动物行为学知识简介(二)——性、双亲行为和婚配制度

1.性冲突和性进化动物生殖细胞的大小有很大区别。雄性的生殖细胞——精子小而多,雌性的生殖细胞——卵子大且营养丰富,数量比精子少得多。所以在生殖过程的初始,两性的生殖投资即相差甚远,然而收获却相等——下一代身上雄雌两性的基因含量永远是相等的。雌雄动物的繁殖策略(最大可能地繁殖下一代)也相应地不同:雄性动物是通过与更多的雌性交配;而雌性动物则通过更快地产卵或幼仔。出于交配的需要引起的同性(往往是雄性)     

多倍体银鲫克隆多样性和双重生殖方式的遗传基础和育种应用

单性物种一般是多倍体,且通过单性生殖方式如雌核生殖、杂种生殖或孤雌生殖繁殖.与其他单性和多倍体物种相比,银鲫具有更高的倍性,为六倍体.它经历了几轮连续的基因组多倍化,还经历了一次额外的、在较近年代发生的基因组复制事件.更为重要的是,银鲫已被证实同时存在雌核生殖和有性生殖双重生殖方式.本文综述了银鲫的多倍化起源、克隆多样性和双重生殖方式的遗传基础,概述了其新品种培育和有关其生殖与早期发育相关基因鉴定的研究进展.已有实验证据表明,银鲫正处于二倍化的进化轨道中.作为一个新的进化发育(Evo-Devo)生物学模型,重点论述了银鲫在重复基因的功能歧化和单性动物的有性起源和进化等方面的研究前景.     

动物生殖抑制研究进展：理论模型、方式和机制

生殖抑制指原本具有生育能力的动物个体因特定外界环境或生理条件而减少或丧失生殖能力的现象,有时是受环境变化的主动调控,更多的是出现在其他个体影响下的被动抑制,极端情况发生在社会性动物的永久性抑制,即永久无法生殖或无法生殖成熟。研究发现非社会性动物也有生殖的推迟及可恢复性的生殖抑制,生殖抑制影响着动物种群数量动态、维持和进化。随着多学科的发展,生殖抑制机理研究取得了诸多进展。从阐述生殖抑制的概念出发,解析生殖抑制的形态、激素和分子生理特征,总结了生殖抑制的原因、作用,终述现有的理论模型以及不同物种方面的最新进展,并就生殖抑制在生物资源管理方面的应用价值进行了展望,旨在丰富生殖抑制的理论,扩展应用实践,为后续的生物资源管理提供理论参考。     

生物学教学中不同脊椎动物红细胞血型的比较和探讨

红细胞血型是以红细胞膜上特异性抗原类型进行区分的,广泛存在于各种动物中。对哺乳纲、鸟纲、爬行纲、两栖纲、鱼纲、圆口纲等不同脊椎动物的红细胞血型系统分类进行了说明、归纳,并对多种动物的血型系统之间存在相似性的现象,以及不同纲的动物之间血型系统的复杂程度进行了比较和探讨。红细胞血型相似程度可反映动物之间的亲缘性,复杂程度可反映不同纲动物之间的进化地位,为生物学教学工作者和研究动物血型的科研工作者提供参考。     

植物大年结实及其与动物贮食行为之间的关系

大年结实(mast seeding)是多生年植物种群周期性同步大量繁殖的一种自然现象。大年结实作为植物适应环境条件、提高繁殖能力的一种策略而备受关注, 但其驱动机制和进化意义尚存在较大争议。在依赖动物扩散种子的植物中, 大年结实被认为是一种调控动物贮食行为、提高种子扩散效率, 并最终增加繁殖成功率的一种策略; 动物介导的植物间互作可能是促进植物共存的进化驱动力。本文简要梳理了大年结实现象的各种假说, 提出了一个包括气候、资源、动植物互作的理解大年结实机制的概念框架, 并着重讨论了大年结实和动物贮食行为之间的关系及其进化和生态意义。建议未来研究需要借助长期生态监测和分子生物学方法, 揭示植物大年结实与动物贮食行为之间的生态与进化过程。     

