight of the constituent polymers results in biomaterial inks that are printable in air and water. The biomaterial ink, initially a viscoelastic fluid, transitions into a viscoelastic solid upon printing due to dehydration (for printing in air) or due to a change in pH and ionic composition (for printing in solution). Consequently, scaffolds printed using the complex coacervate inks are stable without the need for post-printing processing. Fabricated cell culture scaffolds are cytocompatible and show long-term topological stability. These results pave the way to a new class of easy-to-handle tunable biomaterials for biofabrication.
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Bioimpressão , Tinta , Bioimpressão/métodos , Impressão Tridimensional , Reologia , Materiais Biocompatíveis/química , Alicerces Teciduais/química , Engenharia Tecidual/métodos , Hidrogéis/químicaRESUMO
Investment in current reproduction may negatively influence subsequent fitness. Oxidative stress has been proposed as a potential mediator of this trade-off between current and future reproductive success. However, evidence of reproduction causing oxidative stress is limited, possibly owing to compensatory mechanisms that counteract oxidative insults. Here we test the idea that organisms protect against oxidative challenges through a dynamic interaction between behavioural and physiological adjustments at different stages of reproduction. To test this idea, we manipulated maternal care in the mouthbrooding cichlid fish Astatotilapia burtoni by allowing females to continue care (brooders) or by preventing care (non-brooders). We found that brooders depleted the pool of antioxidants as brood care progressed; however, we only observed increased oxidative DNA damage at the early stage of care relative to non-brooders, possibly owing to upregulated antioxidant protection during later stages of care. Most brooders adjusted parental investment by consuming some of their offspring during mouthbrooding. Intriguingly, the level of filial cannibalism was positively related to liver antioxidant function. These changes in antioxidant function and filial cannibalism allow parents to manage the cost of reproduction and are important for our understanding of how oxidative stress mediates life-history trade-offs.
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Canibalismo , Ciclídeos , Feminino , Animais , Antioxidantes/metabolismo , Reprodução/fisiologia , Estresse OxidativoRESUMO
Oxidative stress is a potential cost of social dominance and reproduction, which could mediate life history trade-offs between current and future reproductive fitness. However, the evidence for an oxidative cost of social dominance and reproduction is mixed, in part because organisms have efficient protective mechanisms that can counteract oxidative insults. Further, previous studies have shown that different aspects of oxidative balance, including oxidative damage and antioxidant function, vary dramatically between tissue types, yet few studies have investigated oxidative cost in terms of interconnectedness and coordination within the system. Here, we tested whether dominant and subordinate males of the cichlid Astatotilapia burtoni differ in integration of different components of oxidative stress. We assessed 7 markers of oxidative stress, which included both oxidative damage and antioxidant function in various tissue types (total of 14 measurements). Across all oxidative stress measurements, we found more co-regulated clusters in dominant males, suggesting that components of oxidative state are more functionally integrated in dominant males than they are in subordinate males. We discuss how a high degree of functional integration reflects increased robustness or efficiency of the system (e.g. increased effectiveness of antioxidant machinery in reducing oxidative damage), but we also highlight potential costs (e.g. activation of cytoprotective mechanisms may have unwanted pleiotropic effects). Overall, our results suggest that quantifying the extent of functional integration across different components of oxidative stress could reveal insights into the oxidative cost of important life history events.
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Ciclídeos , Hierarquia Social , Animais , Antioxidantes , Ciclídeos/fisiologia , Masculino , Estresse Oxidativo , Reprodução/fisiologia , Predomínio SocialRESUMO
Although biological systems are more complex and can actively respond to their environment, an effective entry point to the development of a universal theory of biological stress is the physical concepts of stress and strain. If you apply stress to the end of a beam of steel, the strain will accumulate within that steel beam. If the stress is weak then the strain will disappear when the force is removed and the beam will return to its original state of form and functionality. If the stress is more severe, then the strain becomes permanent and the beam will be deformed, potentially losing some degree of functionality. In extremely stressful situations, the beam will break and lose most or all of its original functional capabilities. Although this stress-strain theory applies to the abiotic, stress and strain are also rules of life and directly relate to the form and function of living organisms. The main difference is that life can react and adjust to stress and strain to maintain homeostasis within a range of limits. Here, we summarize the rules of stress and strain in living systems ranging from microbes to multicellular organisms to ecosystems with the goal to identify common features that may underlie a universal biological theory of stress. We then propose to establish a range of experimental, observational, and analytical approaches to study stress across scales, including synthetic microbial communities that mimic many of the essential characteristics of living systems, thereby enabling a universal theory of biological stress to be experimentally validated without the constraints of timescales, ethics, or cost found when studying other species or scales of life. Although the range of terminology, theory, and methodology used to study stress and strain across the scales of life presents a formidable challenge to creating a universal theory of biological stress, working toward such a theory that informs our understanding of the simultaneous and interconnected unicellular, multicellular, organismal, and ecosystem stress responses is critical as it will improve our ability to predict how living systems respond to change, thus informing solutions to current and future environmental and human health challenges.
