植物根系的身份识别能力、作用及影响机制

自然生境中有些植物的繁殖体传播距离有限, 这类植物更有可能与其亲缘相近的植株相邻。植物个体能否识别邻株的身份并做出相应的反应, 会对植物间的相互作用产生重要影响。亲缘选择理论预测亲缘相近的植物间可以通过亲缘识别和选择作用, 有效地削弱彼此间的干扰和竞争, 从而增加适合度。对植物通过根系进行自我/ 非我和亲缘识别能力、作用及其影响机制进行了综述, 同时指出了当前研究中存在的一些疑点和亟待解决的问题, 对植物身份识别研究的发展方向进行了展望。     

非人灵长类的亲缘选择

亲缘选择是动物进化的重要研究领域之一,非人灵长类因具有丰富的社会网络,是亲缘选择研究领域的重要类群。动物进行亲缘选择的前提是亲缘识别,并常通过社会行为的亲缘偏向表现。因此,本文从非人灵长类的亲缘识别机制和亲缘关系对其社会行为的影响两方面进行了综述:熟悉性和表现型匹配是目前普遍认同的非人灵长类亲缘识别机制,同时这两种机制并不相互排斥,它们可能共同在灵长类的亲缘识别中起作用;在非人灵长类中,亲缘关系是影响社会行为模式的主导因子,它影响着多种灵长类个体的友好行为、攻击行为和性行为的选择,同时亲缘偏向行为在不同物种中表现不尽相同,说明亲缘选择理论可以部分解释灵长类的行为,但存在一定的局限性。本文分析了两种亲缘识别机制的异同以及在实际研究中利用亲缘选择理论解释非人灵长
类社会行为的局限及可能原因。目前,对非人灵长类社会中的亲缘选择研究正逐步深入,其中分子遗传学技术的应用是重要的推动力量。同时,依然存在诸如汉密尔顿规则参数估计和新大陆猴的亲缘选择研究案例的难点,有待研究者进一步探究。     

植物亲缘识别的研究进展

植物的亲缘识别(kin recognition)指植物通过识别周边个体与自己的亲缘关系, 调整自身的生长生态策略、促进亲缘个体的生存与繁衍。研究表明, 植物主要通过特定的叶片挥发物、根系分泌物、感光载体等途径, 识别周边个体与自己的亲缘关系, 改变自身形态学策略(如根系大小、根冠比、种子数量等)或者生理代谢策略(次生代谢物质、防御蛋白等), 调整与周边个体的竞争强度, 缓和与近亲缘个体之间的竞争, 加强与远亲缘或非亲缘个体的竞争。同时亲缘识别的强度也受环境因子(养分等)的影响。结合目前的研究进展, 该文分析了导致亲缘识别的研究结果存在差异或争议的主要原因, 认为主要与实验材料的选择、亲缘关系的界定标准、环境条件及测定的指标不统一有关。将来的研究应重点从生理生化、分子、代谢水平上深入研究植物亲缘识别的机理。     

动物辨别亲属的机制

亲缘识别在动物界中是很普遍的现象。不同的动物有不同的手段识别亲属。亲缘辨别的机制主要有空间识别机制,熟悉机制,表形匹配和识别等位基因4种机制,有的动物只有一种机制起作用,有的则是两种机制同时共存或不同时期不同机制。此外,还概述了亲缘辨别的一些功能。     

近缘细菌细胞间的相互识别与相互作用

亲缘识别是细菌细胞间竞争与合作的前提和基础。细菌通过亲缘识别分辨自我细胞和非自我细胞;非同类的细菌细胞相互分离或被排除,而亲缘种群内的细菌细胞进行群体运动、生物膜和子实体形成等社会性合作行为。细菌的自我识别机制可能有助于不同亲缘类群在混杂的自然系统中的共存。近些年来,细菌亲缘识别及相互作用的机制研究工作成为热点,本文总结了近缘细菌细胞间相互识别和作用机制的研究进展。     

动物的亲缘选择和亲缘识别

很多不同类群的动物能够识别亲属和非亲属,甚至远亲和近亲,并表现出行为上的差异,动物怎样完成这种高度准确性分辨是自60年代末期以来日趋活跃的研究领域。亲缘识别(Kin Recognifion)的作用主要在于使重亲行为成为可能,而且可以避免近亲繁殖。但在我国动物亲缘识别的研究几乎还是空白。     

