雄性等级对布氏田鼠雌性选择的影响

利用监控录像以及特定行为记录的方法,研究了雄性等级对布氏田鼠气味选择和个体选择的影响。结果表明不论是气味选择还是个体选择,雌鼠动情状态不影响对雄鼠的选择频次,但是动情雌鼠在自我修饰行为频次上高于非动情雌鼠。此外,在气味和个体选择的优先权方面,动情雌鼠更多的优先选择优势雄鼠的气味和个体并与之交配,而非动情雌鼠更多的先选择从属雄鼠的气味和个体。总之,不论是气味选择还是个体选择,布氏田鼠不仅受到被选择雄鼠的地位影响,还与雌鼠的生理状态有关。     

青春期雌性根田鼠初次择偶行为与雄性优势等级

在实验室条件下，研究了性刚成熟的雌性根田鼠对一对互为同胎同胞的以体重划分为优势和从势的雄性个体的择偶行为，目的在于检验雄性社会等级是否影响其配偶选择。结果发现：(1)根田鼠存在显著的偏雄性二型，而雄性动物优势度与体重大小成正比；(2)在Y字型迷宫中，初次发情雌鼠对优势雄鼠的访问时间、社会探究频次和友好频次等显著大于从属雄鼠；(3)在随后24h的一雌二雄共居中，优势雄优先交配；从属雄的干扰不能中断优势雄鼠的交配。这些结果说明，体重差异可以作为划分雄性优势的指标，社会等级可以作为择偶行为的一个依据。优势雄鼠与处于从属地位的同胞相比，能获得优先交配机会。同胎同胞虽具有遗传相似性，但雄性竞争的结果表现在体重和社会等级上具有显著差异，表明配偶选择在发挥着作用。根田鼠是一雄多雌制，而且具有较大的偏雄性二型，提示社会等级、雌性选择和婚配制度有密切关联[动物学报49(3)：303—309，2003]。     

不同社群条件下雄性布氏田鼠的行为

实验室内观察了布氏田鼠雄性个体之间在3 种不同社群条件下(社群1:2 只雄鼠;社群2:2 只雄鼠与1只非动情雌鼠,社群3:2 只雄鼠与1 只动情雌鼠)的相互行为,结果显示:(1)在嗅闻行为上,不论优势雄鼠还是从属雄鼠在3 组间都存在显著差异,它们的嗅闻频次都是在社群3 中最少、在社群2中居中,在社群1中嗅闻频次最多;(2)在攻击行为频次上,不论是优势雄鼠还是从属雄鼠在3 组中都不存在显著差异;(3)在上跳频次上,优势雄鼠在各组间存在显著差异,从属雄鼠在各组间不存在显著差异; (4) 在自我修饰行为频次上,不论是优势雄鼠还是从属雄鼠,在3 组之间不存在显著差异;(5)在相互友好行为上,3 组间存在显著差异。其中社群2 中的雄性之间的友好行为频次最高,社群3 中次之,社群1 中最低。结果表明,雄性之间的攻击行为并不因为雌性的存在而增强,反而会可能减少,我们推测这可能因为雄性要花费更多的时间用于社群探究和交配,从而减少了雄性之间的斗争。     

雄性根田鼠的同胞竞争及其对同性个体的气味识别

采用两两互作实验并以尿标记来确定共居至成年的2雄性根田鼠同胞在行为竞争上的差异,之后在行为选择箱中分别观察它们对来自熟悉和陌生同性个体的新鲜尿气味的行为反应,以确定气味识别差异。结果表明：①雄性根田鼠两两互作时,尿标记的多少可以作为判断其社会等级的标准;②从属鼠对熟悉气味存在明显偏好,其对熟悉气味的访问时间、接近频次和自我修饰频次都显著高于陌生气味;③优势鼠则优先访问陌生气味,其对陌生气味的访问时间、嗅舔频次、嗅舔时间、自我修饰及反标记均显著高于熟悉气味;④对于陌生气味,优势鼠和从属鼠之间存在明显不同的行为反应模式。本实验结果提示,雄性根田鼠对同性尿气味识别的差异以及对陌生气味的行为反应,可能暗含着其领域防卫的信息。     

