共生理论视角下创新农业生态经济研究范式

共生理论认为异质共生生物在其生存过程中会表现出相互依存的关系。试图将这一论点移植到农业生态经济研究中,用以处理农业生产中生态效益与经济效益之间矛盾的问题。为此,把农业生态经济视为由生态单元与经济单元组成的异质共生体,分析了该共生体的结构与共生模式,并采用Logistic方程与数值模拟方法,探讨农业生态经济共生体的共生机制,揭示共生单元间的演化规律与成长特征。研究表明:共生模式在农业实践上创新应用的主要目标是将共生体从寄生共生模式向互惠共生模式转型,并提出正向转型的激励策略。共生单元的选择必须具有兼容性,共生单元间有明确的共生界面,促使物质、能量和信息的交流,以增加共生体内自由能。共生关系正向发展要遵循共生演进规律,偏利共生模式是演进到互惠共生模式的必经过程。培育互惠共生适存的共生环境对促进共生体演进至关重要。这些研究成果为农业生态经济研究提供一个新范式,并开创一条研究新思路与方法。     

Evolution of dispersal in metacommunities of interacting species

Theoretical studies on the evolution of dispersal in metacommunities are rare despite empirical evidence suggesting that interspecific interactions can modify dispersal behaviour of organisms. To understand the role of species interactions for dispersal evolution, we utilize an individual‐based model of a metacommunity where local population dynamics follows a stochastic version of the Nicholson–Bailey model and dispersal probability is an evolving trait. Our results show that in comparison with a neutral system (commensalism), parasitism promotes dispersal of hosts and parasites, while mutualism tends to reduce dispersal in both partners. Search efficiency of guests (only in the case of parasitism), dispersal mortality and external extinction risk can influence the evolution of dispersal of all partners. In systems composed of two host and two guest species, lower dispersal probabilities evolve under parasitism as well as mutualism than in one host and one guest species systems. This is because of frequency‐dependent modulations of dispersal benefits emerging in such systems for all partners.     

从细胞内共生到多尺度共生：回顾与展望

基于共生概念的历史变化,目前人们普遍接受了广义共生概念。即共生是包含互利共生（mutualism）、偏利共生（commensalism）和拮抗/寄生（antagonism/parasitism）的共生连续体。本文简述了近20年间,全球9次国际共生学术大会取得的重要成果,对细胞内共生、时间、空间以及多种互作尺度共生关系的研究利用进展进行了评述。同时展望了一些活跃共生领域的研究概况,如共生失调 （dysbiosis）、植物-微生物-昆虫三角共生关系（plant-microbe-insect triangle）、细菌-真菌互作（bacterial- fungal interaction,BFI）、菌根菌-真菌内生细菌-植物多方共生联盟（multipartite symbiosis consortium）以及与共生相关微生物组的集合群落（metacommunity）研究及应用等。共生（symbiosis）正成为当代生物学的核心原则,正以一种与更宏大系统方法相一致的概念,从根本上改变了传统上的一些生物学概念,如孤立性的个体（individuality）概念。基因组测序和高通量RNA技术分析揭示,动、植物与共生微生物的重要互作,打破了迄今为止生物个体的特征边界,挑战了这些学科的定义;共生不仅是一对一的互利共生关系,共生实际是多种共生模式的连续共生体。此外,植物-昆虫-微生物互作的三角关系;菌根-真菌-真菌内生细菌-植物的多方联盟等新关系的发现,更把生命科学推向了快速发展的方向。这些科学进展不仅对生物科学的遗传学、免疫学、进化、发育、解剖学和生理学的研究至关重要,拓宽了新的视野,而且对农业中生物制剂研发,人类微生物组的管理及调控,以及对发酵食品及工业微生物生产的设计和管理将产生积极影响。     

Evolutionary double suicide in symbiotic systems

Mutualism is thought to face a threat of coextinction cascade because the loss of a member species could lead to the extinction of the other member. Despite this common emphasis on the perils of such knock-on effect, hitherto, the evolutionary causes leading to extinction have been less emphasised. Here, we examine how extinction could be triggered in mutualism and whether an evolutionary response to partner loss could prevent collateral extinctions, by theoretically examining the coevolution of the host exploitation by symbionts and host dependence on symbiosis. Our model reveals that mutualism is more vulnerable to co-extinction through adaptive evolution (evolutionary double suicide) than parasitism. Additionally, it shows that the risk of evolutionary double suicide rarely promotes the backward evolution to an autonomous (non-symbiotic) state. Our results provide a new perspective on the evolutionary fragility of mutualism and the rarity of observed evolutionary transitions from mutualism to parasitism.     

