基于交叉抚育的雌性根田鼠对雄鼠尿气味的识别

通过交叉抚育建立同巢同胞、同巢非同胞、异巢同胞和异巢非同胞个体组成的室内繁殖种群,在断奶后（80日龄）分别取这些供体的新鲜尿作刺激物,在吕字型观察箱中观察和记录雌性根田鼠对雄鼠气味的行为响应,以研究根田鼠同胞识别的化学通讯机制。结果表明：（1）成年雌性根田鼠对雄性同巢同胞气味的接近潜伏期极显著短于对同巢非同胞气味的接近潜伏期（P〈0.01）,而其对两者的访问时间和嗅舔时间之间的差异并不显著（P〉0.05）;（2）雌鼠对雄性异巢同胞和异巢非同胞气味无明显偏好。其对两者的接近潜伏期、访问时间和嗅舔时间等行为响应均无显著差异（P〉0.05）;（3）雌鼠对雄性同巢非同胞和异巢非同胞的接近潜伏期差异并不明显（P〉0.05）,对两者访问时间和嗅舔时间的差别不大（P〉0.05）;（4）比较雌鼠对异巢同胞和同巢同胞气味的行为响应发现,其对后者的接近潜伏期显著短于前者（P〈0.05）,其对两者访问时间、嗅舔时间之间的差异未达到显著水平（P〉0.05）。这些结果表明,80日龄时,雌性根田鼠具有亲属识别能力,其同胞识别的机制可能为共生熟悉和表型匹配两种模式协同作用。     

雌性根田鼠的亲属识别与配偶选择

在Y字型选择箱中进行3种不同熟悉处理的配偶选择实验以检验动情雌性根田鼠配偶选择时的亲属识别和近交回避。（1）动情雌鼠在熟悉雄性同胞和陌生非同胞之间嗅添频次和时间、及自我修饰时间的差异非常显著（P＜0.01),所作的脊柱前凸反应时间也有显著差异（P＜0.05),这些行为均多针对于陌生非同胞雄性。结果提示动情雌性根田鼠在熟知雄性同胞身份的条件下对陌生非亲属雄性的气味更感兴趣,在择偶中与非亲属雄性交配来实现近交回避。（2）动情雌鼠对陌生雄性非同胞的嗅添频次和时间及所作的脊柱前凸反应时间显著大于陌生同胞雄性（P＜0.05),而针对陌生雄性非同胞自我修饰时间非常显著地大于针对陌生雄性同胞（P＜0.01)。这说明动情雌性根田鼠在择偶时可以识别断奶后分离不熟悉的雄性亲属,而在交配行为上偏向于非亲属雄性。（3）动情雌鼠在熟悉雄性同胞和非同胞之间跃添时间差异明显（P＜0.05),嗅添频次的差异更甚（P＜0.01);所作的脊柱前凸反应时间和社交后自我修饰时间差异也非常显著（P＜0.01),这些行为也均都偏向于非同胞雄性;结果提示断奶后继续熟悉可维持动情雌鼠对雄性亲属的嗅觉识别记忆力,在交配行为上主动回避亲属雄性。故认为：断奶后嗅觉识别持久力是发情雌性根田鼠亲属识别的机制,亲属识别是最初发情的雌性根田鼠配偶选择中近交回避的首要途径和机制,熟悉性是维持雌性根田鼠配偶选择实现近交回避的机制和特征。     

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。     

根田鼠对不同亲属关系异性尿气味的识别

根为检验根田鼠对不同亲属关系异性成体尿气味的识别能力,通过一雄两雌配对实验建立实验种群,从而产生同胞、父系半同胞和陌生个体。在行为选择箱中记录了雌、雄根田鼠对亲属系数分别为0、0.25 和0.5 异性尿气味的行为响应模式。结果如下:雌性根田鼠对3 种不同亲属关系雄鼠气味的接近潜伏期的差异达到极显著水平(P < 0.01),嗅闻时间的差异也达到显著水平(P <0.05),而嗅闻频次和反标记的差异均未达到显著水平(P >0.05)。雄性根田鼠对3 种不同亲属关系雌鼠气味的接近潜伏期和嗅闻时间的差异都未达到显著水平(P >0.05)。对不同亲属关系的气味嗅闻频次和反标记的比较分析表明,三者间的差异也未达到显著水平(P >0.05)。因此,雌性根田鼠能够识别不同亲属关系异性气味并对不同气味表现出不同的行为响应模式;而雄鼠不能识别3 种气味并对其表现出类似的行为模式。     

