根系隔离条件下的谷子亲缘识别

在根系隔离情况下, 通过研究邻株身份(亲缘株、非亲缘株、陌生株)及其与种植密度(高、低)和土壤养分水平(高、低)交互作用对谷子(Setaria italica)形态学特征和生物量分配的影响, 探索谷子地上部分是否能够识别亲缘邻株, 以及谷子的这种亲缘识别能力对环境因子如何响应。结果显示: 1)亲缘组谷子叶生物量分配显著降低, 茎秆显著增粗, 暗示着亲缘组谷子植株间减少竞争, 并增强对当地多风气候的适应。而非亲缘组谷子叶生物量分配显著增加, 表明非亲缘组谷子植株间竞争较强。2)与非亲缘组相比, 陌生组谷子种子生物量分配显著增加, 株高显著减少, 表明陌生组谷子植株通过不对称竞争(与邻株糜(Panicum miliaceum)植株相比, 株高显著增加), 进一步限制邻株(糜)生长, 从而增强竞争能力, 同时, 将更多的生物量投资于繁殖, 增加适合度。3)在高密度种植条件下, 谷子茎生物量和叶生物量分配在各邻株身份处理间无显著差异, 而在低密度种植条件下, 与非亲缘组相比, 亲缘组谷子茎生物量显著增加, 叶生物量分配显著减小; 随着种植间距的增大(种植密度减小), 亲缘组谷子叶生物量分配显著减少, 而非亲缘组和陌生组叶生物量分配在高、低种植密度条件下无显著差异。4)在低土壤养分条件下, 亲缘组和非亲缘组谷子叶生物量分配无显著差异, 前者穗长显著小于后者, 而在高土壤养分条件下, 亲缘组谷子叶生物量分配显著小于非亲缘组, 前者穗长显著大于后者。结果表明, 在根系隔离的情况下,谷子能够识别亲缘邻株, 且谷子地上部分竞争信号在亲缘识别过程中扮演重要角色。较低种植密度和较高土壤养分水平有利于谷子亲缘识别能力的表达。     

Kin recognition in Setaria italica under the condition of root segregation

Aims Kin recognition may play an important part in the performance and productivity of crop plants. However, so far, little is known about whether crop plants can recognize their kin neighbors. The aim of this study was to explore kin recognition in Setaria italica, and its responses to changes in environmental and biological conditions.Methods A field experiment was conducted in the suburb of Shanghai. Setaria italica grew with different neighbors (kin, non-kin and strangers), under the condition of root segregation and different plant densities (high and low) and soil nutrient levels (high and low), respectively. We investigated how neighbor identity and its interactions with plant density and soil nutrient level affected the morphology and biomass allocation of S. italica.Important findings Under the condition of root segregation, 1) Leaf biomass allocation and stem diameter of plants in the kin groups significantly decreased and increased, respectively, suggesting that plants of S. italica in the kin groups reduced inter-individual competition, and adapted to the local windy climate. 2) Compared with the non-kin groups, plants in the stranger groups significantly increased the biomass allocation to seeds, while plant height decreased significantly, suggesting that the plants of S. italica in the stranger groups may reduce the growth of their neighbors due to asymmetric competition (S. italica significantly increased height compared with the neighboring plants, Panicum miliaceum). Therefore, the S. italic plants in the stranger groups allocated more biomass to reproduction and increased fitness than those in non-kin groups. 3) Under the condition of high plant density, no significant differences were found in stem biomass and leaf biomass allocation of plants among different neighbor identity treatments. While under the condition of low plant density, compared with the non-kin groups, biomass allocation to stem and leaf in the kin groups significantly increased and decreased, respectively. As the plant density decreased, plants in the kin groups decreased leaf biomass allocation significantly, while plants in the non-kin and stranger groups did not show such a response. 4) Under the condition of low soil nutrient level, no significant difference was found in leaf biomass allocation between the kin and non-kin groups, while the ear length of plants in the kin groups decreased significantly. Under the condition of high soil nutrient level, the biomass allocated to leaves in the kin groups decreased significantly, while ear length increased significantly compared with the non-kin groups. Therefore, under the condition of root segregation, plants of S. italica showed the ability to recognize their kin neighbors, and the aboveground competitive cues may play important roles in the course of kin recognition in S. italica. Lower plant density and higher soil nutrient level may facilitate the ability of kin recognition in S. italica.     

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.     