桓仁滑蜥生殖行为及生殖策略的研究

桓仁滑蜥（Scincella huanrensis）系卵胎生,1年生殖1次,生殖季节为7月下旬到8月上旬。平均每只雌蜥产仔4．556（2～6）只。雌蜥的分娩在洞穴中进行,分娩时间可持续60-90min,仔蜥在产出时卵膜已破裂。雌性采取的是高投入的生殖策略,平均生殖投入率0．436（0．212～0．526）。相关性回归分析表明：窝仔数与雌蜥体重、体长和生殖投入呈正相关关系;生殖投入与雌蜥的体重和体长不相关;幼蜥的体长、体重与窝仔数呈负相关关系。     

被子植物地下结实和地上/下两型结实的生态适应意义

地下结实和地上/下两型结实是被子植物两类独特的结实方式, 多发生在陆生和草本植物中, 主要生长在缺少水分或光照、土壤扰动频繁及环境波动较为剧烈的生境中。两种结实方式不仅是植物适应性进化的重要方面, 也是选择性进化的产物。其中, 地下结实对于植物在母株附近适宜微环境中保存后代、在极端环境下保持种子活力、逃避地面动物取食和火灾伤害以及延长果实发育时间等方面, 地上/下两型结实对于减少同胞子代及种群内竞争、维持和扩大种群以及提高物种的适应力和进化可塑性等方面, 都具有重要的生态适应意义, 是植物抵御不利生物与非生物环境的两类重要防御策略。但两种结实方式同时也存在着限制果实与种子扩散、影响基因传递与种群遗传结构、加大种群隔离以及提高繁殖代价等进化限制, 对物种的分布、种群增长、迁移、适合度和生活史进化等具有重要影响。目前, 地下和地上/下两型结实现象分别在大约24科57属和13科34属中进行了报道, 其中在菊科、十字花科、豆科和玄参科等类群中两种结实现象同时存在。从系统发育看, 地下结实在木兰分支、单子叶植物分支及双子叶植物分支中均存在, 而地上/下两型结实仅出现在单子叶植物分支和双子叶植物分支中, 在被子植物基部类群(ANITA类群)中两种结实方式均不存在。该文对植物地下和地上/下两型结实的类型、系统进化、繁殖特性和扩散对策进行了介绍, 并对其生态适应意义进行了总结, 以期为深入研究植物结实的进化策略提供参考。     

动物的婚配制度

婚配制度是动物种群个体为获得配偶普遍采取的一种行为策略。一般分为单配制、一雄多雌制、一雌多雄制和混交制。作为动物的一种进化稳定对策,婚配制度又具有一定的可塑性,有时可以逆转,有时又是兼性的。配偶外交配主要存在于单配制物种中特别是单配制鸟类中。其发现表明通过观察个体间联系来确定的“社会性婚配制度”和通过个体实际上的交配对象来描述的“遗传性婚配制度”有可能存在差异。婚配制度影响有害动物的不育控制,在存在繁殖性竞争的情况下,对单配制和一雄多雌制而言,不育效果最佳。     

爬行动物体色的快速变化

快速变色是动物对纷繁复杂的自然界高度适应的策略之一。爬行动物中部分有鳞目动物将快速变色表现得淋漓尽致。主要介绍爬行动物快速变色在伪装、通讯和体温调节方面的作用,阐述其中的变色机理。爬行类等动物快速变色在生理体色变化方面由神经系统调节为主而产生生物色和结构色方面的变化。快速变色这种适应策略可能由伪装方面的自然选择、通讯方面的自然选择或性选择进化而来,也可能受体温调节需求的影响,还可能是由这三者同时产生的选择压力进化而来。     

植物nrDNA ITS致同进化不完全现象及其进化意义

nrDNA是个多基因家族,在基因内往往有成千上万的拷贝,但通常只包含有一种序列,即所谓致同进化.因此,nrDNA ITS被广泛应用于植物系统与进化研究当中.近年来,部分研究发现,一些植物类群的基因组内nrDNA ITS也存在着较高的多样性,即致同进化不完全现象.该文就裸子植物和被子植物中nrDNA ITS存在致同进化不完全现象的原因和特点,nrDNA ITS假基因的识别,以及其进化意义等进行了综述.     

