chemical communication

化学通讯
  • 文章类型: Journal Article
    研究表明,男性体味在女性的择偶过程中起着重要作用,嗅觉能力与女性的性功能有关。尚不清楚的是,在亲密活动中,伴侣的气味类型实际上塑造了女性的体验。因此,这项研究探索了女性与伴侣的各种气味相关的经历,并调查了它们如何影响女性的亲密和性接触。我们对20名单身女性和20名长期恋爱关系的女性进行了半结构化的个人访谈。主题分析揭示了伴侣的四种关键自然气味类型:体臭,汗水,生殖器气味,精液气味。Further,我们已经确定了三种主要的香味气味类型(古龙水,沐浴露,和洗衣代理),并调查了他们在两个亲密(拥抱,接吻,拥抱,并排躺着)和性(性交,口交,射精)上下文。双方的自然气味和香味影响女性的情绪状态(从愉快到不愉快)和行为反应(从方法到避免伴侣)。女性的气味感知通常取决于上下文,因此,即使大多数是负面感知的气味(例如,精液,生殖器气味)通常被接受为性接触的一部分。最后,女性的感知受到伴侣特定汗水的负面影响(工作日后,锻炼,或当伴侣生病时)在亲密接触中。我们的结果强调了伴侣气味感知的复杂性和个体间的变异性。
    Research shows that male body odor plays an important role in women\'s mate choice and that olfactory abilities are associated with women\'s sexual functioning. What remains unclear is what types of partner\'s odor actually shape women\'s experience during intimate activities. This study therefore explored women\'s experience associated with the partner\'s various odors and investigated how they affect women\'s intimate and sexual encounters. We performed semi-structured individual interviews with 20 single women and 20 women in a long-term relationship. Thematic analysis revealed four key natural odor types of the partner: body odor, sweat, genital odor, and semen odor. Further, we have identified three main types of fragrance odor (cologne, shower gel, and laundry agents) and investigated their perception in both intimate (hugging, kissing, cuddling, lying side by side) and sexual (intercourse, oral sex, ejaculation) contexts. Both partner\'s natural odor and fragrance affected women\'s emotional state (ranging from pleasant to unpleasant) and behavioral response (ranging from approach to avoidance of partner). Women\'s odor perception was frequently context-dependent, so that even mostly negatively perceived odors (e.g., semen, genital odor) were often accepted as part of sexual encounter. Finally, women\'s perception was negatively modified by partner\'s specific sweat (after workday, workout, or when the partner is ill) during intimate encounters. Our results highlight the complexity and interindividual variability of partner\'s odor perception.
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  • 文章类型: Journal Article
    社交动物需要跟踪他们群体中的其他个体,以便能够相应地调整他们的行为并促进群体凝聚力。这种识别能力因物种而异,并受到学习和记忆等认知能力的影响。在爬行动物中,尤其是Squamates(蜥蜴,蛇,和蠕虫蜥蜴),化学通信是地域性的关键,繁殖,和其他社会互动。然而,这些社会互动背后的认知过程仍未得到充分研究。在我们的研究中,我们研究了雄性和雌性Tokay壁虎(Gekkogecko)在化学上区分熟悉和不熟悉的交配伴侣的能力。我们的研究结果表明,两性都可以做出这种区分,雄性对熟悉伴侣的气味有更多的反应,女性对陌生伴侣的反应更多。蜥蜴在分离后保持了两到三周的歧视能力,但最多不到六周。这项研究强调了使用气味作为社会刺激来调查蜥蜴的社会认知的功效,一个有希望的途径来更好地了解这些动物的社会认知。
    Social animals need to keep track of other individuals in their group to be able to adjust their behaviour accordingly and facilitate group cohesion. This recognition ability varies across species and is influenced by cognitive capacities such as learning and memory. In reptiles, particularly Squamates (lizards, snakes, and worm lizards), chemical communication is pivotal for territoriality, reproduction, and other social interactions. However, the cognitive processes underlying these social interactions remain understudied. In our study, we examined the ability of male and female Tokay geckos (Gekko gecko) to chemically differentiate familiar and unfamiliar mating partners. Our findings suggest that both sexes can make this distinction, with males responding more to the odour of a familiar mate, and females responding more to unfamiliar mates. The lizards maintained their discriminatory abilities for two to three weeks but not up to six weeks after separation. This research highlights the efficacy of using odours as social stimuli for investigating social cognition in lizards, a promising avenue to better understand social cognition in these animals.