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。     

Kin and sex recognition in a dioecious grass

Plants have evolved complex mechanisms to recognize and respond to the presence of neighboring plants, and the genetic identity of a neighbor has been shown to make a difference in this response. Studies have found that plants are able to differentiate among self- versus non-self and among sibling (kin) competitors. Here, we present data for the dioecious grass Distichlis spicata on seedling recognition of kin and sex. D. spicata exhibits extreme spatial segregation of the sexes (SSS) in the field, and previous work has shown that intra-sexual competition is less than inter-sexual competition in the field. In this experiment, we conducted experiments in the lab, exposing the seedlings to liquid media in which seedlings had been previously grown, rather than have the seedling physically contact one another. We found that inter-sexual interactions caused a decrease in the total dry weight and an increase in root/shoot ratio of the plants compared with intra-sexual interactions. These findings suggest that D. spicata plants can recognize and respond to plant sex and that inter-sexual competition contributes to SSS, even when additional interactions, such as mycorrhizal fungi are controlled, and physical interactions between plants are removed. In the kin recognition analysis, we found that plants paired with another plant from the same mother had significantly greater lateral root number and length than plants paired with non-kin, suggesting that in this highly clonal grass, kin recognition may be an important mechanism in competitive interactions.     

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.     

植物的亲缘选择

亲缘选择是指在一个随机交配群体中的个体基于亲缘关系而以一种非随机性的方式相互作用,其作用结果是亲缘个体得到更大的广义适合度。综述了亲缘选择和亲缘竞争两种观点以及各自的试验支持证据;分析了导致亲缘选择试验结果出现分歧的原因,认为这主要是由于对亲缘选择理解上的模糊以及试验设计的不严谨所致。植物间的亲缘选择研究不仅相对较少,对亲缘选择的机制研究更为欠缺,这就造成了目前对此问题在科学认识上出现不少盲点。综合前期研究,提出今后对亲缘选择的研究应该首先界定"亲缘"程度,同时改良试验设计方案,选择多种不同生境下的物种对亲缘选择进行深入研究,并且考虑环境因子对植物亲缘选择的影响。同时,对植物亲缘识别机制的研究应该从生理生化方面出发,通过定性定量地分析探索植物根系分泌物在植物亲缘识别中的作用和作用途径。     

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.     

Plant adaptation to dynamically changing environment: the shade avoidance response

The success of competitive interactions between plants determines the chance of survival of individuals and eventually of whole plant species. Shade-tolerant plants have adapted their photosynthesis to function optimally under low-light conditions. These plants are therefore capable of long-term survival under a canopy shade. In contrast, shade-avoiding plants adapt their growth to perceive maximum sunlight and therefore rapidly dominate gaps in a canopy. Daylight contains roughly equal proportions of red and far-red light, but within vegetation that ratio is lowered as a result of red absorption by photosynthetic pigments. This light quality change is perceived through the phytochrome system as an unambiguous signal of the proximity of neighbors resulting in a suite of developmental responses (termed the shade avoidance response) that, when successful, result in the overgrowth of those neighbors. Shoot elongation induced by low red/far-red light may confer high relative fitness in natural dense communities. However, since elongation is often achieved at the expense of leaf and root growth, shade avoidance may lead to reduction in crop plant productivity. Over the past decade, major progresses have been achieved in the understanding of the molecular basis of shade avoidance. However, uncovering the mechanisms underpinning plant response and adaptation to changes in the ratio of red to far-red light is key to design new strategies to precise modulate shade avoidance in time and space without impairing the overall crop ability to compete for light.     

Competition intensity along a productivity gradient in a low-diversity grassland

Competition among plants often shifts from roots to shoots as productivity increases and species composition changes. We examined competition in an old field with low diversity to test whether this shift occurred along a productivity gradient without species turnover. Forty plots received one of four nitrogen treatments (0, 5, or 15 g added N m-2 yr-1 or 400 g m-2 yr-1 of sawdust added to immobilize N) annually for 5 yr. All N levels were dominated by the perennial grasses Agropyron cristatum and Bromus inermis. Transplants of Agropyron were grown with all neighbors, roots of neighbors, or no neighbors present to measure total, root, and shoot competition. Transplant growth was 22%-165% higher in subplots without neighbors present, which indicates that competition occurred. Competition from neighbor roots was primarily responsible for suppression of transplant growth over the entire productivity gradient. In contrast to previous field experiments that found either an increase in total competition intensity or a shift from root to shoot competition with increasing productivity, we found neither. Increases in total competition intensity or shifts from root to shoot competition found along other gradients may be caused by changes in species composition and not by increased resources or neighbor biomass. These results suggest that different competitive mechanisms may operate in low-diversity vegetation than in more diverse natural vegetation.     