Multiple paternity is related to adult sex ratio and sex determination system in reptiles

The adult sex ratio (ASR, the proportion of males in the adult population) is an emerging predictor of reproductive behaviour, and recent studies in birds and humans suggest it is a major driver of social mating systems and parental care. ASR may also influence genetic mating systems. For instance male-skewed ASRs are expected to increase the frequency of multiple paternity (defined here as a clutch or litter sired by two or more males) due to higher rates of coercive copulations by males, and/or due to females exploiting the opportunity of copulation with multiple males to increase genetic diversity of their offspring. Here, we evaluate this hypothesis in reptiles that often exhibit high frequency of multiple paternity although its ecological and life-history predictors have remained controversial. Using a comprehensive dataset of 81 species representing all four non-avian reptile orders, we show that increased frequency of multiple paternity is predicted by more male-skewed ASR, and this relationship is robust to simultaneous effects of several life-history predictors. Additionally, we show that the frequency of multiple paternity varies with the sex determination system: species with female heterogamety (ZZ/ZW sex chromosomes) exhibit higher levels of multiple paternity than species with male heterogamety (XY/XX) or temperature-dependent sex determination. Thus, our across-species comparative study provides the first evidence that genetic mating system depends on ASR in reptiles. We call for further investigations to uncover the complex evolutionary associations between mating systems, sex determination systems and ASR.     

The outcome of sperm competition is affected by behavioural and anatomical reproductive traits in a simultaneously hermaphroditic land snail

Sperm competition is important in species with reproductive strategies that involve multiple mating and prolonged sperm storage such as the simultaneously hermaphroditic land snail Cornu aspersum. Double mating trials in this species have revealed that mating order and courtship behaviour affect paternity success. We investigated the effect of behavioural and anatomical reproductive traits on paternity success from triple mating trials. Triple mating resulted in triple fertilization in 58% of the cases whereas zero paternity was observed in 16% of sperm donors. Third sperm donors achieved higher paternity followed by first and second sperm donors. Snails with a longer epiphallus, the spermatophore forming organ, sired more offspring regardless of their mating order. Genetic compatibility between sperm donor and recipient did not influence paternity success. The results of the present study identified mating order and epiphallus length, as traits affecting the outcome of sperm competition in this species.     

Multiple paternity in reptiles: patterns and processes

The evolution of female promiscuity poses an intriguing problem as benefits of mating with multiple males often have to arise via indirect, genetic, effects. Studies on birds have documented that multiple paternity is common in natural populations but strong evidence for selection via female benefits is lacking. In an attempt to evaluate the evidence more broadly, we review studies of multiple paternity in natural populations of all major groups of nonavian reptiles. Multiple paternity has been documented in all species investigated so far and commonly exists in over 50% of clutches, with particularly high levels in snakes and lizards. Marine turtles and lizards with prolonged pair-bonding have relatively low levels of multiple paternity but levels are nevertheless higher than in many vertebrates with parental care. There is no evidence that high levels of polyandry are driven by direct benefits to females and the evidence that multiple paternity arises from indirect genetic benefits is weak. Instead, we argue that the most parsimonious explanation for patterns of multiple paternity is that it represents the combined effect of mate-encounter frequency and conflict over mating rates between males and females driven by large male benefits and relatively small female costs, with only weak selection via indirect benefits. A crucial step for researchers is to move from correlative approaches to experimental tests of assumptions and predictions of theory under natural settings, using a combination of molecular techniques and behavioural observations.     