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  • 文章类型: Journal Article
    电生物膜,在生物电化学系统的同步废水处理和能量回收方面引起了相当大的关注,受化学通讯和钾通道介导的电信号调节。然而,这两种沟通途径如何相互作用尚未得到彻底研究。本研究首先探讨了化学通讯的作用,包括细胞内双-(3'-5')-环二聚一磷酸鸟苷(c-di-GMP)和胞外N-酰基高丝氨酸内酯(AHL)介导的群体感应,通过转录组学和代谢组学的综合分析,在电生物膜形成中。电信号的中断抑制了硫化焦菌生物膜的形成和电活性,这主要归因于生物膜活力和细胞外蛋白/多糖比例的降低。通过转录组学分析,编码c-di-GMP和AHL合成的基因的表达水平上调,通过代谢组学分析,N-丁酰基-L-高丝氨酸内酯的分泌增加证实了电信号中断下化学通讯的增强,从而表明不同信号通路之间存在代偿机制。此外,蛋白质相互作用网络显示了不同信号通路的趋同,与c-di-GMP相关的基因充当中心桥梁。这项研究强调了不同信号通路的相互作用,特别是c-di-GMP信号对不利外部压力的恢复能力,从而为在实际应用中在不利条件下促进电生物膜的形成奠定了基础。
    Electrogenic biofilms, which have attracted considerable attention in simultaneous wastewater treatment and energy recovery in bioelectrochemical systems, are regulated by chemical communication and potassium channel-mediated electrical signaling. However, how these two communication pathways interact with each other has not been thoroughly investigated. This study first explored the roles of chemical communication, including intracellular bis-(3\'-5\')-cyclic dimeric guanosine monophosphate (c-di-GMP) and extracellular N-acyl-homoserine lactone (AHL)-mediated quorum sensing, in electrogenic biofilm formation through an integrated analysis of transcriptomics and metabolomics. Electrical signaling disruption inhibited the formation and electroactivity of Geobacter sulfurreducens biofilm, which was mainly ascribed to the reduction in biofilm viability and extracellular protein/polysaccharide ratio. The upregulation of expression levels of genes encoding c-di-GMP and AHL synthesis by transcriptomic analysis, and the increased secretion of N-butanoyl-L-homoserine lactone by metabolomic analysis confirmed the enhancement of chemical communication under electrical signaling disruption, thus indicating a compensatory mechanism among different signaling pathways. Furthermore, protein-protein interaction network showed the convergence of different signaling pathways, with c-di-GMP-related genes acting as central bridges. This study highlights the interaction of different signaling pathways, especially the resilience of c-di-GMP signaling to adverse external stresses, thereby laying the foundation for facilitating electrogenic biofilm formation under adverse conditions in practical applications.
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  • 文章类型: Journal Article
    嗅觉行为在哺乳动物中具有多种社会功能。气味可能表示有关对交配和繁殖重要的个体属性的信息。嗅觉行为,如气味标记,也可以作为家庭范围或资源防御策略的一部分。我们评估了一对生活和性一夫一妻制灵长类动物嗅觉行为的潜在社会和家庭范围防御功能,Azara的猫头鹰猴(Aotusazarae),在阿根廷查科。这是对猫头鹰猴在野外嗅觉行为的最广泛调查。个体定期进行嗅觉行为(组均值+SD=每小时1.3+0.5)。这些模式通常与圈养猫头鹰猴研究中观察到的模式相当,除了洗尿是野外最常见的行为,而不是气味标记和生殖器检查。大多数嗅觉行为是由成年人执行的,并且在生殖器检查中存在明显的性别差异:几乎所有的生殖器检查都是由成年男性检查成对的成年女性组成。这些发现表明,嗅觉行为在猫头鹰猴的信号传导和协调繁殖中起着重要作用。特别是在女性受孕和怀孕期间。此外,我们的研究表明,这些行为也可以作为一种防御策略,以保持他们的家庭范围的核心区域。这项研究首次评估了嗅觉行为在生殖环境和家庭范围防御中的作用。一夫一妻制的白喉灵长类动物。
    Olfactory behaviors serve a wide variety of social functions in mammals. Odor may signal information about attributes of individuals important for mating and reproduction. Olfactory behaviors, such as scent-marking, may also function as part of home range or resource defense strategies. We assessed the potential social and home range defense functions of olfactory behavior in a pair-living and sexually monogamous primate, Azara\'s owl monkey (Aotus azarae), in the Argentinian Chaco. This is the most extensive investigation of owl monkey olfactory behaviors in the wild. Individuals regularly performed olfactory behaviors (group mean + SD = 1.3 + 0.5 per hour). The patterns were generally comparable to those observed in studies of captive owl monkeys, except that urine washing was the most common behavior in the wild, as opposed to scent-marking and genital inspections. Most olfactory behaviors were performed by adults, and there were striking sex differences in genital inspections: almost all consisted of an adult male inspecting the paired adult female. These findings suggest that olfactory behaviors play an important role in signaling and coordinating reproduction among owl monkeys, particularly during periods of female conception and pregnancy. Additionally, our research indicates that these behaviors may also serve as a defense strategy for maintaining the core area of their home ranges. This study offers the first assessment of the role of olfactory behaviors in reproductive contexts and home range defense in pair-living, monogamous platyrrhine primates.