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Ecossistema , Estresse Fisiológico , Animais , HomeostaseRESUMO
Animals that live in groups often form hierarchies in which an individual's behaviour and physiology varies based on their social rank. Occasionally, a subordinate can ascend into a dominant position and the ascending individual must make rapid behavioural and physiological adjustments to solidify their dominance. These periods of social transition and instability can be stressful and ascending individuals often incur large metabolic costs that could influence their oxidative status. Most previous investigations examining the link between oxidative status and the social environment have done so under stable social conditions and have evaluated oxidative status in a single tissue. Therefore, evaluations of how oxidative status is regulated across multiple tissues during periods of social flux would greatly enhance our understanding of the relationship between oxidative status and the social environment. Here, we assessed how antioxidant capacity in three tissues (brain, gonad, and muscle) varied among dominant, subordinate, and ascending males of the group-living cichlid fish, Neolamprologus pulcher. Antioxidant capacity in the brain and muscle of ascending males was intermediate to that of dominant (highest levels) and subordinate males (lowest levels) and correlated with differences in social and locomotor behaviours, respectively. Gonad antioxidant capacity was lower in ascending males than in dominant males. However, gonad antioxidant capacity was positively correlated with the size of ascending males' gonads suggesting that ascending males may increase gonad antioxidant capacity as they develop their gonads. Overall, our results highlight the widespread physiological consequences of social ascension and emphasize the importance of tissue-specific measures of oxidative status.
Assuntos
Antioxidantes , Ciclídeos , Animais , Ciclídeos/fisiologia , Gônadas/fisiologia , Hierarquia Social , Masculino , MúsculosRESUMO
In many animal societies, dominant individuals have priority access to resources. However, defending high rank can be costly, especially in unstable social hierarchies where there is more intense competition. Oxidative stress has been proposed as a potential cost of social dominance, but few studies have examined this cost in relation to social stability. We studied the cost of social dominance in the cichlid fish Astatotilapia burtoni by manipulating social stability among males in replicate naturalistic communities for 22â weeks. We found that our social stability treatment influenced status-specific patterns in 3 out of 6 measurements of oxidative stress. Specifically, dominant males experienced increased plasma oxidative damage (measured as reactive oxygen metabolites, ROMs) compared with subordinate males in stable hierarchies only. Subordinate males in unstable hierarchies had higher ROMs than their stable community counterparts, but we found no effect of social stability treatment for dominant males. However, dominant males tended to have reduced total antioxidant capacity (TAC) in the liver when compared with subordinate males in unstable hierarchies, suggesting that the cost of social dominance is higher in unstable hierarchies. There were no effects of status and treatment on gonad TAC, muscle TAC or oxidative DNA damage. We conclude that the stability of the social environment influences the relative cost of social dominance in a tissue- and marker-specific manner.
Assuntos
Ciclídeos , Animais , Hierarquia Social , Humanos , Masculino , Estresse Oxidativo , Predomínio Social , Meio SocialRESUMO
Group living confers many benefits while simultaneously exposing group members to intense competition. An individual's rise to prominence within a group may conflict with the overall functioning of the group. There is therefore a complex and dynamic relationship between the behavioral displays that directly benefit an individual, the consequences of these actions for the community, and how they feed back on individual-level fitness. We used a network analysis approach to study the link between behavior, social stability, and steroid hormone levels in replicate communities of the cichlid fish, Astatotilapia burtoni, which live in social groups with a dominance hierarchy. We demonstrate that individual behavior can have direct and indirect effects on the behavior of others while also affecting group characteristics. Our results show that A. burtoni males form stable social networks, where dominant individuals act as hubs for social interactions. However, there was variation in the temporal stability in these networks, and this variation in stability impacted hormone levels. Dominant males had higher testosterone levels, however, the differences in testosterone levels between dominant and subordinate males were greatest in stable communities. In sum, our analyses provide novel insights into the processes by which individual and community properties interact.