布氏田鼠合作贮食行为及其影响因素

合作贮食是社会性动物应对食物匮乏的重要对策,但其内在的生态学机制尚未阐明。本文以布氏田鼠为研究对象,根据亲缘关系划分为亲缘组和非亲缘组。在人工实验箱中录像统计不同组别合作贮食行为占用时间的差异,探究亲缘关系对合作贮食行为的影响,并利用相关性分析探究亲缘组个体对贮食的贡献—收益关系以及与贮食行为相关的个体特征因素。结果表明,在组间水平上,与非亲缘组相比,亲缘组的布氏田鼠表现更多的合作贮食。在亲缘组个体水平上,不同个体对贮食的参与度和贡献度存在很大差异。个体对合作贮食的付出与其取食的收益呈显著负相关关系,并且个体合作贮食贡献越大,其睾丸指数和睾酮含量降低越显著。研究结果说明,亲缘关系的存在有利于布氏田鼠的合作贮食,但合作贮食贡献大的个体,其繁殖受到了抑制。     

亲缘关系与啮齿类动物的社会行为

在漫长的进化过程中 ,动物除了在形态结构、生理生化以及遗传等方面适应周围环境之外 ,也产生了与环境相适应的行为 ,其中种内个体社会联系的行为称为社会行为。由于自然选择提高了个体的生存价值和广义适合度 (ｉｎｃｌｕｓｉｖｅｆｉｔｎｅｓｓ) ,所以如果个体间亲缘程度不同 ,在自然选择的作用下必然会产生不同的结果。亲缘关系与社会行为的研究源于行为生态学 ,其中以Ｈａｍｉｌｔｏｎ[32 ] 提出的亲缘选择和广义适合度理论为代表 ,将亲缘关系和自然选择联系起来 ,扩展了达尔文自然选择理论的范围 ,较好地解释了利他行为 (ａｌｔｒｕｓ…     

不同密度下拟南芥的亲缘选择

亲缘选择理论认为亲缘个体之间通过亲缘认知而相互合作以增加其适合度。前期的相关研究证实了一些植物物种之间具有亲缘选择作用,而且不同物种对亲缘选择的响应方式各有差异。但是,现有的试验物种较少,且未考虑其他环境因子的影响。为了验证拟南芥(Arabidopsis thaliana)不同生态型对亲缘选择的响应方式,本试验选取3个拟南芥生态型[Columbia(Col)、Landsbergerecta(Ler)和Wassileskijia(Ws)]作为研究对象,在3种密度条件下,通过测度植物表型特征(株高、莲座面积、叶面积和种子数等)和各构件生物量特征(根、茎、叶和种子生物量)在亲缘处理组和非亲缘组的差异,来探索密度对亲缘选择结果的影响。结果表明,拟南芥各生态型亲缘组与非亲缘组差异主要表现在地上部分,而地下部分在亲缘和非亲缘组并没有显著差异;3种生态型拟南芥对亲缘选择的响应方式不同,Col主要表现在种子生物量上,Ler主要表现在叶和茎生物量上,Ws主要表现在种子和茎生物量上;拟南芥不同生态型对亲缘选择的响应方式依赖于种植密度,密度是影响拟南芥亲缘选择表现的重要因素。     

动物的亲缘选择

总结了亲缘选择的概念并对亲缘系数的计算方法进行了归纳，对利他行为、适合度、广义适合度、亲缘系数的基本概念进行了阐述，另外还总结了基于亲缘选择的利他行为发生的条件，说明了亲缘选择在人类家族企业中的应用，对亲缘选择理论的学术价值和局限性进行了讨论。     

Fitness consequences of plants growing with siblings: reconciling kin selection, niche partitioning and competitive ability

Plant studies that have investigated the fitness consequences of growing with siblings have found conflicting evidence that can support different theoretical frameworks. Depending on whether siblings or strangers have higher fitness in competition, kin selection, niche partitioning and competitive ability have been invoked. Here, we bring together these processes in a conceptual synthesis and argue that they can be co-occurring. We propose that these processes can be reconciled and argue for a trait-based approach of measuring natural selection instead of the fitness-based approach to the study of sibling competition. This review will improve the understanding of how plants interact socially under competitive situations, and provide a framework for future studies.     

Kin recognition in an annual plant

Kin recognition is important in animal social systems. However, though plants often compete with kin, there has been as yet no direct evidence that plants recognize kin in competitive interactions. Here we show in the annual plant Cakile edentula, allocation to roots increased when groups of strangers shared a common pot, but not when groups of siblings shared a pot. Our results demonstrate that plants can discriminate kin in competitive interactions and indicate that the root interactions may provide the cue for kin recognition. Because greater root allocation is argued to increase below-ground competitive ability, the results are consistent with kin selection.     