不育雄性对布氏田鼠气味选择和个体选择的影响

在Y型迷宫中进行了动情雌性布氏田鼠对结扎不育雄鼠及正常雄鼠气味和个体的选择实验。气味选择实验以动物的巢垫物为气味源,个体选择直接采用结扎雄性和正常雄性作为被选择个体。记录雌性布氏田鼠对选择源的选择频次以及停留时间。结果表明,无论在气味选择或是个体选择中,动情雌性布氏田鼠在选择频次和停留时间上都具有显的正相关性,而且都没有明显表现出对某—气味或个体选择上的偏好。激素测定结扎不育后雄鼠血清中的睾酮和皮质醇含量有听升高．但这种变化没有对雌性布氏田鼠的选择产生影响。     

非繁殖期成年雄性布氏田鼠对群体气体的辨别

本文研究了非繁殖期成年雄性布氏田鼠对陌生群本气味,本群体气体的社会探究行为和尿标记行为,探讨了布氏田鼠利用气味信号进行群体辨别的能力。在熟悉相的辨别实验中,雄鼠对陌生群体气味的社会探究和尿标记行为明显我于本群体气味和对照组,优势个体的探究和尿尿行为显著高于从属个体。实验结果表明,布氏田鼠具体对群体气味的辨别能力,在巢垫物中存在用于群体识别的化学通讯成分,优势鼠的气味标记可能性种内群体间的嗅觉通讯中     

雌性布氏田鼠对雄鼠气味的辨别

通过雌性布氏田导其配偶和陌生雄鼠气味的辨别实验,以及无交配经历的成年雌鼠对社会等级不同的陌生雄鼠气的辨别实验发现,有生育经历的雌鼠对陌生雄鼠气味的探究行为明显多于对配偶气味的控究,但雌鼠在配偶雄鼠气味周围的停留时间、自身修饰行为和何处行为的发生频次和持续地陌生雄鼠,没有交配经历的雌鼠对处于从鼠地位的雄性个体气味探究行为多于对优势雄性的控究,而其他行为表现没有明显差异。表明,雌性布氏田鼠可以利用气味     

非繁殖期成年雄性布氏四鼠对群体气味的辨别

本文研究了非繁殖期成年雄性布氏四鼠对陌生群体气味、本群体气味的社会探究行为和尿标记行为,探讨了布氏四鼠利用气味信号进行群体辨别的能力。在对熟悉相的辨别实验中,雄鼠对陌生群体气昧的社会探究和尿标记行为明显多于本群体气味和对照组,优势个体的探究和尿标记行为显著高于从属个体。实验结果表明：布氏田鼠具有对群体气味的辨别能力;在巢垫物中存在用于群体识别的化学通讯成分,优势鼠的气味标记可能在种内群体间的嗅觉通讯中起着重要的作用。     

不同社会等级雄性根田鼠对自身和同性陌生气味的辨别

采用两两遭遇法确定雄性根田鼠的社会等级,然后以新鲜尿作气味源,在行为选择箱中观察不同社会等级雄性根田鼠对自身气味和非自身气味的行为响应模式,结果表明:种群中从属个体对自身尿液存在明显偏好,其对自身尿液的接近潜伏期显著短于非自身尿液,而访问时间、自我修饰频次都显著高于非自身尿液;种群中优势个体则优先访问非自身尿液,其对非自身尿液的访问时间、嗅舔时间、自我修饰及反标记均显著高于自身尿液;对非自身尿液,不同社会等级雄性个体之间存在明显不同的行为反应模式。这说明,不同社会等级雄性根田鼠具有自我识别的能力且模式不同,同时其对非自身尿气味响应模式的差异也与自身的社会等级有关。     