Genetic analysis of the mating system and opportunity for sexual selection in northern water snakes (Nerodia sipedon)

We used data collected over 3 years at two study sites to quantifythe rates and consequences of multiple paternity and to determinethe opportunity for selection on male and female northern watersnakes (Nerodia sipedon). We sampled litters from 45 femalesthat gave birth to 811 offspring. Using eight microsatelliteDNA loci (probability of exclusion of nonparental males >0.99), we assigned paternity to 93% of neonates from one studypopulation and 69% of neonates from the other population. Observationsof participation in mating aggregations predicted individual reproductive success poorly for two reasons. First, males regularlycourted nonreproductive females. Second, more than half ofall sexually mature males obtained no reproductive successeach year, despite the fact that many of them participatedin mating aggregations. The number of sires per litter ranged from one to five, with 58% of all litters sired by more thanone male. Multiple paternity increased with female size, apparentlyboth because bigger females mated with more males and becausethe larger litters of big females provide paternity opportunitiesto more males. Multiple paternity was also more prevalent inyears with shorter mating seasons. We detected no advantage to multiple paternity in reducing either the number of unfertilizedovules or stillborn young. Despite the majority of males siringno young each year, some males fathered young with as manyas three different females in one year. Male reproductive successincreased by more than 10 offspring for each additional mate,whereas female success increased by fewer than 2 offspring foreach additional mate. The opportunity for sexual selectionwas more than five times higher in males than females.     

Multiple paternity in an aggregate breeding amphibian: the effect of reproductive skew on estimates of male reproductive success

Aggregate, or explosive, breeding is widespread among vertebrates and likely increases the probability of multiple paternity. We assessed paternity in seven field-collected clutches of the explosively breeding spotted salamander (Ambystoma maculatum) using 10 microsatellite loci to determine the frequency of multiple paternity and the number of males contributing to a female's clutch. Using the Minimum Method of allele counts, multiple paternity was evident in 70% of these egg masses. Simple allele counts underestimate the number of contributing males because this method cannot distinguish multiple fathers with common or similar alleles. Therefore, we used computer simulations to estimate from the offspring genotypes the most likely number of contributing fathers given the distributions of allele frequencies in this population. We determined that two to eight males may contribute to A. maculatum clutches; therefore, multiple paternity is a common strategy in this aggregate breeding species. In aggregate mating systems competition for mates can be intense, thus differential reproductive success (reproductive skew) among males contributing to a female's clutch could be a probable outcome. We use our data to evaluate the potential effect of reproductive skew on estimates of the number of contributing males. We simulated varying scenarios of differential male reproductive success, ranging from equal contribution to high reproductive skew among contributing sires in multiply sired clutches. Our data suggest that even intermediate levels of reproductive skew decrease confidence substantially in estimates of the number of contributing sires when parental genotypes are unknown.     

Unique multiple paternity in the endangered big‐headed turtle (Platysternon megacephalum) in an ex situ population in South China

Understanding the mating system and reproductive strategies of an endangered species is critical to the success of captive breeding. The big‐headed turtle (Platysternon megacephalum) is one of the most threatened turtle species in the world. Captive breeding and reintroduction are necessary to re‐establish wild populations of P. megacephalum in some of its historical ranges in China, where the original populations have been extirpated. However, the captive breeding of P. megacephalum is very difficult and this may be due to its mysterious reproductive strategies and special behavior (e.g., aggressive temperament and territoriality). In this study, we achieved successful captive breeding of P. megacephalum by creating a habitat that mimics natural conditions and then investigated its mating system using microsatellite makers. A total of 16 clutches containing 79 eggs of P. megacephalum were collected, and 52 were hatched successfully over two breeding seasons. Of the 15 effective clutches, 6 clutches (40%) exhibited multiple paternity. There was no significant correlation between clutch size and multiple paternity, and no significant difference in hatching success between multiple‐sired and single‐sired clutches. However, there was significant correlation between male body size and the number of offspring, with higher‐ranked males contributing to more clutches. Our results provide the first evidence of multiple paternity and male hierarchy in P. megacephalum. These findings suggest that multiple paternity and male hierarchy should be considered in captive breeding programs for P. megacephalum, and creating a habitat that mimics natural conditions is an effctive way to achieve successful captive breeding and investigate the mating systems of this species.     