Recent advance of kin recognition in plant

Plants have the ability to discriminate kin members from strangers in competitive interactions and show altruistic behavior towards related individuals. Studies have showed that plants recognize their neighbors and adjust their ecological strategy mainly through leaf volatiles, root secretions and photographic carrier. The target plants can modify their morphological traits (root size, root:shoot ratio, seed numbers etc.) or metabolism characteristics (secondary metabolites, defense-related proteins etc.) when groups of plants shared common resources, so as to minimize competition with close relatives. The density of kin recognition is influenced by environmental conditions. The main reasons for controversial experimental results of kin recognition are associated with plant materials, standard of kin selection, ecological factors and measured indices. Further studies are required to understand the mechanisms of kin interactions in plants from physiological, biochemical, molecular and metabolic levels.     

Kin recognition: Competition and cooperation in Impatiens (Balsaminaceae)

The ability to recognize kin is an important element in social behavior and can lead to the evolution of altruism. Recently, it has been shown that plants are capable of kin recognition through root interactions. Here we tested for kin recognition in a North American species of Impatiens that has a high opportunity of growing with kin and responds strongly to aboveground competition. We measured how the plants responded to the aboveground light quality cues of competition and to the presence of root neighbors and determined whether the responses depended on whether the neighbors were siblings or strangers. The study families were identified by DNA sequencing as members of the same species, provisionally identified as Impatiens pallida (hereafter I. cf. pallida). We found that I. cf. pallida plants were capable of kin recognition, but only in the presence of another plant's roots. Several traits responded to relatedness in shared pots, including increased leaf to root allocation with strangers and increased stem elongation and branchiness in response to kin, potentially indicating both increased competition toward strangers and reduced interference (cooperation) toward kin. Impatiens cf. pallida responded to both competition cues simultaneously, with the responses to the aboveground competition cue dependent on the presence of the belowground competition cue.     

Relatives growing together: pair density and kinship

Kin selection is often used to explain the evolution altruism towards relatives through favouring the evolution of kin recognition. However, it remains unclear whether kin recognition is affected by plant pair density and different degrees of relatedness. A two-factor experimental design of kinship (three kinship degrees including siblings, closely related strangers and distantly related strangers) and pair density (including relative small, medium and large pair densities) was conducted in this study. Plant competitive traits including rosette size, specific leaf area (SLA), stem elongation, root and leaf allocation, seed biomass and vegetative biomass were measured to reflect interactions among plants living with different relatives of Arabidopsis thaliana accessions [Columbia (Col-0), Landsberg erecta (Ler) and Wassileskijia (Ws)] in three different pair densities. The SLA only showed kinship effect, and siblings showed higher SLA than non-siblings in each pair density. The plant total biomass was only affected by pair density, which increased with decreases of pair density. The rosette size, stem elongation, root allocation and leaf allocation showed interactive effects of kinship and pair density. In the large pair density, the rosette size of siblings was lower than distantly related strangers, compared to closely related strangers; the stem elongation and root allocation were lower, while the seed biomass of siblings was higher than the closely or distantly related strangers. In the medium pair density, plants living with siblings or with closely related neighbours showed higher root allocation than with distantly related neighbours. In the small pair density, the plant rosette acted similarly to of which in the large pair density, and siblings showed higher root allocation than the two strangers. The other traits in each pair density showed no significant differences among kinship treatments. A relatively large pair density achieves kin recognition by deducing root competition ability and mutual shading, with increased efficient light capture and fitness. Similar such root and efficient light capture strategies were selected in medium pair density. Except the efficient light capture strategies, small pair density also displays allocation trade-offs between roots (decreased) and leaves (increased) for siblings. Moreover, kin responses on those attributes are also adjusted by kinship degree. Thus, kinships and pair density are important variables to mediate kin interactions.     

Kinship rivalry does not trigger specific allocation strategies in Lupinus angustifolius

Background and Aims

Research on the ability of plants to recognize kin and modify plant development to ameliorate competition with coexisting relatives is an area of very active current exploration. Empirical evidence, however, is insufficient to provide a sound picture of this phenomenon.

Methods

An experiment was designed to assess multi-trait phenotypic expression in response to competition with conspecifics of varied degrees of genealogical relatedness. Groups of siblings, cousins and strangers of Lupinus angustifolius were set in competition in a pots assay. Several whole-plant and organ-level traits, directly related to competition for above- and below-ground resources, were measured. In addition, group-level root proliferation was measured as a key response trait to relatedness to neighbours, as identified in previous work.