The ecological and medical aspects of the symbiosis between Escherichia coli and man

In this work different variants of the symbiosis of E. coli with a human body are analyzed, and the symbiotic relationships between them are shown to follow the type mutualism, commensalism, parasitism and habitation. The authors emphasize that the multiplicity of variants of bacteria-host relationships is based on the phenotypic polymorphism of E. coli clones (clone lines). Taking into account their ecological (symbiotic) features and biomedical importance, all E. coli clones are divided into 4 groups (clusters): mutualists as nonpathogenic organisms; commensals as potential pathogens (causing extraintestinal E. coli infections); parasites as real pathogens (causing acute intestinal infections); "occasional" symbionts of man. The proposition on the cluster structure of E. coli as a species is formulated.     

Biodiversity and evolution in host-parasite associations

Evolutionary relationships in heterospecific associations (parasitoidism, parasitism, commensalism and mutualism) are analysed through a game theory model defined in terms of fitness of hosts and parasites. In front of the game solutions (i.e. ESS) which present a great diversity of evolutionary patterns, we envisage co-evolution between hosts and parasites through the evolution of its two fundamental parameters (i.e. host's resistance and parasite's virulence). We then discuss the reciprocal influence of hosts and parasites on their respective biodiversity.     

Evolution of mutualistic symbiosis: A differential equation model

In geological history, rapid speciation, called adaptive radiation, has occurred repeatedly. The origins of such newly developing taxa often evolved from the symbiosis of different species. Mutualistic symbioses are generally considered to evolve from parasitic relationships. As well as the previous model of host population with discrete generations, a differential equation model of host population with overlapping generations shows that vertical transmission, defined as the direct transfer of infection from a parent host to its progeny, is an important factor which can stimulate reduction of parasite virulence. Evolution of the vertical transmission rate from both points of view, the parasite and the host, is analyzed. There is a critical level of the rate, below which an evolutionary conflict arises (the parasite would want an increase in the rate while the host would not), and above which both species would correspond to increase the rate. Therefore, once the parasite dominates the evolutionary race so as to overcome this critical level, one-way evolution begins toward a highly mutualistic relationship with a high vertical transmission rate, possibly creating a new organism through symbiosis with perfect vertical transmission. Changes in other parameters may decrease the critical level, initiating one-way evolution. However, changes in traits, probably developed through a long interrelationship in parasitism, do not necessarily induce the evolution of mutualism. Establishment of the ability to make use of metabolic and digestive wastes from the partner certainly facilitates the evolution of mutualism, while improvements in reproductive efficiency of parasites and reduction of negative effects from exploitation in hosts on the contrary disturb mutualism.     

Ecological correlates of feather mite prevalence in passerines

The relationship between host ecology and feather mite prevalence was analysed in birds. Feather mites are small arthropods (fam. Pterolichoidea and Analgoidea) commonly found on birds, although the nature of their interactions with the host (commensalism, mutualism or parasitism), still remains unclear. Host body mass and migratory behaviour were unrelated to feather mite prevalence. Contrary to expectation, there was no differences in mite prevalence between colonial and solitary-breeding species. However, winter sociality was associated with increased prevalence, suggesting that winter and breeding sociality affected the distribution patterns of feather mites in different ways. Plumage dichromatism was negatively correlated with feather mite prevalence, a result that is opposite to that predicted by the Hamilton and Zuk hypothesis for the evolution of host secondary sexual characteristics in relation to parasitism.     