Patterns of multiple paternity and maternity in fishes

The characterization of patterns of multiple mating is a major facet of molecular ecology and is paramount to understanding the evolution of behaviours associated with parental care and mate choice. Over the last 15 years, fishes have been particularly well studied with respect to multiple maternity and paternity thanks to the widespread application of microsatellite markers. The present review focusses on the impressive literature on genetic parentage in fishes. In studies of natural populations, we find that multiple paternity is extremely common across fish species, whereas rates of multiple maternity are much more variable. In species with nest defence, for example, rates of multiple maternity are strongly bimodal, and the occurrence of multiple dams per brood is either rare or the rule. The sex of the care‐giving parent is correlated with the rate of multiple parentage: when males provide uniparental care, rates of multiple paternity are low compared to rates of multiple maternity; when females provide parental care, either alone or assisted by males, rates of multiple paternity are highly variable, whereas rates of multiple maternity are quite low. These patterns may reflect conflicts between the reproductive interests of males and females. We also find that fishes in which females brood the offspring internally display much higher rates of multiple paternity compared to mammals or birds, whereas reptiles are intermediate. Male‐nesting fish species, however, show rates of multiple paternity more similar to those found in other vertebrates. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 735–760.     

Turtle mating systems: behavior, sperm storage, and genetic paternity

As evidenced by the articles in this volume, a recent increase in interest in the mating systems of poikilothermic vertebrates has focused primarily on fishes, a few amphibians, and squamate reptiles. Turtles by contrast have received relatively little attention, yet they display a wide variety of mating behaviors and life-history characteristics that make them excellent candidates for addressing several aspects of genetic parentage that should contribute to a broader understanding of animal reproductive strategies. Here we focus on genetic studies of the mating systems and reproductive patterns of turtles, specifically with respect to multiple paternity and long-term sperm storage. These phenomena highlight the importance of a temporally extended perspective on patterns of individual reproductive success.     

社会性单配制鸟类婚外父权的发生和影响因素

一雌一雄单配制鸟类中,雌性个体与配偶外雄性发生交配的行为称为婚外交配,继而导致了婚外受精产生婚外子代的现象称为产生了婚外父权。婚外父权广泛存在于鸟类中,针对其发生和影响因素已经成为了鸟类行为生态学研究的热点。本文收集了近十年社会性单配制鸟类婚外父权方面的研究文献,从婚外父权的发生及其影响因素两个方面综述了单配制鸟类婚外父权的研究进展。婚外父权发生原因的探讨主要包括：1、从两性的角度探讨雌雄两性在婚外行为中不同的进化繁殖策略。雄性策略旨在增加自身的繁殖输出;有关雌性策略则提出了确保受精假说、食物供给假说、遗传利益假说等,但目前尚存争议;2、在遗传利益假说中较常见的又分为3个假说：“优秀基因”假说、“遗传相容性”假说和“遗传多样性”假说,该三种假说是针对雌性从遗传方面获得的利益而提出的,不断有报道指出雌性配偶选择会被潜在的雄性遗传特性所影响;3、非遗传利益——母系效应影响婚外父权的进化。一些研究指出遗传质量参数,如体重、身体大小、存活率和免疫应答等方面可能会存在母系效应。婚外父权发生的影响因素这里主要指环境因素,包括繁殖同步性、繁殖密度、栖息地环境、产卵及孵化时机等。由于物种不同,受到环境压力不同,导致婚外父权发生率千差万别。最后本文针对未来的研究方向做出了展望。尽管近十年的研究进一步解释了鸟类婚外父权现象,但是该领域仍然存在并且产生了许多新的未解决的问题,而相关实验操作和理论的完善是深入探讨这些问题的关键。     

Multiple paternity in wild-caught Drosophila mojavensis

Female remating frequency and sperm allocation patterns can strongly influence levels of sperm competition and reproductive success in natural populations. In the laboratory, Drosophila mojavensis males transfer very few sperm per copulation and females remate often, suggesting multiple paternity should be common in nature. Here, we examine female sperm loads, incidence of multiple paternity, and sperm utilization by genotyping progeny from 20 wild-caught females at four highly polymorphic microsatellite loci. Based on indirect paternity analyses of 814 flies, we found evidence for high levels of multiple paternity coupled with relatively low reproductive output, consistent with the high levels of female remating predicted in this sperm-limited species. Overall, we found little evidence for last -- male sperm precedence though some temporal variation in sperm utilization was observed, consistent with laboratory findings.