雄性布氏田鼠对不同熟悉程度和动情状态下雌鼠气味的辨别

通过雄性布氏田鼠（Microtus brandti)对配偶和陌生雌鼠气味的辨别实验发现：（1）雌鼠动情状况对雄鼠气味行为反应的影响,当配偶雌鼠处于动情期时,被试雄鼠对动情陌生雌鼠巢垫物的溴闻和舔舐的持续时间都明显多于对非动情陌生雌鼠巢垫物的嗅闻和舔舐时间,而其他行为没有明显判别当配偶雌鼠处于非动情期时,雄鼠对动情的和非动情的陌生雌鼠的气味行为反应都没有明显判别当陌生雌鼠处于动情期时,被试雄鼠对动情配偶气味的嗅闻时间明显多于对非动情配偶的嗅闻时间,而其他行为反应没有明显差异;当陌生雌鼠处于非动情期时,被试雄鼠对动情配偶雌鼠巢垫物的嗅闻时间,挖掘频次,舔舐频次和时间都明显多于对非动情配偶气味的嗅闻,而且雄鼠在动情配偶气味周围搔扒体侧行为的发生频次和持续时间也多于在非动情配偶气味周围的搔扒行为;（2）熟悉性对雄鼠气味行为反应的影响,当配偶雌鼠和陌生雌鼠都处于动情期或都处于非动情期时,被试雄鼠对陌生雌鼠气味的探究行为（嗅问、挖掘和舔噬）和搔扒体侧行为明显多于对配偶雌鼠气味的探究;雄鼠访问陌生雌鼠气味源箱的频次和在其中的停留时间也都多于访问配偶雌鼠气味源箱的频次和停留时间;当配偶雌鼠处于动情期而陌生雌鼠处于非动情期时,被试雄鼠对配偶气味的嗅闻,挖掘和自身修饰行为明显多于对陌生雌鼠气味的行为反应,进入配偶气味源箱中的访问频次也多于进入陌生雌鼠气味源箱的访问频次。结果说明：雌鼠与被试雄鼠的熟悉程度和雌鼠的动情状况直接影响雄性布氏田鼠的社会探究和气味选择行为,即雄鼠偏好并选择动情雌鼠和陌生雌鼠,雄性布氏田鼠对雌鼠气味的行为反应也表现出田鼠属动物典型的多配制种类特征。     

Serum testosterone, male dominance, and aggression in captive groups of vervet monkeys (Cercopithecus aethiops sabaeus)

The relationship of serum testosterone concentration to male dominance rank and frequency of aggression was investigated in stable vervet monkey social groups, each containing two or three adult males, several adult females, and their offspring. Dominance relationships were determined by noting an animal's success in intermale aggressive encounters. A striking finding was the marked within-subject variation in testosterone concentration: 5- to 10-fold fluctuations were often observed on successive days. When all 15 groups were considered together, testosterone concentration was unrelated to dominance rank. Although mean testosterone concentration for all dominant males was higher than the mean for all subordinate males, this difference was not significant. In a subset of 4 groups, the rate of aggression initiated was significantly correlated with same-day testosterone in dominant but not in subordinate males.     

Ritualized submission and the reduction of aggression in an invertebrate

Ritualized behaviors that signify acceptance of a dominance relationship and reduce aggression between rivals are a common feature of vertebrate social behavior. Although some invertebrates, including crayfish, lobsters, and ants, display dominance postures, more complex dominance rituals and their effects on fitness have not been reported. We found that crayfish display such a complex ritual, when two males engaged in pseudocopulatory behavior to signify their dominance relationship. This was followed by a reduction in aggression and an increased likelihood of the subordinate's survival. Pseudocopulation was initiated by the eventual dominant and could be accepted or refused by the eventual subordinate. The frequency of aggressive behavior declined significantly during the first hour in all pairs that pseudocopulated but remained high in pairs that did not. Whereas all the subordinate members of pairs that pseudocopulated survived the initial 24 hr of pairing, half of subordinates that did not pseudocopulate were killed during that time. This differential mortality indicates that the reduction of aggression induced by the pseudocopulatory ritual directly enhances the differential survival of male crayfish that engage in this behavior.     