Parasitoids, polydnaviruses and endosymbiosis

Symbiotic associations traditionally have been treated as evolutionary curios rather than as a major source of evolutionary innovation. Recent research on a wide variety of organisms is changing this view and is breaking down the barriers between the traditional categories of parasitism, commensalism and mutualism, to produce a more flexible view of multispecific interactions. An especially abundant, but little discussed, mutualism exists between parasitoid wasps in the superfamily Ichneumonoidea and a novel form of DNA viruses known as polydnaviruses. Mutualisms between viruses and eukaryotes are not often reported, although as many as 100 000 species of organisms may exhibit this unusual association. In this review Jim Whitfield considers what is known about the parasitoid-polydnavirus relationship and how (and from what) it might have arisen.     

Stability analysis of commensal and mutual relations with competitive assimilation in continuous mixed culture

The population dynamics of continuous mixed cultures with pure commensalism, commensalism plus competitive assimilation, pure mutualism, and mutualism plus competitive assimilation was disused. The population does not display oscillatory phenomena where there is a single interaction of commensalism or mutualism. Damped oscillations take place when two interactions (such commensalism and competitive assimilation or mutualism and competitive assimilation) coexist in the continuous mixed culture. The stability of these systems was discussed in detail.     

Cyanobacterial lichen symbiosis: the fungal partner as an optimal harvester

Lichen symbiosis has been traditionally treated as a model case of mutualism in which both partners, the fungus and the photobiont, gain benefits reciprocally. Some recent evidence, however, supports an alternative view that lichen symbiosis may represent an association largely controlled by the commensal or even parasitic fungal partner. The latter gains photosynthates from the photobiont (algae and/or cyanobacteria) which may not always substantially benefit from the symbiosis. We analyze from this perspective how a lichen fungus may maximize photosynthetic gains in bipartite and tripartite associations. We treat the frequency of nitrogen-fixing cells called heterocysts in cyanobacteria and the relative proportion of green algal cells vs. that of cyanobacteria per unit fungus as the variables to be manipulated for maximal carbon gain. The model predicts that even with a negligible cost of cephalodia (compartments containing cyanobacteria) it is in the interest of the tripartite lichen, first, to increase the heterocyst frequency, and second, keep the relative number of cyanobacteria considerably lower than that of green algae. Hence, the lichen fungus achieves higher fitness by making the cyanobacterial partner to specialize on N fixation. The available empirical data support these predictions as the reported heterocyst frequencies in bipartite lichens range from 2 to 8%, and in tripartite lichens between 10 and 55%. It is concluded that interaction asymmetry (i.e. commensalism or parasitism rather than mutualism) provides a sound basis to understand the high phenotypic plasticity expressed by fungi-forming bipartite and tripartite associations with cyanobacteria and green algae.     

Ecological genomics of mutualism decline in nitrogen-fixing bacteria

Anthropogenic changes can influence mutualism evolution; however, the genomic regions underpinning mutualism that are most affected by environmental change are generally unknown, even in well-studied model mutualisms like the interaction between legumes and their nitrogen (N)-fixing rhizobia. Such genomic information can shed light on the agents and targets of selection maintaining cooperation in nature. We recently demonstrated that N-fertilization has caused an evolutionary decline in mutualistic partner quality in the rhizobia that form symbiosis with clover. Here, population genomic analyses of N-fertilized versus control rhizobium populations indicate that evolutionary differentiation at a key symbiosis gene region on the symbiotic plasmid (pSym) contributes to partner quality decline. Moreover, patterns of genetic variation at selected loci were consistent with recent positive selection within N-fertilized environments, suggesting that N-rich environments might select for less beneficial rhizobia. By studying the molecular population genomics of a natural bacterial population within a long-term ecological field experiment, we find that: (i) the N environment is indeed a potent selective force mediating mutualism evolution in this symbiosis, (ii) natural variation in rhizobium partner quality is mediated in part by key symbiosis genes on the symbiotic plasmid, and (iii) differentiation at selected genes occurred in the context of otherwise recombining genomes, resembling eukaryotic models of adaptation.     