植物种间和种内的化学作用

植物间相互作用是生态学基础科学问题之一,植物能感受和识别共存同种或异种植物,进而调整生长、繁殖和防御策略。植物种间和种内的感受和识别大多是由植物产生释放的次生物质所介导,这类化学识别通讯可以启动相应的植物化感作用机制。近年发现,植物亲属间也存在着化学识别、地下根系通讯调控地上开花繁殖等植物种间和种内的化学作用关系。目前植物通过地上挥发物介导的植物化学作用已基本澄清,但根分泌物介导的植物地下化学作用机制及其信号物质还所知甚少。地下化学作用不仅决定根系侵入(接近)和躲避(排斥)行为,也能调控地上开花时间和花期。这样,植物间的化学作用还涉及植物地下和地上的协调互作。本文以植物化感作用和植物化学识别通讯及相应的化感物质和信号物质为基点,从植物亲属识别、根系化学识别和行为模式、地下化学作用调控地上开花繁殖3个方面综述植物种间和种内化学作用的研究进展,为全面理解植物间相互作用提供新视野。     

谷子的亲缘识别能力及其与种植密度和土壤养分水平的关系

为探明作物是否具有识别邻株身份的能力以及这种能力是否受到环境因子的调控,通过大田试验,研究邻株身份(亲缘株、非亲缘株和陌生株)、种植密度和土壤养分水平的交互效应对谷子(Setaria italica)地上部分生物量分配的影响。结果表明,谷子与亲缘株为邻时的净繁殖生物量分配和种子生物量分配,比与非亲缘株为邻时显著提高,且营养生物量分配显著降低(P<0.05)。在高种植密度条件下,亲缘组谷子的穗长、净繁殖生物量分配和种子生物量分配显著大于非亲缘组,而营养生物量分配显著小于非亲缘组(P<0.05)。随着土壤养分水平提高,亲缘组谷子的种子生物量分配显著增加,营养生物量分配显著减少(P<0.05)。由此推断,谷子具有对亲缘邻株的识别能力,且这种能力受种植密度和土壤养分水平的调控,在高种植密度和高土壤养分水平条件下,谷子的亲缘邻株识别能力较强。     

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.     

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.     

A test of alternative hypotheses for kin recognition in cannibalistic tiger salamanders

The function of kin recognition is controversial. We investigatedthe adaptive significance of kin discrimination in cannibalistictiger salamander larvae, Ambystoma tigrinum. Previous laboratoryexperiments show that cannibals preferentially consume lessrelated individuals. We hypothesized that this example of kinrecognition (1) is a laboratory artifact, (2) is a by-productof sibship-specific variation in escape responses, because cannibalsfrom families with rapid responses may be more likely to cannibalize slowlyescaping non-kin, (3) is an epiphenomenon of species recognition,(4) functions in disease avoidance, because kin may be moreinfectious than non-kin, or (5) is favored by kin selection.We evaluated these five hypotheses by using laboratory and fieldexperiments to test specific predictions made by each hypothesis.We rejected hypotheses 1-4 above because (1) kin recognitionwas expressed in the wild, (2) escape responses did not reliablypredict whether a cannibal would ingest kin or non-kin, (3)kin recognition was not most pronounced in populations wheretiger salamanders co-occur with other species of salamanders,and (4) non-kin prey were more likely than kin to transmit pathogensto cannibals. However, we established that the necessary conditionfor kin selection, Hamilton's rule, was met. Thus, our resultsimplicate kin selection as the overriding reason that cannibalistictiger salamanders discriminate kin.     

Differential effect of inter- and intraspecific competition on the performance of invasive and native Taraxacum species

Inter- and intraspecific competitive abilities are significant determinants of invasive success and the ecological impact of non-native plants. We tested two major hypotheses on the competitive ability of invasive species using invasive (Taraxacum officinale) and native (T. platycarpum) dandelions: differential interspecific competitive ability between invasive and native species and the kin recognition of invasive species. We collected seeds from two field sites where the two dandelion species occurred nearby. Plants were grown alone, with kin (plants from the same maternal genotype) or strangers (plants from different populations) of the same species, or with different species in a growth chamber, and the performance at the early developmental stage between species and treatments was compared. The invasive dandelions outcompeted the native dandelions when competing against each other, although no difference between species was detected without competition or with intraspecific competition. Populations of native species responded to interspecific competition differently. The effect of kinship on plant performance differed between the tested populations in both species. A population produced more biomass than the other populations when grown with a stranger, and this trend was manifested more in native species. Our results support the hypothesis that invasive plants have better competitive ability than native plants, which potentially contributes to the establishment and the range expansion of T. officinale in the introduced range. Although kin recognition is expected to evolve in invasive species, the competitive ability of populations rather than kinship seems to affect plant growth of invasive T. officinale under intraspecific competition.     