Beyond aggression: Androgen-receptor blockade modulates social interaction in wild meerkats

In male vertebrates, androgens are inextricably linked to reproduction, social dominance, and aggression, often at the cost of paternal investment or prosociality. Testosterone is invoked to explain rank-related reproductive differences, but its role within a status class, particularly among subordinates, is underappreciated. Recent evidence, especially for monogamous and cooperatively breeding species, suggests broader androgenic mediation of adult social interaction. We explored the actions of androgens in subordinate, male members of a cooperatively breeding species, the meerkat (Suricata suricatta). Although male meerkats show no rank-related testosterone differences, subordinate helpers rarely reproduce. We blocked androgen receptors, in the field, by treating subordinate males with the antiandrogen, flutamide. We monitored androgen concentrations (via baseline serum and time-sequential fecal sampling) and recorded behavior within their groups (via focal observation). Relative to controls, flutamide-treated animals initiated less and received more high-intensity aggression (biting, threatening, feeding competition), engaged in more prosocial behavior (social sniffing, grooming, huddling), and less frequently initiated play or assumed a ‘dominant’ role during play, revealing significant androgenic effects across a broad range of social behavior. By contrast, guarding or vigilance and measures of olfactory and vocal communication in subordinate males appeared unaffected by flutamide treatment. Thus, androgens in male meerkat helpers are aligned with the traditional trade-off between promoting reproductive and aggressive behavior at a cost to affiliation. Our findings, based on rare endocrine manipulation in wild mammals, show a more pervasive role for androgens in adult social behavior than is often recognized, with possible relevance for understanding tradeoffs in cooperative systems.     

Influence of reproductive season and rank on fecal glucocorticoid levels in free-ranging male Verreaux's sifakas (Propithecus verreauxi)

Studies in anthropoid primates and other mammals suggest that reproductive season, rank, reproductive skew, aggression received, and social support are the major factors influencing glucocorticoid output. In which way these are also affecting adrenal function in lemurid primates has been studied rarely. Here, we examine the influence of reproductive season and rank on glucocorticoid output in male sifakas (Propithecus verreauxi), a species characterized by high breeding seasonality, a hierarchy among males and extreme reproductive skew towards dominant males. We established a fecal assay for non-invasively monitoring adrenal activity and collected 315 fecal samples during the reproductive and birth season from 10 male sifakas living in 5 groups in Western Madagascar. We found a significant effect of season on glucocorticoid output, with males exhibiting higher fecal glucocorticoid levels during the reproductive compared to the birth season in conjunction with an increase in overall aggression rates during the former period. Moreover, our data indicate a significant effect of rank on adrenocortical activity with dominant males exhibiting higher glucocorticoid levels than subordinate males in the reproductive season. However, dominant males did not differ significantly in rates of initiated or received aggression and rates of affiliative behavior from subordinates but showed significantly lower rates of submission. Given their highly formalized dominance relationships, we conclude that higher glucocorticoid output in dominant males during the 4-month reproductive season is likely related to higher energetic demands necessary to cope with the challenges of male reproduction rather than to physical demands of increased fighting frequency to maintain dominance status. High rank in sifakas may thus carry high costs, which, however, may be outweighed by monopolization of almost all paternities. In sum, our data generally support the findings on the relationship between environmental and social factors and glucocorticoid output found in non-lemurid primates.     