The fine line between mutualism and parasitism: complex effects in a cleaning symbiosis demonstrated by multiple field experiments

Ecological theory and observational evidence suggest that symbiotic interactions such as cleaning symbioses can shift from mutualism to parasitism. However, field experimental evidence documenting these shifts has never been reported for a cleaning symbiosis. Here, we demonstrate shifts in a freshwater cleaning symbiosis in a system involving crayfish and branchiobdellid annelids. Branchiobdellids have been shown to benefit their hosts under some conditions by cleaning material from host crayfish's gill filaments. The system is uniquely suited as an experimental model for symbiosis due to ease of manipulation and ubiquity of the organisms. In three field experiments, we manipulated densities of worms on host crayfish and measured host growth in field enclosures. In all cases, the experiments revealed shifts from mutualism to parasitism: host crayfish growth was highest at intermediate densities of branchiobdellid symbionts, while high symbiont densities led to growth that was lower or not significantly different from 0-worm controls. Growth responses were consistent even though the three experiments involved different crayfish and worm species and were performed at different locations. Results also closely conformed to a previous laboratory experiment using the same system. The mechanism for these shifts appears to be that branchiobdellids switched from cleaning host gills at intermediate densities of worms to consuming host gill tissue at high densities. These outcomes clearly demonstrate shifts along a symbiosis continuum with the maximum benefits to the host at intermediate symbiont densities. At high symbiont densities, benefits to the host disappear, and there is some evidence for a weak parasitism. These are the first field experimental results to demonstrate such shifts in a cleaning symbiosis.     

Mutualism and parasitism: the yin and yang of plant symbioses

Plants are solar-powered sugar factories that feed a multitude of other organisms. Many of these organisms associate directly with host plants to gain access to the plant's photosynthates. Such symbioses encompass a wide collection of styles ranging from mutualistic to commensal and parasitic. Among these, the mutualistic arbuscular mycorrhizal (AM) symbiosis is one of the evolutionarily oldest symbioses of plants, relying on the formation of an intimate relationship between fungi of the Glomeromycota and roots of the majority of vascular flowering plants. In this symbiosis, the fungus intracellularly colonizes living root cells, implying the existence of an extreme form of compatibility. Interestingly, molecular events that happen in the plant in response to mycorrhizal colonization also occur in other beneficial and, as recently shown, even antagonistic plant symbioses. Thus, basic 'compatibility modules' appear to be partially conserved between mutualism and parasitism.     

Lotus hosts delimit the mutualism–parasitism continuum of Bradyrhizobium

Symbioses are modelled as evolutionarily and ecologically variable with fitness outcomes for hosts shifting on a continuum from mutualism to parasitism. In a classic example, rhizobia fix atmospheric nitrogen for legume hosts in exchange for photosynthetic carbon. Rhizobial infection often enhances legume growth, but hosts also incur interaction costs because of root tissues and or metabolites needed to support symbionts in planta. Rhizobia exhibit genetic variation in symbiotic effectiveness, and ecological changes in light or mineral nitrogen availability can also alter the benefits of rhizobial infection for hosts. The net effects of symbiosis thus can range from mutualistic to parasitic in a context‐dependent manner. We tested the extent of the mutualism–parasitism continuum in the legume–rhizobium symbiosis and the degree to which host investment can shape its limits. We infected Lotus strigosus with sympatric Bradyrhizobium genotypes that vary in symbiotic effectiveness. Inoculations occurred under different mineral nitrogen and light regimes spanning ecologically relevant ranges. Net growth benefits of Bradyrhizobium infection varied for Lotus and were reduced or eliminated dependent on Bradyrhizobium genotype, mineral nitrogen and light availability. But we did not detect parasitism. Lotus proportionally reduced investment in Bradyrhizobium as net benefit from infection decreased. Lotus control occurred primarily after infection, via fine‐scale modulation of nodule growth, as opposed to control over initial nodulation. Our results show how divestment of symbiosis by Lotus can prevent shifts to parasitism.     