Assuntos
Ciclídeos , Agressão , Animais , Hierarquia Social , Hormônios , Humanos , Masculino , Comportamento Social , Predomínio Social , Rede SocialRESUMO
Natural selection has been shown to drive population differentiation and speciation. The role of sexual selection in this process is controversial; however, most of the work has centered on mate choice while the role of male-male competition in speciation is relatively understudied. Here, we outline how male-male competition can be a source of diversifying selection on male competitive phenotypes, and how this can contribute to the evolution of reproductive isolation. We highlight how negative frequency-dependent selection (advantage of rare phenotype arising from stronger male-male competition between similar male phenotypes compared with dissimilar male phenotypes) and disruptive selection (advantage of extreme phenotypes) drives the evolution of diversity in competitive traits such as weapon size, nuptial coloration, or aggressiveness. We underscore that male-male competition interacts with other life-history functions and that variable male competitive phenotypes may represent alternative adaptive options. In addition to competition for mates, aggressive interference competition for ecological resources can exert selection on competitor signals. We call for a better integration of male-male competition with ecological interference competition since both can influence the process of speciation via comparable but distinct mechanisms. Altogether, we present a more comprehensive framework for studying the role of male-male competition in speciation, and emphasize the need for better integration of insights gained from other fields studying the evolutionary, behavioral, and physiological consequences of agonistic interactions.
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The color of an athlete's uniform may have an effect on psychological functioning and consequently bias the chances of winning contests in sport competition. Several studies reported a winning bias for judo athletes wearing a blue outfit relative to those wearing a white outfit. However, we argue there is no winning bias and that previous studies were confounded and based on small and specific data sets. We tested whether blue biases winning in judo using a very extensive judo data set (45,874 contests from all international judo tournaments between 2008 and 2014). In judo, the first called athlete for the fight used to wear the blue judogi but this was changed to the white judogi in 2011. This switch enabled us to compare the win bias before and after this change to isolate the effect of the color of the judogi. We found a significant win bias for the first called athlete, but this effect was not significantly related to the color of the judogi. The lack of a significant win effect of judogi color suggests that blue does not bias winning in judo, and that the blue-white pairing ensures an equal level of play. Our study shows the importance of thoroughly considering alternative explanations and using extensive datasets in color research in sports and psychology.
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To facilitate the ongoing transition toward a circular economy, the availability of renewable materials for additive manufacturing becomes increasingly important. Here, we report the successful fabrication of complex shaped prototypes from biobased acrylate photopolymer resins, employing a commercial stereolithography apparatus (SLA) 3D printer. Four distinct resins with a biobased content ranging from 34 to 67% have been developed. All formulations demonstrated adequate viscosity and were readily polymerizable by the UV-laser-based SLA process. Increasing the double-bond concentration within the resin results in stiff and thermally resilient 3D printed products. High-viscosity resins lead to high-resolution prototypes with a complex microarchitecture and excellent surface finishing, comparable to commercial nonrenewable resins. These advances can facilitate the wide application of biobased resins for construction of new sustainable products via stereolithographic 3D printing methods.
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The melanocortin system is a neuroendocrine system that regulates a range of physiological and behavioural processes. We examined the extent to which the melanocortin system simultaneously regulates colour and behaviour in the cichlid fish Astatotilapia burtoni We found that yellow males are more aggressive than blue males, in line with previous studies. We then found that exogenous α-melanocyte-stimulating hormone (α-MSH) increases yellowness of the body and dispersal of xanthophore pigments in both morphs. However, α-MSH had a morph-specific effect on aggression, with only blue males showing an increase in the rate of aggression. Exogenous agouti signalling peptide (ASIP), a melanocortin antagonist, did not affect coloration but reduced the rate of aggression in both colour morphs. Blue males had higher cortisol levels than yellow males. Neural gene expression of melanocortin receptors (mcr) and ligands was not differentially regulated between colour morphs. In the skin, however, mc1r and pro-opiomelanocortin (pomc) ß were upregulated in blue males, while asip 1 was upregulated in yellow males. The effects of α-MSH on behaviour and body coloration, combined with morph-specific regulation of the stress response and the melanocortin system, suggest that the melanocortin system contributes to the polymorphism in behaviour and coloration in A. burtoni.
