Priority Effects Research Articles

Effects of Phenology on Plant Community Assembly and Structure

Phenology—the timing of critical stages of growth and reproduction and the transitions between them—determines environmental conditions and biotic interactions. Hence, phenology is a key functional trait influencing organisms’ survival and fitness; however, the role of phenology in community assembly processes has been less considered. Here we review the importance of phenology in environmental and biotic filtering, structuring priority effects, and species coexistence in the context of the assembly of native communities, as well as in invasions and restoration. We highlight the complexity of the life-history aspect of phenology, which makes simple trade-offs—such as between growth timing and competitive ability—part of larger plant strategies shaped by a framework of risk, reward, and investment over multiple timescales. Embracing this complexity could yield insights into how phenology shapes communities.

Transcriptional Responses to Priority Effects in Nectar Yeast.

Priority effects, where the order and timing of species arrival influence the assembly of ecological communities, have been observed in a variety of taxa and habitats. However, the genetic and molecular basis of priority effects remains unclear, hindering a better understanding of when priority effects will be strong. We sought to gain such an understanding for the nectar yeast Metschnikowia reukaufii commonly found in the nectar of our study plant, the hummingbird-pollinated Diplacus (Mimulus) aurantiacus. In this plant, M. reukaufii can experience strong priority effects when it reaches flowers after other nectar yeasts, such as M. rancensis. After inoculation into two contrasting types of synthetic nectar simulating early arrival of M. rancensis, we conducted whole-transcriptome sequencing of 108 strains of M. reukaufii. We found that several genes were differentially expressed in M. reukaufii strains when the nectar had been conditioned by growth of M. rancensis. Many of these genes were associated with amino acid metabolism, suggesting that M. reukaufii strains responded molecularly to the reduction in amino acid availability caused by M. rancensis. Furthermore, investigation of expression quantitative trait loci (eQTLs) revealed that genes involved in amino acid transport and resistance to antifungal compounds were enriched in some genetic variants of M. reukaufii. We also found that gene expression was associated with population growth rate, particularly when amino acids were limited. These results suggest that intraspecific genetic variation in the ability of nectar yeasts to respond to nutrient limitation and direct fungal competition underpins priority effects in this microbial system.

The Synergistic Effects of Different Phosphorus Sources: Ferralsols Promoted Soil Phosphorus Transformation and Accumulation

Phosphorus (P) application can enhance soil P availability and alter P fractions. However, the P accumulation and transformation of different P sources in low-phosphorus red soil remain unclear. Two-year (2018–2019) field experiments were conducted to investigate the effects of five P source treatments (CK—no phosphorus; SSP—superphosphate; MAP—calcium–magnesium phosphate; DAP—monoammonium phosphate; and CMP—diammonium phosphate) on the P accumulation of maize and soil P fractions in low-P red soil using the Hedley Sequential Method. The results showed that P application significantly increased P uptake, Olsen-P, total phosphorus, and most of the soil P fractions. Compared to the CMP, MAP, and DAP treatments, SSP had a relatively higher P accumulation and labile P pool, with a slightly lower moderately labile P pool. The SSP treatment mainly increased soil-available P content and crop P uptake by increasing the labile P pool (resin-P and NaHCO3-Pi) and reducing the moderately labile P pool and non-labile P pool. The P activation coefficient (PAC%) and Olsen-P were positively correlated with labile P (resin-P, NaHCO3-Pi, and NaHCO3-Po) and moderately labile P (NaOH-Pi and 1 M HCl-Pi) and negatively correlated with Fe2O3 and Al2O3. The results suggest that SSP has a priority effect on the crop P uptake and soil P availability in low-P red soil.

Effects of parental care on skin microbial community composition in poison frogs.