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。     

根田鼠气味识别的性二型

以新鲜尿和粪作气味源,在行为选择箱中观察根田鼠的行为识别模式,结果表明：雌鼠对陌生同性尿、粪气味源行为识别的差异不显,雄鼠对陌生同性尿、粪气味源的行为识别在多个指标上存在显差异;根田鼠对陌生异性尿、粪的行为响应模式无明显差异;除嗅舔时间外,雌、雄鼠对粪气味的行为识别在其他指标上不存在明显差异;比较雌、雄鼠对尿刺激的行为识别发现,雄鼠对尿刺激的访问频次和反标记显大于雌鼠,雌性嗅舔时间显大于雄性,其他方面二之间并无显差异。因此,雌、雄性根田鼠对粪气味的行为识别模式不存在性别差异;对尿气味的行为识别模式存在性别差异。     

熟悉性对棕色田鼠和根田鼠择偶行为的影响

以两性配对共居至交配或生仔作不同熟悉处理,在Y字型选择箱中分别观察棕色田鼠,根田鼠的择偶行为和配偶识别。结果如下：（1）棕色田鼠雌雄两性均表现出对熟悉异性或配偶的喜好倾向;（2）在根 中,只有雌性表现出对熟悉雄鼠或配偶的选择倾向,雄鼠无明显的选择取向;（3）每种雌性田鼠对2种熟悉性不同的同种雄性的嗅舔时间均有显著差异性,而每种雄性田鼠对2种熟悉性不同的同种雌性的嗅舔时间均无显著差异性。这些结果表明：（1）熟悉性对棕色田鼠择偶行为的影响大于对根田鼠的影响;（2）嗅舔时间差别可反映不同田鼠的配偶识别能力,熟悉性对棕色田鼠配偶识别能力的影响大于对根田鼠的影响;（3）2种田鼠配偶识别的性别差异的一致性,提示这2种田鼠的雌性配偶识别能力均强于雄性田鼠,而雌鼠的配偶识别能力可能取决于雌鼠受酱或分娩刺激的生理状态;（4）棕色田鼠室内所表现的择偶行为的两性熟悉性特征与其单配制有关,根田鼠所表现的择偶行为的雌性熟悉性特征与其一雄多雌制有关。     

雌性根田鼠断奶后对亲本尿气味的记忆

为研究亲子分开后雌性根田鼠对亲本气味的记忆持续时间，分别在未分开（20日龄）、分开10 d（30日龄）、20 d（40日龄）、30 d（50日龄）、40 d（60日龄）时，以新鲜尿作气味源，在行为观察箱中记录雌性根田鼠对不同气味源的行为响应模式，结果表明：(1) 未分开时，雌鼠遭遇父本气味时自我修饰的频次极显著高于陌生雄鼠气味，在分开10 d时，雌鼠接近父本气味的频次显著多于接近陌生雄鼠气味的频次，其对前者反标记显著少于后者；(2)分开20 d后，雌鼠对父本和陌生雄鼠气味的行为响应无明显差异；(3) 未分窝时，雌性根田鼠幼仔对母本和陌生雌鼠气味的行为响应无差异；（4）分开10—40 d时，雌性根田鼠对母本和陌生雌鼠气味表现出不同的行为响应模式。以上结果表明，在亲子分开10 d时，雌鼠仍能识别父本和陌生雄鼠的气味；分开20 d后，雌鼠不再能够识别父本和陌生雄鼠的气味；在亲子分开40 d时，雌鼠仍能识别母本和陌生雌鼠的气味。因此，雌鼠对父本气味的嗅觉记忆时间可以持续到亲子分开10—20 d之间；而对母本气味的嗅觉记忆时间则可以持续到亲子分开40 d以上。     

