Biological markets: supply and demand determine the effect of partner choice in cooperation,mutualism and mating

The formation of collaborating pairs by individuals belonging to two different classes occurs in the contexts of reproduction and intea-specific cooperation as well as of inter-specific mutualism. There is potential for partner choice and for competition for access to preferred partners in all three contexts. These selective forces have long been recognised as important in sexual selection, but their impact is not yet appreciated in cooperative and mutualistic systems. The formation of partnerships between members of different classes has much in common with the conclusion of trade agreements in human markets with two classes of traders, like producers and consumers, or employers and employees. Similar game-theoretical models can be used to predict the behaviour of rational traders in human markets and the evolutionarily stable strategies used in biological markets. We present a formal model in which the influence of the market mechanism on selection is made explicit. We restrict ourselves to biological markets in which: (1) Individuals do not compete over access to partners in an agonistic manner, but rather by outcompeting each other in those aspects that are preferred by the choosing party. (2) The commodity the partner has to offer cannot be obtained by the use of force, but requires the consent of the partner. These two restrictions ensure a dominant role for partner choice in the formation of partnerships. In a biological market model the decision to cooperate is based on the comparison between the offers of several potential partners, rather than on the behaviour of a single potential partner, as is implicitly assumed in currently accepted models of cooperation. In our example the members of one class A offer a commodity of fixed value in exchange for a commodity of variable value supplied by the other class, B. We show that when the B-class outnumbers the A-class sufficiently and the cost for the A-class to sample the offers of the B-class are low, the choosiness of the A-class will lead to selection for the supply of high value commodities by the B-class (Fig. 3a). Under the same market conditions, but with a high sampling cost this may still be the evolutionariy stable outcome, but another pair of strategies proves to be stable too: relaxed choosiness of class A coupled with low value commodities supplied by class B (Fig. 3b). We give a number of examples of mating, cooperative and mutualistic markets that resemble the low sampling cost situation depicted in Fig. 3a.     

Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Floral scents: their roles in nursery pollination mutualisms

Mutualisms are interspecies interactions in which each participant gains net benefits from interacting with its partner. In nursery pollination mutualisms, pollinators reproduce within the inflorescence they pollinate. In these systems, each partner depends directly on the other for its reproduction. Therefore, the signal responsible for partner encounter is crucial in these horizontally transmitted mutualisms, in which the association between specific partners must be renewed at each generation. As in many other interspecies interactions, chemical signals are suspected to be important in the functioning of these mutualisms. We synthesized and compared the published data available on the role of floral scents in the functioning of the 16 known independently evolved nursery pollination mutualisms. So far, attraction of pollinators to their specific hosts has been investigated in only seven of these systems, and the majority of the studies have been conducted on one of them, fig/fig wasp interactions. While such unevenness of the information limits the potential for meta-analysis, some patterns emerge from this review concerning the role of flower volatiles in maintaining the specificity of pollinator attraction, in signaling the appropriate phenological stage for pollinator visit, in attracting the pollinator toward the rewardless sex in dioecious plant species and in aiding the location and exploitation of resources by parasites and predators associated with these mutualisms. Finally, we highlight new perspectives on the evolution of signals in these diversified systems depending on the age and the degree of specificity of the interaction, and on the effect of phylogenetic inertia on the evolutionary dynamics of plant signals.     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Plant-mediated and nonadditive effects of two global change drivers on an insect herbivore community

Warmer temperatures can alter the phenology and distribution of individual species. However, differences across species may blur community-level phenological responses to climate or cause biotic homogenization by consistently favoring certain taxa. Additionally, the response of insect communities to climate will be subject to plant-mediated effects, which may or may not overshadow the direct effect of rising temperatures on insects. Finally, recent evidence for the importance of interaction effects between global change drivers suggests that phenological responses of communities to climate may be altered by other drivers. We used a natural temperature gradient (generated by elevation and topology), combined with experimental nitrogen fertilization, to investigate the effects of elevated temperature and globally increasing anthropogenic nitrogen deposition on the structure and phenology of a seminatural grassland herbivore assemblage (lepidopteran insects). We found that both drivers, alone and in combination, severely altered how the relative abundance and composition of species changed through time. Importantly, warmer temperatures were associated with biotic homogenization, such that herbivore assemblages in the warmest plots had more similar species composition than those in intermediate or cool plots. Changes in herbivore composition and abundance were largely mediated by changes in the plant community, with increased nonnative grass cover under high treatment levels being the strongest determinant of herbivore abundance. In addition to compositional changes, total herbivore biomass more than doubled under elevated nitrogen and increased more than fourfold with temperature, bearing important functional implications for herbivores as consumers and as a prey resource. The crucial role of nonnative plant dominance in mediating responses of herbivores to change, combined with the frequent nonadditive (positive and negative) effects of the two drivers, and the differential responses of species, highlight that understanding complex ecosystem responses will benefit from multifactor, multitrophic experiments at community scales or larger.     