Parent-offspring interactions constitute the first contact of many newborns with their environment, priming community assembly of microbes through priority effects. Early exposure to microbes can have lasting influences on the assembly and functionality of the host's microbiota, leaving a life-long imprint on host health and disease. Studies of the role played by parental care in microbial acquisition have primarily focused on humans and hosts with agricultural relevance. Anuran vertebrates offer the opportunity to examine microbial community composition across life stages as a function of parental investment. In this study, we investigate vertical transmission of microbiota during parental care in a poison frog (Family Dendrobatidae), where fathers transport their offspring piggyback-style from terrestrial clutches to aquatic nurseries. We found that substantial bacterial colonization of the embryo begins after hatching from the vitelline envelope, emphasizing its potential role as microbial barrier during early development. Using a laboratory cross-foster experiment, we demonstrated that poison frogs performing tadpole transport serve as a source of skin microbes for tadpoles on their back. To study how transport impacts the microbial skin communities of tadpoles in an ecologically relevant setting, we sampled frogs and tadpoles of sympatric species that do or do not exhibit tadpole transport in their natural habitat. We found more diverse microbial communities associated with tadpoles of transporting species compared to a non-transporting frog. However, we detected no difference in the degree of similarity between adult and tadpole skin microbiotas, based on whether the frog species exhibits transporting behavior or not. Using a field experiment, we confirmed that tadpole transport can result in the persistent colonization of tadpoles by isolated microbial taxa associated with the caregiver's skin, albeit often at low prevalence. This is the first study to describe vertical transmission of skin microbes in anuran amphibians, showing that offspring transport may serve as a mechanism for transmission of parental skin microbes. Overall, these findings provide a foundation for further research on how vertical transmission in this order impacts host-associated microbiota and physiology.

New insights into the merging behavior of pedestrians: Quantification of lane competition level

Merging behavior is a complex crowd movement pattern caused by certain architectural features and has triggered serious crowd accidents worldwide. It is not uncommon to find such merging structures on the floor plans of some underground infrastructures, which deserve more detailed studies on special crowd movement patterns to enhance the safety level. Researchers have demonstrated that the unstable emotions and competitive behaviors of pedestrians during underground evacuation need to be taken into account due to the enclosed environment. However, there are no relevant studies focused on the competition characteristics among pedestrians during the merging process in a quantitative way. The comprehensiveness and systematization of the geometric features of typical merging structures, as well as the way to create the perception of competition and conflict among pedestrians during real emergencies also need to be improved. In this paper, we carried out a series of controlled experiments in symmetric and asymmetric merging structures with different channel widths and incentive levels to investigate crowd merging behavior with competition characteristics. The density and delay time within the merging area, as well as the time interval and lane changes before and after merging are analyzed. By introducing and modifying Simpson’s Diversity Index, we proposed a new method for quantifying the competition level among pedestrians during the merging process. It is concluded that the performance of the merging area is sensitive to the channel widths and the merging layouts, and the negative effect of unbalanced walking priority is more significant under high incentive level. The lane competition level during the merging process is correlated with the effects of various geometric features, movement layouts, and incentive levels. The method proposed in this paper brings new insights into the interpretation of merging scenarios and more studies on pedestrian dynamics. We hope that this study could serve as supplementary work for other modeling or field experiment methods to jointly provide more insights into the crowd management strategies in some underground infrastructures.

Phenology mediates direct and indirect interactions among co-occurring invasive plant species.

Why nonnative invasive plant species commonly co-occur, despite their competitive superiority and propensity to displace native species, remains a paradox in invasion biology. Negative interactions among competitively dominant invaders are potentially alleviated by two understudied mechanisms: seasonal priority effects, where phenological separation weakens the effect of competition on species with early phenology; and indirect facilitation, where competition between two species is mitigated by a third species. Although phenological separation has been speculated as a mechanism for explaining co-occurrence patterns of invasive plants, it has never been directly tested. In a greenhouse experiment, we tested the effect of phenological separation on direct and indirect interactions between three co-occurring invasive plant species found in the riparian forests of North America. These species have distinct natural phenological separation with reproduction in early spring (Ficaria verna), mid-spring (Alliaria petiolata), and late summer (Microstegium vimineum). When phenology was experimentally synchronized, direct pairwise interactions among invasive species were overwhelmingly negative, asymmetric, and unlikely to promote co-occurrence. However, increasing phenological separation generated seasonal priority effects, which weakened the effect of competition on species with early phenology. Furthermore, the addition of a third species generated indirect facilitative effects, which balanced competitive outcomes among the two weakest competitors. Based on these findings, we conclude that phenological separation modulates the strength of both seasonal priority effects and indirect facilitation within species interaction networks and may promote the co-occurrence of three common invasive species within this study system. We articulate how future studies can test the external validity of these findings in more complex environmental conditions and with a larger range of invasive plants.