雄性柴达木根田鼠断奶后对亲本尿气味的记忆

为了研究亲子分开后雄性柴达木根田鼠(Microtus limnophylus)对亲本尿气味的记忆持续时间,分别在未分开(20日龄),以及分开10d(30日龄)、20d(40日龄)、30d(50日龄)、40d(60日龄)时,以新鲜尿作为气味源,在行为观察箱中记录雄性柴达木根田鼠对不同气味源的行为响应模式.结果表明:(1)未分开时,雄鼠对父本气味的接近频次显著高于陌生雄鼠气味;分开10d时,雄鼠对父本气味的访问时间显著多于对陌生雄鼠气味的访问时间;分开20d时,雄鼠对陌生雄鼠气味的接近潜伏期极显著短于父本,对陌生雄鼠气味的访问时间极显著长于父本,其遭遇父鼠气味时的自我修饰频次显著少于陌生雄鼠气味.(2)分开30d后,雄鼠对父本和陌生雄鼠气味的行为响应没有明显差异.以上结果表明,在亲子分开20d时,雄鼠仍能识别父本与陌生雄鼠的气味;在分开30d后,雄鼠不再能够识别父本与陌生雄鼠的气味.(3)在未分窝时,雄性柴达木根田鼠幼仔对母本和陌生雌鼠气味的行为响应没有任何差异.(4)在分开10d时,雄性柴达木根田鼠对母本和陌生雌鼠气味表现出不同的行为响应模式;分开20～40d时,雄鼠对母本与陌生雌鼠气味的行为响应没有任何差异.以上结果表明,在亲子分开10d时,雄鼠仍能识别母本与陌生雌鼠的气味;在分开20d后,雄鼠不再能够识别母本与陌生雌鼠的气味.因此,雄鼠对父本气味的嗅觉记忆时间可以持续到亲子分开20～30d之间;其对母本气味的嗅觉记忆时间可以持续到亲子分开10d时.     

熟悉性对棕色田鼠和根田鼠择偶行为的影响

以两性配对共居至交配或生仔作不同熟悉处理,在Y字型选择箱中分别观察棕色田鼠、根田鼠的择偶行为和配偶识别.结果如下:(1)棕色田鼠雌雄两性均表现出对熟悉异性或配偶的喜好倾向;(2)在根田鼠中,只有雌性表现出对熟悉雄鼠或配偶的选择倾向,雄鼠无明确的选择取向;(3)每种雌性田鼠对2种熟悉性不同的同种雄性的嗅舔时间均有显著差异性,而每种雄性田鼠对2种熟悉性不同的同种雌性的嗅舔时间均无显著差异性.这些结果表明:(1)熟悉性对棕色田鼠择偶行为的影响大于对根田鼠的影响;(2)嗅舔时间差异可反映不同田鼠的配偶识别能力,熟悉性对棕色田鼠配偶识别能力的影响大于对根田鼠的影响;(3)2种田鼠配偶识别的性别差异的一致性,提示这2种田鼠的雌性配偶识别能力均强于雄性田鼠,而雌鼠的配偶识别能力可能取决于雌鼠受交配或分娩刺激的生理状态; (4)棕色田鼠室内所表现的择偶行为的两性熟悉性特征与其单配制有关,根田鼠所表现的择偶行为的雌性熟悉性特征与其一雄多雌制有关.     

The ontogeny of kin recognition in tadpoles of the toadBufo melanostictus (Anura; Bufonidae)

The ontogeny of kin recognition and influence of social environment on the development of kin recognition behaviour was experimentally investigated in tadpoles of Bufo melanostictus that lived in aggregations and showed low larval dispersion. Embryos and tadpoles of the toad were reared as (i) kin only, (ii) with kin and non-kin (separated by a mesh screen), and (iii) in isolation. They were tested for the ability to discriminate between (i) familiar siblings and unfamiliar non-siblings, (ii) familiar siblings and familiar non-siblings and, (iii) unfamiliar siblings and unfamiliar non-siblings. All tadpoles were fed on boiled spinach before conducting trials. Preference of test tadpoles to associate near the end compartments whether empty or containing members of specific stimulus groups was assessed using a rectangular choice tank. When tested in tanks with empty end compartments, the test tadpoles showed random distribution and thus no bias for the apparatus or the procedure. In the presence of kin/non-kin in the end compartments a significantly greater number of test tadpoles spent the majority of the time near familiar or unfamiliar kin rather than near familiar or unfamiliar non-kin. Kin discrimination ability persisted throughout larval development. Familiarity with siblings is not required for discriminating kin from non-kin, and kin discrimination ability is not modified following exposure to non-kin. Also, involvement of dietary cues is unlikely to be the prime mechanism of kin recognition inB. melanostictus unlike in some other anurans.     