Mutualistic stability in the arbuscular mycorrhizal symbiosis: exploring hypotheses of evolutionary cooperation

The 450-million-year-old symbiosis between the majority of land plants and arbuscular mycorrhizal fungi (AMF) is one of the most ancient, abundant, and ecologically important mutualisms on Earth. Yet, the evolutionary stability of mycorrhizal associations is still poorly understood, as it follows none of the constraints thought to stabilize cooperation in other well-known mutualisms. The capacity of both host and symbiont to simultaneously interact with several partners introduces a unique dilemma; detecting and punishing those exploiting the mutualism becomes increasingly difficult if these individuals can continue to access resources from alternative sources. Here, we explore four hypotheses to explain evolutionary cooperation in the arbuscular mycorrhizal symbiosis: (1) pseudo-vertical transmission and spatial structuring of plant and fungal populations leading to local adaptation of partners; (2) luxury resource exchange in which plants trade surplus carbon for excess fungal nutrients; (3) partner choice allowing partners to associate with better cooperators; and (4) host and symbiont sanctions which actively reward good partners and punish less cooperative ones. We propose that mycorrhizal cooperation is promoted by an exchange of surplus resources between partners and enforced through sanctions by one or both partners. These mechanisms may allow plant and fungal genotypes to discriminate against individuals employing exploitative strategies, promoting patterns of partner choice. Together these selection pressures provide a framework for understanding the stabilization of mycorrhizal cooperation over evolutionary time.     

Bottom-up effects of plant genotype on aphids, ants, and predators

Theory predicts that bottom-up ecological forces can affect community dynamics, but whether this extends to the effects of heritable plant variation on tritrophic communities is poorly understood. In a field experiment, I contrasted the effects of plant genotype (28 genotypes; 1064 plants), aphid density, and the presence/absence of mutualistic ants in affecting the per capita population growth of a specialist aphid herbivore, as well as the effects of plant genotype on the third trophic level. Plant genotype strongly affected aphid population growth rate, explaining 29% of the total variation in growth rate, whereas aphid density and ant-aphid interactions explained substantially less variation (< 2%) in aphid population growth rate. Plant genotype also had direct and indirect effects on the third trophic level, affecting the abundance of aphid-tending ants and the richness of predators. Multiple regression identified several heritable plant traits that explained 49% of the variation in aphid growth rate and 30% of the variation in ant abundance among plant genotypes. These bottom-up effects of plant genotype on tritrophic interactions were independent of the effects of either initial aphid density or the presence/absence of mutualistic ants. This study shows that plant genotype can be one of the most important ecological factors shaping tritrophic communities.     

A century of increasing pine density and associated shifts in understory plant strategies

We analyzed one of the longest-term ecological data sets to evaluate how forest overstory structure is related to herbaceous understory plant strategies in a ponderosa pine forest. Eighty-two permanent 1-m2 chart quadrats that were established as early as 1912 were remeasured in 2007. We reconstructed historical forest structure using dendrochronological techniques. Ponderosa pine basal area increased from an average of 4 m2/ha in the early 1900s to 29 m2/ha in 2007. Understory plant foliar cover declined by 21%, species richness declined by two species per square meter, and functional diversity also declined. The relative cover of C4 graminoids decreased by 18% and C3 graminoids increased by 19%. Herbaceous plant species with low leaf and fine root nitrogen concentrations, low specific leaf area, high leaf dry matter content, large seed mass, low specific root length, short maximum height, and early flowering date increased in relative abundance in sites where pine basal area increased the most. Overall, we observed a long-term shift in composition toward more conservative shade- and stress-tolerant herbaceous species. Our analysis of temporal changes in plant strategies provides a general framework for evaluating compositional and functional changes in terrestrial plant communities.     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Tradable fuel economy credits: Competition and oligopoly

Corporate average fuel economy (CAFE) regulations specify minimum standards for fuel efficiency that vehicle manufacturers must meet independently. We design a system of tradeable fuel economy credits that allows trading across vehicle classes and manufacturers with and without considering market power in the credit market. We perform numerical simulations to measure the potential cost savings from moving from the current CAFE system to one with stricter standards, but that allows vehicle manufacturers various levels of increased flexibility. We find that the ability for each manufacturer to average credits between its cars and trucks provides a large percentage of the potential savings. As expected, the greatest savings come from the greatest flexibility in the credit system. Market power lowers the potential cost savings to the industry as a whole, but only modestly. Loss in efficiency from market power does not eliminate the gains from credit trading.     