Assuntos
Comportamento Animal , Ciclídeos/fisiologia , Melanocortinas/fisiologia , Pigmentação , Receptores de Melanocortina/fisiologia , Agressão , Animais , Cor , Proteínas de Peixes/fisiologia , Masculino , Pró-Opiomelanocortina/fisiologia , Comportamento SocialRESUMO
The main motivation for development of biobased polymers was their biodegradability, which is becoming important due to strong public concern about waste. Reflecting recent changes in the polymer industry, the sustainability of biobased polymers allows them to be used for general and engineering applications. This expansion is driven by the remarkable progress in the processes for refining biomass feedstocks to produce biobased building blocks that allow biobased polymers to have more versatile and adaptable polymer chemical structures and to achieve target properties and functionalities. In this review, biobased polymers are categorized as those that are: (1) upgrades from biodegradable polylactides (PLA), polyhydroxyalkanoates (PHAs), and others; (2) analogous to petroleum-derived polymers such as bio-poly(ethylene terephthalate) (bio-PET); and (3) new biobased polymers such as poly(ethylene 2,5-furandicarboxylate) (PEF). The recent developments and progresses concerning biobased polymers are described, and important technical aspects of those polymers are introduced. Additionally, the recent scientific achievements regarding high-spec engineering-grade biobased polymers are presented.
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The detection of nuclear radiation necessitates the availability of new generations of tunable blue emitting fluorophores with high emission quantum yields. Here we show that pyridyl-1,2,4-triazole based diphenyl boron complexes can provide highly tuneable emission through facile modification of the C5 position of the 1,2,4-triazolato ring. The series of complexes prepared show a wide range of emission from near-UV to green, enabling fine control over the spectral overlap with detectors used in scintillator technology.
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The melanocortin system is a neuroendocrine machinery that has been associated with phenotypic diversification in a number of vertebrate lineages. Central to the highly pleiotropic melanocortin system is the pro-opiomelanocortin (pomc) gene family, a family of pre-prohormones that each give rise to melanocyte stimulating hormone (MSH), adrenocorticotropic releasing hormone (ACTH), ß-lipotropin hormone, and ß-endorphin. Here we examine the structure, tissue expression profile, and pattern of cis transcriptional regulation of the three pomc paralogs (α1, α2, and ß) in the model cichlid fish Astatotilapia burtoni and other cichlids, teleosts, and mammals. We found that the hormone-encoding regions of pomc α1, pomc α2 and pomc ß are highly conserved, with a few notable exceptions. Surprisingly, the pomc ß gene of cichlids and pomacentrids (damselfish) encodes a novel melanocortin peptide, ε-MSH, as a result of a tandem duplication of the segment encoding ACTH. All three genes are expressed in the brain and peripheral tissues, but pomc α1 and α2 show a more spatially restricted expression profile than pomc ß. In addition, the promoters of each pomc gene have diverged in nucleotide sequence, which may have facilitated the diverse tissue-specific expression profiles of these paralogs across species. Increased understanding of the mechanisms regulating pomc gene expression will be invaluable to the study of pomc in the context of phenotypic evolution.
Assuntos
Evolução Biológica , Proteínas de Peixes/genética , Regulação da Expressão Gênica , Família Multigênica , Pró-Opiomelanocortina/genética , Elementos Reguladores de Transcrição/genética , Hormônio Adrenocorticotrópico/genética , Sequência de Aminoácidos , Animais , Peixes/genética , Hormônios Estimuladores de Melanócitos/genética , Dados de Sequência Molecular , Especificidade de Órgãos , Filogenia , Regiões Promotoras Genéticas/genética , Homologia de Sequência de AminoácidosRESUMO
Studies on birds and mammals indicate that sexual traits may signal superior health because active immunity, like inflammatory responses to infections, is suppressive to the production of androgens that facilitate the expression of these traits. Here we test this possible pathway for honest signaling in a teleost species, Sarotherodon galilaeus, by activating the immune system with sheep red blood cells (SRBC), which is a non-pathogenic T- and B-cell stimulating antigen. Two weeks after the start of treatment adult males injected with SRBC showed a significant increase in antibody production in comparison with control males. The variation in specific antibody production was negatively related with variation in both testosterone and 11-ketotestosterone levels. This suggests that investment in immune protection is incompatible with increased activity of the hypothalamic-pituitary-gonadal axis. However, opposite to our expectation no difference in androgen levels was found between placebo and SRBC treatment suggesting that immune activation did not cause androgen suppression in our studied species.