Kin recognition by larval wood frogs (Rana sylvatica): effects of diet and prior exposure to conspecifics

Summary We investigated kin recognition by larval wood frogs (Rana sylvatica) in blind laboratory experiments using spatial affinity as a recognition assay. Tadpoles reared with full-sibs displayed a significant preference for familiar full-sibs over unfamiliar non-kin, but failed to discriminate between unfamiliar full-sibs and unfamiliar paternal half-sibs. Tadpoles reared in social isolation (with or without maternal egg jelly) from the two-celled embryonic stage displayed a significant preference for unfamiliar full-sibs over unfamiliar non-kin. Tadpoles reared on a meat diet with their full-sibs: 1) exhibited a significant preference for unfamiliar full-sibs fed meat over unfamiliar non-kin fed meat, 2) failed to discriminate between unfamiliar full-sibs fed lettuce and unfamiliar non-kin fed meat, 3) exhibited a significant preference for unfamiliar non-kin fed meat over unfamiliar non-lin fed lettuce, 4) failed to discriminate between unfamiliar full-sibs fed meat and unfamiliar full-sibs fed lettuce, and 5) displayed a significant spatial preference for odors associated with meat (a familiar food) over odors associated with lettuce (an unfamiliar food). Our results, together with those of Cornell et al. (1989), indicate that the recognition cue of larval R. sylvatica has both genetic and environmental (dietary) components. Our findings establish that previous exposure to maternal egg jelly, kin, or conspecifics is not necessary for the development of kin recognition ability in larval R. sylvatica. Our results are more consistent with the self-learning of recognition cues (a form of phenotype matching) than with a recognition mechanism that involves a genetically fixed recognition template. Finally, our results indicate that increasing similarity between the recognition template and perceived cue does not necessarily result in increasing spatial affinity for kin.     

Odor-based recognition of familiar and related conspecifics: a first test conducted on captive Humboldt penguins (Spheniscus humboldti)

Studies of kin recognition in birds have largely focused on parent-offspring recognition using auditory or visual discrimination. Recent studies indicate that birds use odors during social and familial interactions and possibly for mate choice, suggesting olfactory cues may mediate kin recognition as well. Here, we show that Humboldt penguins (Spheniscus humboldti), a natally philopatric species with lifetime monogamy, discriminate between familiar and unfamiliar non-kin odors (using prior association) and between unfamiliar kin and non-kin odors (using phenotype matching). Penguins preferred familiar non-kin odors, which may be associated with the recognition of nest mates and colony mates and with locating burrows at night after foraging. In tests of kin recognition, penguins preferred unfamiliar non-kin odors. Penguins may have perceived non-kin odors as novel because they did not match the birds' recognition templates. Phenotype matching is likely the primary mechanism for kin recognition within the colony to avoid inbreeding. To our knowledge this is the first study to provide evidence of odor-based kin discrimination in a bird.     

Kin recognition in Bufo scaber tadpoles: ontogenetic changes and mechanism

Ontogenetic changes in kin-recognition behavior, effect of social environment on kin-recognition ability, and use of visual and chemical cues in kin recognition have been studied in tadpoles of Bufo scaber after rearing them with kin, in mixed groups, or in isolation from Gosner stage 12 (gastrula). By use of a rectangular choice tank the tadpoles were tested for their ability to choose between (a) familiar siblings and unfamiliar non-siblings, (b) unfamiliar siblings and familiar non-siblings, and (c) unfamiliar siblings and unfamiliar non-siblings. When tested without any stimulus groups in the end compartments of the tank, random distribution was observed for the tadpoles and no bias for the apparatus or the procedure. In the presence of kin and non-kin in the end compartments, significantly more tadpoles spent most of their time near kin (familiar or unfamiliar) rather than near non-kin during early larval stages, up to stage 37. After stage 37 (characterized by the differentiation of toes), test tadpoles showed no preference to associate with kin, suggesting an ontogenetic shift in the kin-recognition ability in B. scaber. In experiments involving selective blockade of visual or chemical cues the test tadpoles preferentially associated near their kin on the basis of chemical rather than visual cues. These findings suggest that familiarity with siblings is not necessary for kin recognition and that kin-recognition ability is not modified after exposure to non-kin by mixed rearing. The findings for B. scaber indicate a self referent phenotype matching mechanism of kin recognition which is predominantly aided by chemical rather than visual cues.     