Pinewood nematode induced changes in the assembly process of gallery microbiomes benefit its vector beetle's development.

Microbiomes play crucial roles in insect adaptation, especially under stress such as pathogen invasion. Yet, how beneficial microbiomes assemble remains unclear. The wood-boring beetle Monochamus alternatus, a major pest and vector of the pine wilt disease (PWD) nematode, offers a unique model. We conducted controlled experiments using amplicon sequencing (16S rRNA and ITS) within galleries where beetles and microbes interact. PWD significantly altered bacterial and fungal communities, suggesting distinct assembly processes. Deterministic factors like priority effects, host selection, and microbial interactions shaped microbiome composition, distinguishing healthy from PWN-infected galleries. Actinobacteria, Firmicutes, and Ophiostomataceae emerged as potentially beneficial, aiding beetle's development and pathogen resistance. This study unveils how nematode-induced changes in gallery microbiomes influence beetle's development, shedding light on microbiome assembly amid insect-pathogen interactions. Insights gleaned enhance understanding of PWD spread and suggest novel management strategies via microbiome manipulation.IMPORTANCEThis study explores the assembly process of gallery microbiomes associated with a wood-boring beetles, Monochamus alternatus, a vector of the pine wilt disease (PWD). By conducting controlled comparison experiments and employing amplicon approaches, the study reveals significant changes in taxonomic composition and functional adaptation of bacterial and fungal communities induced by PWD. It identifies deterministic processes, including priority effects, host selection, and microbial interactions, as major drivers in microbiome assembly. Additionally, the study highlights the presence of potentially beneficial microbes such as Actinobacteria, Firmicutes, and Ophiostomataceae, which could enhance beetle development and resistance to pathogens. These findings shed light on the intricate interplay among insects, microbiomes, and pathogens, contributing to a deeper understanding of PWD prevalence and suggesting innovative management strategies through microbiome manipulation.

A strong priority effect in the assembly of a specialized insect-microbe symbiosis.

Host-associated microbial communities underpin critical ecosystem processes and human health, and their ability to do so is determined in turn by the various processes that shape their composition. While selection deterministically acts on competing genotypes and species during community assembly, the manner by which selection determines the trajectory of community assembly can differ depending on the sequence by which taxa are established within that community. We document this phenomenon, known as a priority effect, during experimental colonization of a North American insect pest, the squash bug Anasa tristis, by its betaproteobacterial symbionts in the genus Caballeronia. Our study demonstrates how stark, strain-level variation can emerge in specialized host-microbe symbioses simply through differences in the order by which strains colonize the host. Understanding the mechanistic drivers of community structure in host-associated microbiomes can highlight both pitfalls and opportunities for the engineering of these communities and their constituent taxa for societal benefit.

Effects of search priority on working memory-guided search for real objects: Evidence from eye-movements

Effects of search priority on working memory-guided search for real objects: Evidence from eye-movements

Amphibian hotspots in Iran: Effectiveness of current protected area and priorities for conservation planning