When competition does not matter: grassland diversity and community composition

We examined whether the intense root competition in a rough fescue grassland plant community in central Alberta, Canada, was important in structuring plant species diversity or community composition. We measured competition intensity across gradients of species richness, evenness, and community composition, using pairs of naturally occurring plants of 12 species. One plant in each pair was isolated from neighbors to measure competition; community structure and environmental conditions were also measured at each pair. We used structural equation modeling to examine how competition influenced community structure. Competition intensity was unrelated to species richness and community composition, but increased competition intensity was associated with a slight decline in evenness. Size-symmetric root competition was probably unimportant in structuring this plant community because there are no feedback mechanisms through which size-symmetric competition can magnify small initial differences and eventually lead to competitive exclusion. In plant communities with little shoot competition, competition and community structure should be unlinked regardless of competition intensity. In more productive systems, we propose that interactions between root and shoot competition may indirectly structure communities by altering the overall asymmetry of competition.     

Kin discrimination and altruism in the larvae of a solitary insect

Kin selection theory predicts altruism between related individuals, which requires the ability to recognize kin from non-kin. In insects, kin discrimination associated with altruistic behaviour is well-known in clonal and social species but in very few solitary insects. Here, we report that the solitary larvae of a non-social insect Aleochara bilineata Gyll. (Coleoptera; Staphylinidae) show kin discrimination and sibling-directed altruistic behaviour. Larvae superparasitize more frequently the hosts parasitized by non-kin individuals than those hosts parasitized by siblings. Kin discrimination probably occurs by self-referent phenotype matching, where an individual compares its own phenotype with that of a non-familiar related individual, a mechanism rarely demonstrated in animals. The label used to recognize kin from non-kin corresponds to substances contained in the plug placed on the hosts by the resident larvae during the parasitization process. Kin competition induced by a limited larval dispersion may have favoured the evolution of kin recognition in this solitary species.     

Growing with siblings: a common ground for cooperation or for fiercer competition among plants?

Recent work has shown that certain plants can identify their kin in competitive settings through root recognition, and react by decreasing root growth when competing with relatives. Although this may be a necessary step in kin selection, no clear associated improvement in individual or group fitness has been reported to qualify as such. We designed an experiment to address whether genetic relatedness between neighbouring plants affects individual or group fitness in artificial populations. Seeds of Lupinus angustifolius were sown in groups of siblings, groups of different genotypes from the same population and groups of genotypes from different populations. Both plants surrounded by siblings and by genotypes from the same population had lower individual fitness and produced fewer flowers and less vegetative biomass as a group. We conclude that genetic relatedness entails decreased individual and group fitness in L. angustifolius. This, together with earlier work, precludes the generalization that kin recognition may act as a widespread, major microevolutionary mechanism in plants.     

Fast seedling root growth leads to competitive superiority of invasive plants

The competitive superiority of invasive plants plays a key role in the process of plant invasions, enabling invasive plants to overcome the resistance of local plant communities. Fast aboveground growth and high densities lead to the competitive superiority of invasive species in the competition for light. However, little is understood of the role belowground root competition may play in invasion. We conducted an experiment to test the effect of root growth on the performance of an invasive shrub Cassia alata, a naturalized, non-invasive shrub Corchorus capsularis, and a native shrub Desmodium reticulatum. We compared seedling growth of the three species and their competitive ability in situ. The roots of the C. alata seedlings grew much faster than those of C. capsularis and D. reticulatum during the entire growth period although C. alata had shorter shoots than D. reticulatum. Furthermore, C. alata showed an apparent competition advantage compared to the other two species as evidenced by less biomass reduction in intraspecific competition and higher competitive effects in interspecific competition. Our study reveals that fast seedling root growth may be important in explaining the competitive advantages of invasive plants. Future studies should pay more attention to the belowground traits of invasive plants, the trade-off between shoot and root growth, and the role of root competition in affecting the population dynamics of invasive plants and the structures of invaded communities.