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  • 文章类型: Journal Article
    Many animals exchange chemicals during courtship and mating. In some amphibians, sexual chemical communication is mediated by pheromones produced in male breeding glands that are transferred to the female\'s nostrils during mating. This has been mostly studied in salamanders, despite frogs having similar glands and courtship behaviours suggestive of chemical communication. In Neotropical poison frogs (Dendrobatidae and Aromobatidae), males of many species develop breeding glands in their fingers, causing certain fingers to visibly swell. Many also engage in cephalic amplexus, whereby the male\'s swollen fingers are placed in close contact with the female\'s nares during courtship. Here, we investigate the possible roles of swollen fingers in pheromone production using whole-transcriptome sequencing (RNAseq). We examined differential gene expression in the swollen versus non-swollen fingers and toes of two dendrobatid species, Leucostethus brachistriatus and Epipedobates anthonyi, both of which have specialised mucous glands in finger IV, the latter of which has cephalic amplexus. The overwhelming pattern of gene expression in both species was strong upregulation of sodefrin precursor-like factors (SPFs) in swollen fingers, a well-known pheromone system in salamanders. The differentially expressed SPF transcripts in each species were very high (>40), suggesting a high abundance of putative protein pheromones in both species. Overall, the high expression of SPFs in the swollen fingers in both species, combined with cephalic amplexus, supports the hypothesis that these traits, widespread across members of the subfamilies Colostethinae and Hyloxalinae (ca. 141 species), are involved in chemical signalling during courtship.
    Muchos animales intercambian sustancias químicas durante el cortejo y el apareamiento. En algunos anfibios, la comunicación química sexual está mediada por feromonas producidas en las glándulas reproductoras de los machos que se transfieren a las hembras durante el apareamiento. Esto se ha estudiado sobre todo en salamandras, a pesar de que las ranas tienen glándulas similares y comportamientos de cortejo que sugieren una comunicación química. En las ranas venenosas neotropicales (Dendrobatidae y Aromobatidae), los machos de muchas especies desarrollan glándulas en los dedos, lo que hace que algunos dedos se vean hinchados. Asimismo, varias especies presentan amplexo cefálico, comportamiento de cortejo en el cual los dedos hinchados entran en estrecho contacto con las narinas y boca de la hembra. En este estudio investigamos las posibles funciones de los dedos hinchados en la producción de feromonas mediante la secuenciación del transcriptoma completo (RNAseq). Examinamos la expresión génica diferencial en los dedos hinchados y no hinchados de dos especies de dendrobátidos, Leucostethus brachistriatus y Epipedobates anthonyi, ambos con glándulas mucosas especializadas en el dedo IV, y esta última especie, con amplexo cefálico. El patrón abrumador de expresión génica en ambas especies fue la alta expression de Sodefrin Precursor‐Like Factor (SPF) en los dedos hinchados, un sistema de feromonas ampliamente conocido en las salamandras. El número de transcritos SPF expresados diferencialmente en cada especie fue muy elevado (>40), lo que sugiere una gran abundancia de feromonas proteicas putativas en ambas especies. En general, la elevada expresión de SPF en los dedos hinchados en ambas especies, combinada con el amplexo cefálico, apoya la hipótesis de que estos rasgos, muy extendidos entre los miembros de las subfamilias Colostethinae e Hyloxalinae (aprox 141 especies), están implicados en la señalización química durante el cortejo.