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Androgênios/sangue , Androgênios/imunologia , Ciclídeos/imunologia , Sistema Imunitário/imunologia , Adjuvantes Imunológicos , Animais , Anticorpos/sangue , Ciclídeos/sangue , Masculino , Testosterona/análogos & derivados , Testosterona/sangueRESUMO
The diverse cichlid species flocks of the East African lakes provide a classical example of adaptive radiation. Territorial aggression is thought to influence the evolution of phenotypic diversity in this system. Most vertebrates mount hormonal (androgen, glucocorticoid) responses to a territorial challenge. These hormones, in turn, influence behavior and multiple aspects of physiology and morphology. Examining variation in competition-induced hormone secretion patterns is thus fundamental to an understanding of the mechanisms of phenotypic diversification. We test here the hypothesis that diversification in male aggression has been accompanied by differentiation in steroid hormone levels. We studied two pairs of sibling species from Lake Victoria belonging to the genera Pundamilia and Mbipia. The two genera are ecologically differentiated, while sibling species pairs differ mainly in male color patterns. We found that aggression directed toward conspecific males varied between species and across genera: Pundamilia nyererei males were more aggressive than Pundamilia pundamilia males, and Mbipia mbipi males were more aggressive than Mbipia lutea males. Males of both genera exhibited comparable attack rates during acute exposure to a novel conspecific intruder, while Mbipia males were more aggressive than Pundamilia males during continuous exposure to a conspecific rival, consistent with the genus difference in feeding ecology. Variation in aggressiveness between genera, but not between sibling species, was reflected in androgen levels. We further found that M. mbipi displayed lower levels of cortisol than M. lutea. Our results suggest that concerted divergence in hormones and behavior might play an important role in the rapid speciation of cichlid fishes.
Assuntos
Ciclídeos/fisiologia , Comportamento Competitivo/fisiologia , Hormônios/sangue , Comportamento Social , Agressão/fisiologia , Comportamento Agonístico/fisiologia , Análise de Variância , Animais , Feminino , Hibridização Genética , Hidrocortisona/sangue , Masculino , Pigmentação , Especificidade da Espécie , Esteroides/sangue , Territorialidade , Testosterona/análogos & derivados , Testosterona/sangueRESUMO
Previous winning experience increases the probability of winning a subsequent contest. However, it is not clear whether winning probability is affected only by the outcome of the contest (winning or losing) or whether fighting experience itself is also sufficient to induce this effect. We investigated this question in the East African cichlid fish Pundamilia spec. To create an unresolved conflict we allowed males to fight their own mirror image prior to a real fight against a size-matched non-mirror-stimulated control male. When males fight their own mirror image, the image's response corresponds to the action of the focal animal, creating symmetrical fighting conditions without the experience of losing or winning. We found that mirror-stimulated males were more likely to win an ensuing contest than control males. Interestingly, in this species mirror stimulation also induced an increase in circulating androgens, which is consistent with the hypothesis that stimulation of these sex steroids during aggressive encounters may prepare the animal for subsequent encounters. Our results suggest that fighting experience alone coupled with an androgen response, increases the likelihood of winning, even in the absence of a winning experience.
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Agressão/fisiologia , Ciclídeos/fisiologia , Comportamento Competitivo/fisiologia , Conflito Psicológico , Dominação-Subordinação , Probabilidade , Animais , Comportamento Animal , Masculino , Estimulação Luminosa , Prática Psicológica , Fatores de TempoRESUMO
It has been suggested that intrasexual competition can be a source of negative frequency-dependent selection, causing agonistic character displacement and facilitating speciation and coexistence of (sibling) species. In this paper we synthesise the evidence that male-male and female-female competition contributes to cichlid diversification, showing that competition is stronger among same-coloured individuals than those with different colours. We argue that intrasexual selection is more complex because there are several examples where males do not bias aggression towards their own type. In addition, sibling species or colour morphs often show asymmetric dominance relationships. We briefly discuss potential mechanisms that might promote the maintenance of covariance between colour and aggression-related traits even in the face of gene-flow. We close by proposing several avenues for future studies that might shed more light on the role of intrasexual competition in cichlid diversification.