The identification of conservation priority areas for amphibians in Iran remains unexplored. Although protected areas play a vital role in biodiversity conservation, 40 percent of amphibians are at risk of extinction. The present study aimed to identify the priority areas for amphibian conservation and assess the effectiveness of the current protected conservation areas. 2192 species occurrence points for 23 amphibians, including 16 species of Anura and 7 species of Caudata, were curated. These occurrence points in addition to bioclimatic, anthropogenic, and geographical variables were used to model the species distribution. Then, the richness and rarity maps of amphibians were determined and hotspots identified based on the top 10 % area of the country with the highest richness and rarity values. Finally, Iran's protected areas were overlaid to evaluate the current coverage of hotspots and identify future conservation priorities. Although approximately 12 % of Iran is currently protected, the present study findings indicate that the current network of protected areas is ineffective for amphibian conservation; over 90 % of hotspots for amphibians lie outside the current national protected area network. The most important hotspots are located in the Caspian Hyrcanian Mixed Forest, the western margin of the North Zagros, and the Central Zagros Mountains Forest-Steppe ecoregions of the country. Among different types of protected areas that overlap with amphibian hotspots, protected areas and wildlife refuges respectively ranked the highest in quantity and size, while national parks ranked the lowest. In the current study, a baseline of top candidate areas for expanding protected areas where habitats can be managed to protect amphibians in Iran was provided. To further improve protected areas' coverage, priority conservation areas in the Zagros Mountains Forest-Steppe hotspots were recommended.

Primary succession of Bifidobacteria drives pathogen resistance in neonatal microbiota assembly

Human microbiota assembly commences at birth, seeded by both maternal and environmental microorganisms. Ecological theory postulates that primary colonizers dictate microbial community assembly outcomes, yet such microbial priority effects in the human gut remain underexplored. Here using longitudinal faecal metagenomics, we characterized neonatal microbiota assembly for a cohort of 1,288 neonates from the UK. We show that the pioneering neonatal gut microbiota can be stratified into one of three distinct community states, each dominated by a single microbial species and influenced by clinical and host factors, such as maternal age, ethnicity and parity. A community state dominated by Enterococcus faecalis displayed stochastic microbiota assembly with persistent high pathogen loads into infancy. In contrast, community states dominated by Bifidobacterium, specifically B. longum and particularly B. breve, exhibited a stable assembly trajectory and long-term pathogen colonization resistance, probably due to strain-specific functional adaptions to a breast milk-rich neonatal diet. Consistent with our human cohort observation, B. breve demonstrated priority effects and conferred pathogen colonization resistance in a germ-free mouse model. Our findings solidify the crucial role of Bifidobacteria as primary colonizers in shaping the microbiota assembly and functions in early life.

Asymmetric Biotic Interactions Cannot Be Inferred Without Accounting for Priority Effects.

Understanding biotic interactions is a crucial goal in community ecology and species distribution modelling, and large strides have been made towards improving multivariate computational methods with the aim of quantifying biotic interactions and improving predictions of species occurrence. Yet, while considerable attention has been given to computational approaches and the interpretation of these quantitative tools, the importance of sampling design to reveal these biotic interactions has received little consideration. This study explores the influential role of priority effects, that is, the order of habitat colonisation, in shaping our ability to detect biotic interactions. Using a simple set of simulations, we demonstrate that commonly used cross-sectional co-occurrence data alone cannot be used to make reliable inferences on asymmetric biotic interactions, even if they perform well in predicting the occurrence of species. We then show how sampling designs that consider priority effects can recover the asymmetric effects that are lost when priority effects are ignored. Based on these findings, we urge for caution when drawing inferences on biotic interactions from cross-sectional binary co-occurrence data, and provide guidance on sampling designs that may provide the necessary data to tackle this longstanding challenge.

Decomposing socioemotional wealth: the effect of restricted and extended socioemotional priorities on family firms’ internationalization

Decomposing socioemotional wealth: the effect of restricted and extended socioemotional priorities on family firms’ internationalization

Patterns of spawning and settlement of reef fishes as strategic responses to post-settlement competition