The roles of sensory traps in the origin, maintenance, and breakdown of mutualism

Sensory traps are signal mimics that elicit out-of-context behaviors by exploiting the adaptive, neural responses of signal receivers. Sensory traps have long been invoked in studies of mate and prey attraction, but the possible roles of sensory traps in mutualisms (cooperation between species) have yet to be thoroughly examined. Our review identifies four candidate roles for sensory traps in the evolution of mutualistic interactions: reassembly, error reduction, enforcement, and cost reduction. A key consequence of sensory traps is that they limit the applicability of partner choice and biological market models of mutualism. We conclude by suggesting that an important research topic in the evolution of cooperation should be to identify any mechanisms that increase the truthfulness of communication between cooperating species.     

Sperm exchange in a simultaneous hermaphrodite

Many simultaneously hermaphroditic animals cross-inseminate when copulating, even though unilateral sperm exchange is functionally possible. The question is why reciprocity is the rule. This study addresses sperm exchange in a hermaphroditic planarian flatworm, Dugesia polychroa, in which self-fertilisation does not occur. We investigated the availability of self sperm for inseminating a partner, how self sperm affects the likelihood of sperm donation, and whether sperm donation depends on reciprocation by the partner. The amount of self sperm depended on body size, duration of isolation and experimental date, and animals that had more self sperm were more likely to mate. Depletion of allosperm (sperm received from partners) played only an indirect role, suggesting that animals copulate more to donate sperm rather than to replenish allosperm reserves. Among 60 copulating pairs, reciprocal insemination was more common (63.3%) than unilateral transfer (18.3%). A surprising 18.3% did not exchange sperm in either direction. Individuals were more likely to behave like their partners, resulting in more symmetrical matings (either reciprocity or no sperm exchange) than asymmetrical matings (unilateral sperm donation). The amount of self sperm donated during a copulation depended mainly on the amount available before the copulation and not on the amount received from the partner. The results suggest that conditional sperm exchange, or sperm trading, takes place in D. polychroa. Received: 31 May 1997 / Accepted after revision: 1 December 1997     

A review of hypotheses for the functions of avian duetting

Avian duets are striking for the remarkable precision with which duetting partners sometimes coordinate their songs. Duetting species are taxonomically diverse, and the form of their duets varies. The reasons some birds duet when most do not remains unclear despite numerous hypotheses for its function. I review work done so far on duetting, discuss evidence for and against hypotheses for its functions, and highlight approaches useful for future research. The four hypotheses that appear most promising are that individuals join their partners songs to form duets: (1) to avoid being usurped from a partnership, (2) to prevent their partner being usurped, (3) as a collaborative display in defence of some resource, or (4) to signal commitment to their partner. These hypotheses are not mutually exclusive, and duetting is likely to have multiple roles both within and among species. However, much basic research is still required. Characteristics of duets have rarely been quantified in detail, and information about variability among species in the precision of duetting is necessary, not only to test hypotheses about function, but also to define duetting more precisely. Quantifying the relative frequencies of alternative vocal strategies (for example, remaining silent when a partner sings versus joining in to form a duet) between species and in different contexts will help to determine why partners coordinate their songs to form duets. Furthermore, social systems and sex roles in duetting species are poorly understood, yet understanding these is critical to determining the functions of avian duetting.Communicated by A. Cockburn     

A simple population theory for mutualism by the use of lattice gas model

The population dynamics of species interactions provides valuable information for life sciences. Lotka-Volterra equations (LVEs) are known to be the most popular model, and they are mainly applied to the systems of predation and competition. However, LVEs often fail to catch the population dynamics of mutualism; the population sizes of species increase infinitely under certain condition (divergence problem). Furthermore, LVEs never predicts the Allee effect in the systems of obligate mutualism. Instead of LVEs, several models have been presented for mutualism; unfortunately, they are rather complicated. It is, therefore, necessary to introduce a simpler theory for mutualism. In the present paper, we apply the lattice gas model which corresponds to the mean-field theory of the usual lattice model. The derived equations are cubic and contain only essential features for mutualism. In the case of obligate mutualism, the dynamics exhibits the Allee effect, and it is almost the same as in the male-female systems. In our model, the population sizes never increase infinitely, because our model contains not only intra- but also interspecific competitions. If the density of one species increases disproportionately in respect of its mutual partners, then this might imply downward pressure on the population abundance of the mutual partner species and such feedback would eventually act as a controlling influence on the population abundance of either species. We discuss several assumptions in our model; in particular, if both species can occupy in each cell simultaneously, then the interspecific competition disappears.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