Key Results

No major significant phenotypic differences were found between individuals and groups that could be assigned to the gradient of relatedness used here. This occurred in univariate models, and also when multi-trait interactions were evaluated through multi-group comparisons of Structural Equation Models. Root proliferation was higher in phenotypically more heterogeneous groups, but phenotypic heterogeneity was independent of the relatedness treatments of the experiment, and root proliferation was alike in the neighbourhoods of siblings, cousins and strangers.

Conclusions

In contrast to recent findings in other species, genealogical relatedness to competing neighbours has a negligible impact on the phenotypic expression of individuals and groups of L. angustifolius. This suggests that kin recognition needs further exploration to assess its generality, the ecological scenarios where it might have been favoured or penalized by natural selection, and its preponderance in different plant lineages.     

Kin recognition, not competitive interactions, predicts root allocation in young Cakile edentula seedling pairs

? Recent studies have demonstrated sibling vs stranger differences in group root allocation in plants, suggesting that plants have the potential for kin discrimination in competition. However, morphology differences could potentially be generated by competition-based mechanisms. Here, we tested these hypotheses for the sibling vs stranger differences in root allocation in Cakile edentula. ? Seeds were planted in pairs of either kin (siblings) or strangers, from all combinations of eight families, to give eight kin (sibling) and 28 stranger pair identities. Because the species has a seed dimorphism, the 10 replicates of each pair identity included both seed types. Root allocation, size inequality between seedlings in a pair, and competitive ability were derived from measures of biomass and height. ? Cakile edentula seedlings demonstrated the same kin recognition response previously observed in juvenile plants, with lower root allocation in kin pairs than stranger pairs. The seed dimorphism was not associated with root allocation. ? The two competitive mechanisms, genetic differences in competitive ability and increased size inequality in stranger groups, did not explain the root allocation differences in these seedlings. Kin recognition offered the most probable explanation for the differences in root allocation between sibling and stranger pairs.     

Kin distribution of amphibian larvae in the wild

According to kin selection theory, the location of an individual with respect to its relatives can have important ramifications for its fitness. Perhaps more than any other vertebrate group, anuran amphibian larvae have been the subject of many experiments on this topic. Some anuran species have been shown in the laboratory to recognize and associate with their siblings and half-siblings. However, due to the difficulty of identifying sibships, no kinship studies with anuran larvae have been conducted in the wild. Here, we use microsatellite analysis to show that wood frog (Rana sylvatica) tadpoles were nonrandomly distributed in two ponds with respect to their relatives. In one pond, the tadpoles were significantly clumped with their siblings or half-siblings as expected from other published laboratory studies on this species. However, in another pond, the tadpoles were significantly nonrandomly dispersed from their siblings or half-siblings. This is the first example of kin repulsion of nonreproductive animals in the wild and the first time a species has been shown to display both aggregation and repulsion under different circumstances. These results suggest that kin distribution is context dependent and demonstrate the importance of testing kin selection hypotheses under natural conditions.     

Inclusive fitness,asymmetric competition and kin selection in plants

The findings that some plants alter their competitive phenotype in response to genetic relatedness of its conspecific neighbour (and presumed competitor) has spurred an increasing interest in plant kin‐interactions. This phenotypic response suggests the ability to assess the genetic relatedness of conspecific competitors, proposing kin selection as a process that can influence plant competitive interactions. Kin selection can favour restrained competitive growth towards kin, if the fitness loss from reducing own growth is compensated by increased fitness in the related neighbour. This may lead to positive frequency dependency among related conspecifics with important ecological consequences for species assemblage and coexistence. However, kin selection in plants is still controversial. First, many studies documenting a plastic response to neighbour relatedness do not estimate fitness consequences of the individual that responds, and when estimated, fitness of individuals grown in competition with kin did not necessarily exceed that of individuals grown in non‐kin groups. Although higher fitness in kin groups could be consistent with kin selection, this could also arise from mechanisms like asymmetric competition in the non‐kin groups. Here we outline the main challenges for studying kin selection in plants taking genetic variation for competitive ability into account. We emphasize the need to measure inclusive fitness in order to assess whether kin selection occurs, and show under which circumstances kin selected responses can be expected. We also illustrate why direct fitness estimates of a focal plant, and group fitness estimates are not suitable for documenting kin selection. Importantly, natural selection occurs at the individual level and it is the inclusive fitness of an individual plant – not the mean fitness of the group – that can capture if a differential response to neighbour relatedness is favoured by kin selection.