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  • 文章类型: Journal Article
    一些丽鱼科鱼释放含有尿液的化学线索,降低对手的攻击性。识别侵略调节信息素的生物测定包括评估尿液部分对镜像或与另一男性相互作用的行为的影响。然而,许多这些方法可能是耗时的,需要许多鱼。本研究的目的是使用两种方法评估雄性莫桑比克罗非鱼(Oreochromismosambicus)对雄性尿液的行为,目的是简化生物测定:对镜像的侵略(镜像测定)和真正的对手,其中泌尿生殖器乳头使用手术丝来防止排尿。结果证实了两种实验方法中显性男性尿液的减轻侵略作用。在镜像试验中进行十分钟的咬或15分钟的拍尾行为,或与真实对手互动的5分钟的手术扩张或15分钟的横向显示是必要的,以检测对显性男性尿液的攻击行为的统计学显着减少。我们还发现,在相同条件下,具有从属地位的男性对镜子发起攻击行为的潜伏期比优势者低,即使鱼被隔离了1周。然而,在真正的对手试验中未发现潜伏期的这种差异.我们得出的结论是,在生物测定指导的信息素鉴定中,在真实对手战斗中5分钟的可操作扩张行为或在镜像试验中10分钟的咬人行为是测试尿液部分中攻击行为所需的最短时间。
    Some cichlid fishes release urine-containing chemical cues that lower aggression in their opponents. Bioassays to identify the aggression-modulating pheromone include assessing the effect of urine fractions on the behavior towards a mirror image or in interactions with another male. However, many of these methods can be time-consuming and require many fish. The objective of the present study was to assess the behavior of male Mozambique tilapia (Oreochromis mossambicus) towards male urine using two methods with the intent of simplifying the bioassays: aggression towards a mirror image (mirror assay) and real opponents in which the urogenital papilla was tied using surgical silk to prevent urination. The results confirm the aggression-reducing effect of dominant male urine in both experimental approaches. Ten minutes of biting or 15 min of tail-beating behaviors in the mirror assay, or 5 min of opercular expansion or 15 min of lateral display in interactions with real opponents were necessary to detect a statistically significant reduction in aggressive behavior towards dominant male urine. We also found that males with subordinate status had lower latency to initiate aggressive behaviors towards the mirror than dominants in the same condition, even though fish had been isolated for 1 week. However, no such differences in latency were found in the real opponent assay. We conclude that 5 min of opercular expansion behavior in real opponent fights or 10 min of biting behavior in the mirror assay are the shortest times necessary to test aggressive behavior in urine fractions in bioassay-guided identification of pheromones.
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  • 文章类型: Journal Article
    有毒河豚积聚河豚毒素(TTX),一种众所周知的神经毒素,以携带TTX的生物为食,并用它来保护自己免受捕食者的攻击。我们以前的研究表明,有毒的河豚被5,6,11-三脱氧河豚毒素(TDT)所吸引,一种无毒的TTX类似物,与TTX同时积累在有毒河豚及其猎物中。此外,使用免疫组织化学靶向神经元活性标记进行活性标记表明,TDT激活绿色斑点河豚的隐窝嗅觉感觉神经元(OSN)。然而,个体隐窝OSN是否能对TDT产生生理响应尚待确定。通过电穿孔在OSN中表达GCaMP6,我们成功地确定了一组独特的椭圆形OSN,当暴露于绿色斑点河豚的TDT时,它们表现出特定的钙反应。这些椭圆形OSN对氨基酸(AAs)没有反应,作为硬骨鱼的食物气味线索。此外,嗅觉上皮中对TDT敏感的OSN的椭圆形形态和表面定位与隐窝OSN非常相似。这些发现表明,在绿色斑点河豚中,隐窝OSN可以特异性检测到TDT。TDT气味可以充当化学引诱物,用于寻找同种有毒河豚和喂养携带TTX的生物以进行有效的中毒。
    Toxic puffers accumulate tetrodotoxin (TTX), a well-known neurotoxin, by feeding on TTX-bearing organisms and using it to defend themselves from predators. Our previous studies have demonstrated that toxic puffers are attracted to 5,6,11-trideoxytetrodotoxin (TDT), a nontoxic TTX analog that is simultaneously accumulated with TTX in toxic puffers and their prey. In addition, activity labeling using immunohistochemistry targeting neuronal activity marker suggests that TDT activates crypt olfactory sensory neurons (OSN) of the green spotted puffer. However, it remains to be determined whether individual crypt OSNs can physiologically respond to TDT. By employing electroporation to express GCaMP6s in OSNs, we successfully identified a distinct group of oval OSNs that exhibited a specific calcium response when exposed to TDT in green spotted puffers. These oval OSNs showed no response to amino acids (AAs), which serve as food odor cues for teleosts. Furthermore, oval morphology and surface positioning of TDT-sensitive OSNs in the olfactory epithelium closely resemble that of crypt OSNs. These findings further substantiate that TDT is specifically detected by crypt OSNs in green spotted puffer. The TDT odor may act as a chemoattractant for finding conspecific toxic puffers and for feeding TTX-bearing organisms for effective toxification.