阿米巴-细菌互作：进化、生态与环境效应

阿米巴是原生动物的主要类群,是陆地和水生生态系统的关键组成部分。阿米巴与细菌之间有着密切而复杂的相互作用。一方面,阿米巴通过捕食直接影响细菌群落与多样性,增强细菌活性。另一方面,细菌也进化出抵抗捕食的机制来抵抗甚至感染阿米巴,反向影响阿米巴的生长和多样性。近年来,阿米巴-细菌互作的研究开始受到了广泛关注。本文总结了阿米巴-细菌互作的进化历史、生态关系(捕食、偏利共生、寄生和互利共生)以及其对环境的潜在影响,旨在更好地理解阿米巴-细菌互作这一研究领域,为其他原生动物-微生物之间的研究提供新的思路,也为探究宿主-微生物互作的机制提供参考。     

Cooperation strategies, signals and symbiosis

The authors of models concerning cooperation found an interesting application point in mutualism and symbiosis. Phenomena which are to be found in symbiosis are integrated by the various models in relation to the 'Prisoner's Dilemma': 'cost-benefit' ratio components, signals, neighbourhood interaction or reciprocity. Reciprocal altruism is found in symbiosis, but others strategies seem more to reflect human societies, where choice is unrestricted. It is very difficult to give a value to cost and benefit equilibriums. The biological environment interferes and a model based on a dominant strategy is often limited or insufficient to report on the living system's evolution.     

Parabiotic ants: the costs and benefits of symbiosis

1. Mutualisms are important drivers of co‐evolution and speciation. However, they typically imply costs for one or both partners. Each partner consequently tries to maximise benefits and minimise costs. Mutualisms can therefore develop towards commensalism or parasitism if one partner fails to provide sufficient benefits. This is particularly likely in diffuse interactions, where multiple species can associate with each other. If costs and benefits of a species vary with the identity of the partner species, this may result in a geographical mosaic of co‐evolution. 2. In the present study, inter‐specific interactions in two parabiotic associations of ants were studied (Hymenoptera: Formicidae). One Crematogaster species was associated with one of two closely related Camponotus species. We assessed cost and benefits by studying behavioural interactions, foraging behaviour, and nest defence in the associations. 3. While parabioses had been shown to be mutualistic, evidence was found for exploitation and aggressive competition between species. In spite of apparent costs of being exploited, we found no benefits for one partner (Crematogaster). The magnitude of potential costs to Crematogaster varied between the two Camponotus species. 4. We conclude that the cost/benefit ratio for Crematogaster varies between the two Camponotus partners, and between environmental conditions. Parabiosis can thus fluctuate between mutualism, commensalism, and parasitism, with Crematogaster being the species that may have higher costs than benefits. 5. We suggest that geneflow in the Crematogaster population hinders local adaptation to the resulting mosaic of locally varying selection pressures. This study demonstrates how diffuse interactions and environmental variation can result in a complex of local selection pressures.     

A simple model of host-parasite evolutionary relationships. Parasitism: compromise or conflict?

The evolutionary biology of host-parasite relationships are considered here using a simple game-theory model in which hosts play against parasite and vice versa. In this model, the players can choose between two strategies (aggressive or not aggressive) and the utility of the game is envisaged in terms of fitness and selective costs. The game solutions suggest that the two types of confrontation are encountered in symbiotic relationships and thus constitute two Evolutionary Stable Strategies (ESS). These observations lead us to discuss: (i) the status of different kinds of symbiotic relationships (i.e. parasitoidism; parasitism, commensalism and mutualism) related to selective costs and (ii) the position of coevolution in this game theory context.     

Boundary lines in symbiosis forms

Symbiosis can take different forms (parasitism, mutualism, commensalism, etc.) but boundaries between different types of symbiotic interactions are not well defined. The kinds of symbiotic associations between organisms cannot however be restricted to isolated and distinct categories. These associations are part of a broad continuum in which it is difficult to know where one type of association ends and another begins. Moreover, different scientists use the same term to mean different things or different terms to mean the same thing. This can obscure what is biologically important and what is not. This communication proposes a new classification scheme, which simply and comprehensively illustrates relationships between the various kinds of associations. The scheme illustrates relationships clearly and highlights the continuum between types of associations. It further indicates where modifications to the scheme are possible over time. The classification of the association between two organisms can be reduced to two factors: 1) the impact incurred by the host (benefit or damage) and 2) the relative duration of the association (RDA), i.e. the ratio of the duration of the association to the life expectancy of the symbiont. The conceptual figure provides concrete examples and illustrates some relationships that can change during different life stages. This figure should help teachers and students in the understanding of symbiosis, and could be a starting point for future discussions in the continuously developing research fields studying ecological and evolutionary implications of symbiotic relationships.