Serum testosterone, agonistic behavior, and dominance in inbred strains of mice

Social situations in which male mice establish dominant/subordinate relationships were utilized in an attempt to correlate circulating testosterone (T) titer with agonistic behavior. Two long-term (several months) and two short-term (3- and 5-day) situations in which dominance was verified by severity of body scarring or individual aggression scores indicated no consistent correlation of dominance with serum T levels.     

Effects of Dominant/Subordinate Social Status on Formalin-Induced Pain and Changes in Serum Proinflammatory Cytokine Concentrations in Mice

Current investigations regarding social stress primarily focus on the health consequences of being in stressful social hierarchies. The repetitive nature of social conflicts seems to favor an induction of hyperalgesia or hypoalgesia, both in rodents and humans. Additionally, social conflicts may affect the immune system. In order to better establish the pain and immune responses to stress, the present study implemented a sensory contact model on 32 male BALB/c mice. Subsequent to establishing a dominance/submissive social relationship, each mouse was injected with formalin (20 μl, 2%) and their pain behavior was scored and serum concentrations of proinflammatory cytokines IL-1 and IL-6, and corticosterone were also measured. Test results revealed that subordinate mice were hypoalgesic during chronic phase of formalin test compared to control and dominant mice (P<0.05). On the other hand, subordinate mice were hyperalgesic compared to dominant mice during the whole acute phase of formalin test (P<0.05). Corticosterone, IL-1 and IL-6 concentrations were much higher in serum of dominant and subordinate mice than in the control group (p<0.05). The results indicated that, although both dominant and subordinate animals displayed an increase in serum corticosterone and proinflammatory cytokines during social interactions, their response to pain perception differently was affected with the social status.     

The effects of temporary removal of the alpha male on the behavior of subordinate male vervet monkeys

The effects of temporary removal of the alpha male on the behavior of subordinate adult male vervet monkeys were evaluated. Twelve subordinate males from six captive multimale, multifemale vervet monkey groups were observed in two conditions: when all group members were present and when the alpha male was temporarily removed from each group. In the absence of the alpha male, subordinate males initiated more affiliative behavior and increased the amount of time spent in proximity to females but their rates of aggression were unaltered. Increased affiliative behavior was selectively directed to high-ranking females and their offspring. Under removal conditions, subordinate male behavior did not resemble that of alpha males in intact conditions: they differed in their proximity to and affiliative behavior towards other group members. In the absence of the alpha male, females increased their aggression towards subordinate males. These observations suggest that the presence of alpha males strongly inhibits subordinate males' behavior. When the constraints of the alpha male's presence are removed, subordinate males rapidly engage in behavior that may enhance their likelihood of attaining high rank. In combination with prior studies, the data also indicate that the behaviors involved in the maintenance of high rank by alpha males differ from those subordinates use to acquire dominance. Finally the current study supports the view that aggression by female vervets may be highly influential in determining male ascendency to dominant rank.     

Social status,immune response and parasitism in males: a meta-analysis

In male vertebrates, two conflicting paradigms—the energetic costs of high dominance rank and the chronic stress of low rank—have been proposed to explain patterns of immune function and parasitism. To date, neither paradigm has provided a complete explanation for status-related differences in male health. Here, we applied meta-analyses to test for correlations between male social status, immune responses and parasitism. We used an ecoimmunological framework, which proposes that males should re-allocate investment in different immune components depending on the costs of dominance or subordination. Spanning 297 analyses, from 77 studies on several vertebrate taxa, we found that most immune responses were similar between subordinate and dominant males, and neither dominant nor subordinate males consistently invested in predictable immune components. However, subordinate males displayed significantly lower delayed-type hypersensitivity and higher levels of some inflammatory cytokines than dominant males, while dominant males exhibited relatively lower immunoglobulin responses than subordinate males. Despite few differences in immunity, dominant males exhibited consistently higher parasitism than subordinate males, including protozoan blood parasites, ectoparasites and gastrointestinal helminths. We discuss our results in the context of the costs of dominance and subordination and advocate future work that measures both parasitism and immune responses in wild systems.