Settlement is a critical transition in the life history of reef fish, and the timing of this event can have a strong effect on fitness. Key factors that influence settlement timing are predictable lunar cyclic variation in tidal currents, moonlight, and nocturnal predation risk as larvae transition from pelagic to benthic environments. However, populations typically display wide variation in the arrival of settlers over the lunar cycle. This variation is often hypothesized to result from unpredictable conditions in the pelagic environment and bet-hedging by spawning adults. Here, we consider the hypothesis that the timing of spawning and settlement is a strategic response to post-settlement competition. We use a game theoretic model to predict spawning and settlement distributions when fish face a tradeoff between minimizing density-independent predation risk while crossing the reef crest vs. avoiding high competitor density on settlement habitat. In general, we expect competition to spread spawning over time such that settlement is distributed around the lunar phase with the lowest predation risk, similar to an ideal free distribution in which competition spreads competitors across space. We examine the effects of overcompensating density dependence, age-dependent competition, and competition among daily settler cohorts. Our model predicts that even in the absence of stochastic variation in the larval environment, competition can result in qualitative divergence between spawning and settlement distributions. Furthermore, we show that if competitive strength increases with settler age, competition results in covariation between settler age and settlement date, with older larvae settling when predation risk is minimal. We predict that competition between daily cohorts delays peak settlement, with priority effects potentially selecting for a multimodal settlement distribution.

Intraspecific priority effects in response to egg hatching delay in a pond-breeding salamander

As reproduction phenologies shift with climate change, populations can experience intraspecific priority effects, wherein early hatching cohorts experience an advantage over late-hatching cohorts, resulting in altered demography. Our study objective was to identify how variation in egg hatching phenology alters intraspecific interactions in small-mouthed salamanders, Ambystoma texanum. We addressed two research questions: (Q1) How are demographic responses altered by variation in the temporal duration of hatching between cohorts, and (Q2) How does the seasonality of hatching delays affect demographic responses? We manipulated hatching phenologies of A. texanum eggs and reared larvae in outdoor mesocosms to metamorphosis. For Q1, hatching delay exhibited non-linear relationships with survival and body size, with the greatest asynchrony in cohort additions resulting in the highest mortality and largest body sizes. For Q2, hatching delay effects were stronger (i.e., survival was lower and body sizes larger) when they occurred later in the season, potentially due to temperature differences that larvae experienced. Overall, our results demonstrate that changes in intraspecific interactions due to phenological shifts can be context-dependent, depending on the strength (i.e., temporal duration) and seasonality of such processes. Identifying context-dependencies of phenological shifts will be critical for predicting changes in organismal demographics with climatic shifts.

Ecological filtering drives rapid spatiotemporal dynamics in fish skin microbiomes.

Skin microbiomes provide vital functions, yet knowledge about the drivers and processes structuring their species assemblages is limited-especially for non-model organisms. In this study, fish skin microbiome was assessed by high throughput sequencing of amplicon sequence variants from metabarcoding of V3-V4 regions in the 16S rRNA gene on fish hosts subjected to the following experimental manipulations: (i) translocation between fresh and brackish water habitats to investigate the role of environment; (ii) treatment with an antibacterial disinfectant to reboot the microbiome and investigate community assembly and priority effects; and (iii) maintained alone or in pairs to study the role of social environment and inter-host dispersal of microbes. The results revealed that fish skin microbiomes harbour a highly dynamic microbial composition that was distinct from bacterioplankton communities in the ambient water. Microbiome composition first diverged as an effect of translocation to either the brackish or freshwater habitat. When the freshwater individuals were translocated back to brackish water, their microbiome composition converged towards the fish microbiomes in the brackish habitat. In summary, external environmental conditions and individual-specific factors jointly determined the community composition dynamics, whereas inter-host dispersal had negligible effects. The dynamics of the microbiome composition was seemingly non-affected by reboot treatment, pointing towards high resilience to disturbance. The results emphasised the role of inter-individual variability for the unexplained variation found in many host-microbiome systems, although the mechanistic underpinnings remain to be identified.

Scarification under the canopy of Japanese oak: should pre-harvest treatment coincide with the masting year of acorns?