Effects of long-term nitrogen addition and precipitation changes on CH4 flux in an alpine steppe北大核心CSCD

土壤是甲烷(CH4)重要的源和汇.氮沉降和降水格局变化正在急剧改变土壤碳循环,进而可能对土壤CH4通量造成深刻影响.高寒生态系统是巨大的碳库,对氮沉降和降水变化十分敏感.然而,目前多数研究集中在短期实验上,缺乏对长期氮沉降和降水变化背景下CH4通量的响应及其调控因素的认识.以青藏高原高寒草原为研究对象,在2013年搭建模拟氮沉降和降水格局改变实验平台.基于静态箱–气相色谱法测定2020年生长季(5-10月)土壤CH4通量.结果显示,高寒草原土壤呈CH4的汇.氮添加没有显著改变生长季和植物生长高峰CH4通量.然而,降水变化显著改变了生长季和植物生长高峰CH4通量,其中降水增加(+50%降水)降低了CH4的吸收(分别为–16%和–45%),降水减少(–50%降水)增强了CH4的吸收(分别为+73%和+33%).进一步研究发现,与植物属性和功能基因丰度相比,土壤环境因子主导了CH4通量变化(解释率>90%).其中CH4通量与土壤含水量和温度显著正相关,与土壤pH显著负相关.综上所述,在未来全球变化情景下,降水格局改变更能调节青藏高原高寒草原CH4通量的变化.(图6表1参37)     

Compositional analysis of overdispersed counts using generalized estimating equations

Multivariate abundance data are commonly collected in ecology, and used to explore questions of “community composition”—how relative abundance of different taxa changes with environmental conditions. In this paper, we propose a log-linear marginal modeling approach for analyzing such compositional count data, via generalized estimating equations. This method exploits the multiplicative nature of log-linear models for counts, by reparameterizing models that describe marginal effects on mean abundance. This allows partitioning into “main effects” and compositional effects, which is appealing for interpretation. We apply the proposed approach to reanalyze compositional counts of benthic invertebrates from Delaware Bay, and data of invertebrate communities inhabiting Acacia plants in eastern Australia. In both cases we resort to a resampling approach to make inferences about regression parameters, because the number of clusters was not large compared to cluster size.     

Uncertainty and the choice of pollution control instruments

This paper compares the relative efficiencies of pollution taxes, pollution standards, and the auctioning of pollution rights when the marginal damage function or marginal control cost are subject to uncertainty. In the first case, we find that all instruments yield the same expected social surplus. In the latter case, the choice of the optimal instrument depends, in general, on the relative elasticities of the marginal damage and marginal expected cost functions, on the way in which uncertainty enters the model, and on the distribution of the error term. Policy conclusions are derived.     

A framework for allocating conservation resources among multiple threats and actions

Land managers decide how to allocate resources among multiple threats that can be addressed through multiple possible actions. Additionally, these actions vary in feasibility, effectiveness, and cost. We sought to provide a way to optimize resource allocation to address multiple threats when multiple management options are available, including mutually exclusive options. Formulating the decision as a combinatorial optimization problem, our framework takes as inputs the expected impact and cost of each threat for each action (including do nothing) and for each overall budget identifies the optimal action to take for each threat. We compared the optimal solution to an easy to calculate greedy algorithm approximation and a variety of plausible ranking schemes. We applied the framework to management of multiple introduced plant species in Australian alpine areas. We developed a model of invasion to predict the expected impact in 50 years for each species-action combination that accounted for each species’ current invasion state (absent, localized, widespread); arrival probability; spread rate; impact, if present, of each species; and management effectiveness of each species-action combination. We found that the recommended action for a threat changed with budget; there was no single optimal management action for each species; and considering more than one candidate action can substantially increase the management plan's overall efficiency. The approximate solution (solution ranked by marginal cost-effectiveness) performed well when the budget matched the cost of the prioritized actions, indicating that this approach would be effective if the budget was set as part of the prioritization process. The ranking schemes varied in performance, and achieving a close to optimal solution was not guaranteed. Global sensitivity analysis revealed a threat's expected impact and, to a lesser extent, management effectiveness were the most influential parameters, emphasizing the need to focus research and monitoring efforts on their quantification.