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  • 文章类型: Journal Article
    嗅觉在生命的各个阶段都是健康和社会性的重要媒介,然而,它在我们的血统中受到的关注有限。嗅觉始于子宫内,参与儿童社会纽带的建立,青春期后的浪漫和性关系。气味在食品评估和避免危险方面发挥着关键作用;在现代社会中,它也指导我们的消费者行为。感觉能力通常随着年龄的增长而下降,并可能受到疾病的影响,对健康和福祉产生影响。这里,我们批判性地回顾了我们目前对人类嗅觉交流的理解,以驳斥过时的观念,即我们的嗅觉重要性不高。我们提供了嗅觉生物学的摘要,对嗅觉在整个生命过程中的重要性进行前瞻性概述,最后概述了这一领域的局限性和未来发展方向。
    The sense of smell is an important mediator of health and sociality at all stages of life, yet it has received limited attention in our lineage. Olfaction starts in utero and participates in the establishment of social bonds in children, and of romantic and sexual relationships after puberty. Smell further plays a key role in food assessment and danger avoidance; in modern societies, it also guides our consumer behavior. Sensory abilities typically decrease with age and can be impacted by diseases, with repercussions on health and well-being. Here, we critically review our current understanding of human olfactory communication to refute outdated notions that our sense of smell is of low importance. We provide a summary of the biology of olfaction, give a prospective overview of the importance of the sense of smell throughout the life course, and conclude with an outline of the limitations and future directions in this field.
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  • 文章类型: Journal Article
    表皮俱乐部细胞(ECCs)存在于许多硬骨鱼中。为了证明它们在鱼表皮中的存在,它们与许多功能相关联。一个建议的功能是在捕食事件期间与特定物种通信,因为这些细胞可能在破裂时被动释放物质,这可能发生在捕食过程中。我们确定了成人红衣主教tetra的身体皮肤中ECCs的存在和分布,Parachirodonaxelrodi(Schultz,1956年),并在实验室环境中分析了动物对同种皮肤提取物的行为反应。通过常规组织学和免疫组织化学证实动物表皮中ECCs的鉴定和分布。我们的结果表明:ECCs存在于整个身体的皮肤中;从头部到尾部的背侧观察到高密度,在鳍的插入和覆盖它们的表皮中;腹侧分布比背侧分布更广泛,更分散。用同种异体的皮肤制剂处理P.axelrodi标本导致动物的行为变化:它们表现出不稳定的游泳运动,他们表现出避免刺激应用的区域,并且减少了移动所花费的时间。总的来说,这些结果使我们得出结论,P.axelrodi在整个身体具有ECCs,在高度暴露于捕食事件的区域(背侧区域和鳍)中存在更多。暴露于同种皮肤提取物的动物在其他物种中表现出被描述为抗捕食性的行为的显着增加。这支持了这样的假设,即ECCs可能是当皮肤损伤发生时被动释放的化学警报提示的起源,提醒其他群体注意捕食的风险。
    Epidermal club cells (ECCs) are present in many species of teleost fish. In an attempt to justify their presence in the epidermis of fish, they have been associated with numerous functions. One proposed function is communication with conspecifics during a predation event, as these cells may passively release substances upon rupture, which may occur during predation. We identified the presence and distribution of ECCs in the body skin of adult cardinal tetra, Paracheirodon axelrodi (Schultz, 1956) and analyzed the animal\'s behavioral response to conspecific skin extract in a laboratory setting. The identification and distribution of ECCs in the epidermis of the animals were confirmed by conventional histology and immunohistochemistry. Our results demonstrated that: ECCs are present in the skin of the entire body; a high density is observed in the dorsal side from head to tail, in the insertion of the fins and in the epidermis covering them; and ventral distribution is less extensive and more dispersed than dorsal distribution. Treatment of P. axelrodi specimens with skin preparations of conspecifics resulted in behavioral changes in the animals: they showed erratic swimming movements, they showed avoidance of the area of stimulus application and they decreased the time spent moving. Overall, these results allow us to conclude that P. axelrodi possesses ECCs throughout the body, with a greater presence in areas of high exposure to predation events (dorsal area and fins). Animals exposed to conspecific skin extract showed a significant increase in behaviors described as anti-predatory in other species. This supports the hypothesis that ECCs may be the origin of chemical alarm cues that are passively released when skin damage occurs, alerting the rest of the group to the risk of predation.