Kin recognition in golden hamsters: evidence for kinship odours

Differential treatment of kin and non-kin has been well documented, but much remains unclear about how kin are recognized. If kin are recognized by a phenotype-matching mechanism, there must be a correlation between genetic relatedness and the similarity of cues used for recognition. A habituation technique was used with golden hamsters,Mesocricetus auratus, to investigate the relative similarity of the odour quality of flank gland secretions from siblings and unrelated individuals. Hamsters discriminated between the odours of their own, same-sex siblings but also treated these odours as similar compared to odours of non-siblings (experiment 1). They did not discriminate between the flank gland odours of unfamiliar siblings from another family (experiment 2). They also did not discriminate between the flank gland odours of unfamiliar, paternal half-siblings from another family (experiment 3). These results indicate that subjects perceived odours from genetically similar individuals as similar and provide evidence for kinship odour cues. The discrimination between the flank gland odours of subjects’ own siblings, however, indicates that hamsters learn the subtle differences between the odours of their close kin, probably through experience with siblings in the nest. When only volatile components from flank gland secretions were available to subjects (experiment 4), they again discriminated between the odours of their own siblings, suggesting that the volatile components from the flank gland secretion were sufficient for recognition of individual litter-mates.     

Kin recognition by tadpoles and froglets of the wood frog Rana sylvatica

Summary We investigated kin recognition by the wood frog Rana sylvatica in blind laboratory experiments using spatial proximity as a recognition assay. Tadpoles were tested for the ability to discriminate between: 1) familiar full-sibs and unfamiliar non-kin, 2) unfamiliar paternal half-sibs and unfamiliar non-kin, and 3) familiar and unfamiliar full-sibs. Tadpoles discriminated full- and paternal half-sibs from unrelated conspecifics, but did not discriminate between familiar and unfamiliar full-sibs. Froglets from the same laboratory population were tested for the ability to discriminate between 1) familiar full-sibs and unfamiliar non-kin, and 2) unfamiliar paternal half-sibs and unfamiliar non-kin. Froglets preferentially associated with full- and half-sibs over unrelated conspecifics. Our results show that familiarity, i.e., prior association, is not necessary for kin recognition in tadpoles and froglets. The ability of tadpoles and froglets to recognize unfamiliar paternal half-sibs demonstrates that a common maternal factor is not necessary for kin recognition, and indicates that the recognition cue has a genetic component. Our results add to the increasing evidence that a variety of vertebrate and invertebrate animals have the ability to recognize unfamiliar kin by using genetically specified recognition cues.     

Kin recognition in Rana cascadae tadpoles: maternal and paternal effects

Cascades frog (Rana cascadae) tadpoles preferentially associate with full siblings over half sibling and half siblings over non-siblings when reared with siblings or as isolates. These tadpoles can use cues of maternal or paternal origin in distinguishing siblings from non-siblings, but maternal cues are preferred over paternal cues. This suggests that a hierarchy of cue importance may exist. Our results are consistent with both a phenotypic matching and a genetic recognition system of kin recognition. Thus, both learned and innate components may play a role in R. cascadae sibling recognition. Kin recognition may facilitate preferential treatment of kin, such as cooperation in food finding or in warning against predators, and therefore those individuals behaving altruistically in kin groups can increase their inclusive fitness.     

Ovarian hormones influence odor cues emitted by female meadow voles, Microtus pennsylvanicus.

During the spring-summer breeding season female meadow voles emit odors that are preferred by males, whereas in the autumn-winter season of reproductive quiescence females emit odors that are not preferred by males, but are attractive to females. The effects of daylength and ovarian hormones on salience of female odors were determined by assaying male responses to odors. Females housed in long and short photoperiods transmitted odors that elicited responses similar to those of spring and autumn female voles, respectively. The odor cues emitted by ovariectomized (OVX) females, irrespective of photoperiodic history, were similar to those generated by females during the nonbreeding season. In the absence of ovarian hormones, long daylengths were not sufficient to induce females to broadcast the spring odors preferred by males. Spring-type odor cues were, however, emitted by OVX voles housed in either photoperiod and treated with estradiol. Ovarian hormones appear necessary and sufficient to generate breeding season odor cues and sufficient to induce production of such cues during the nonbreeding season. We conclude that daylength affects odor cues emitted by females by altering ovarian hormone activity.