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  • 文章类型: Journal Article
    动物社会使用巢穴识别来防止社会作弊者和寄生虫。在大多数社会昆虫社会中,个体识别和排除任何非巢穴,表皮碳氢化合物作为识别线索的作用是有据可查的。一些安布罗西亚甲虫生活在合作繁殖社会中,养殖的真菌文化很难建立,但一旦建立就具有很高的价值。因此,可以选择不支付巢穴基础费用而潜入巢穴的社会作弊者。因此,nestmate识别也有望存在于ambrosia甲虫中,但是到目前为止,还没有人调查过这种行为及其潜在的机制。在这里,我们研究了合作繁殖的ambrosia甲虫Xyleborinussaxesenii中的巢穴识别能力,结合行为观察和角质层碳氢化合物分析。萨克森氏X的实验室巢暴露于来自同一人群的外国成年雌性,另一个种群和另一个物种。观察到外国女性的生存和行为。还观察了接收个体的行为。我们预计,增加遗传距离将导致化学谱的距离增加,行为排斥水平和可能的死亡率增加。种群之间的化学特征不同,并且与其他高度社交的昆虫一样可变。然而,我们发现只有很少的证据表明外国人的行为排斥。种群间捐赠者的产巢率高于对照捐赠者,但是,无论是他们的行为还是巢内接受者的行为,都没有对任何一种治疗方法中的外国个体表现出任何反应。这些结果表明,角质层烃剖面可用于通讯和巢穴识别,但是,在X.saxesenii中不存在非巢穴的行为排除,或者激动的遭遇是如此罕见或微妙,以至于我们的方法无法检测到它们。需要更多的研究来进一步调查这一点。
    Animal societies use nestmate recognition to protect against social cheaters and parasites. In most social insect societies, individuals recognize and exclude any non-nestmates and the roles of cuticular hydrocarbons as recognition cues are well documented. Some ambrosia beetles live in cooperatively breeding societies with farmed fungus cultures that are challenging to establish, but of very high value once established. Hence, social cheaters that sneak into a nest without paying the costs of nest foundation may be selected. Therefore, nestmate recognition is also expected to exist in ambrosia beetles, but so far nobody has investigated this behavior and its underlying mechanisms. Here we studied the ability for nestmate recognition in the cooperatively breeding ambrosia beetle Xyleborinus saxesenii, combining behavioural observations and cuticular hydrocarbon analyses. Laboratory nests of X. saxesenii were exposed to foreign adult females from the same population, another population and another species. Survival as well as the behaviours of the foreign female were observed. The behaviours of the receiving individuals were also observed. We expected that increasing genetic distance would cause increasing distance in chemical profiles and increasing levels of behavioural exclusion and possibly mortality. Chemical profiles differed between populations and appeared as variable as in other highly social insects. However, we found only very little evidence for the behavioural exclusion of foreign individuals. Interpopulation donors left nests at a higher rate than control donors, but neither their behaviours nor the behaviours of receiver individuals within the nest showed any response to the foreign individual in either of the treatments. These results suggest that cuticular hydrocarbon profiles might be used for communication and nestmate recognition, but that behavioural exclusion of non-nestmates is either absent in X. saxesenii or that agonistic encounters are so rare or subtle that they could not be detected by our method. Additional studies are needed to investigate this further.
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