ABSTRACT The regeneration of Japanese oak (Quercus crispula Blume) has turned even more pressing in recent years. The pre-harvest treatment, which accumulates advanced seedlings, is essential for its natural regeneration in stands with a dense dwarf bamboo cover. Such pre-harvest treatment is considered particularly effective when done in conjunction with a mast year of acorns, but it is difficult to adjust the operation to specific timing. If there is no mast year for several years after the pre-harvest treatment, there is concern that the “priority effect” from other plant species may inhibit Japanese oak regeneration. The objective of this study was to determine whether scarification under the canopy of Japanese oaks, treated several years before the masting of acorns, is effective for its regeneration. Seedlings and vegetation were surveyed at sites scarified four years before (PRE) and just before masting (MAST) to compare the effects of treatments, i.e. differences in scarification years on the number and/or size of tall trees and other plant species. PRE could limit the predominance of dwarf bamboos and birches, and sufficient number of seedlings (>10 stems m−2) for future development was observed in the fifth year after masting. Since Japanese oak generally has a masting cycle of less than 5 years at most, we can conclude that the timing of pre-harvest treatment is not constrained by the masting year. Further studies are required to determine the details of priority effects of competitive plants on oak seedlings to establish a natural forest management for Japanese oak.

Order of arrival and nutrient supply alter outcomes of co-infection with two fungal pathogens.

A pathogen arriving on a host typically encounters a diverse community of microbes that can shape priority effects, other within-host interactions and infection outcomes. In plants, environmental nutrients can drive trade-offs between host growth and defence and can mediate interactions between co-infecting pathogens. Nutrients may thus alter the outcome of pathogen priority effects for the host, but this possibility has received little experimental investigation. To disentangle the relationship between nutrient availability and co-infection dynamics, we factorially manipulated the nutrient availability and order of arrival of two foliar fungal pathogens (Rhizoctonia solani and Colletotrichum cereale) on the grass tall fescue (Lolium arundinaceum) and tracked disease outcomes. Nutrient addition did not influence infection rates, infection severity or plant biomass. Colletotrichum cereale facilitated R. solani, increasing its infection rate regardless of their order of inoculation. Additionally, simultaneous and C. cereale-first inoculations decreased plant growth and-in plants that did not receive nutrient addition-increased leaf nitrogen concentrations compared to uninoculated plants. These effects were partially, but not completely, explained by the duration and severity of pathogen infections. This study highlights the importance of understanding the intricate associations between the order of pathogen arrival, host nutrient availability and host defence to better predict infection outcomes.

Combined toxicity of Cd and aniline to soil bacteria varying with exposure sequence

Joint toxicity of organic-metal co-contamination can vary depending on organisms, toxicants, and even the sequence of exposure. This study examines how the combined toxicity of aniline (An) and cadmium (Cd) to soil bacteria in microcosms changes when the order of contaminant introduction is altered. Through analyzing biodiversity, molecular ecological network, functional redundancy, functional genes and pathways, we find the treatment of Cd followed by An brings about the strongest adverse impact to the bacterial consortium, followed by the reverse-ordered exposure and the simple mixture of the two chemicals. On the level of individual organisms, exposure sequence also affects the bacteria that are otherwise resistant to the standalone toxicity of both An and Cd. The dynamic behavior of aniline-cadmium composite is interpreted by considering the tolerance of organisms to individual chemicals, the interactions of the two toxicants, the recovery time, as well as the priority effect. The overall effect of the composite contamination is conceptualized by treating the chemicals as environmental filters screening the growth of the community.

Practical applications of soil microbiota to improve ecosystem restoration: current knowledge and future directions.

Soil microbiota are important components of healthy ecosystems. Greater consideration of soil microbiota in the restoration of biodiverse, functional, and resilient ecosystems is required to address the twin global crises of biodiversity decline and climate change. In this review, we discuss available and emerging practical applications of soil microbiota into (i) restoration planning, (ii) direct interventions for shaping soil biodiversity, and (iii) strategies for monitoring and predicting restoration trajectories. We show how better planning of restoration activities to account for soil microbiota can help improve progress towards restoration targets. We show how planning to embed soil microbiota experiments into restoration projects will permit a more rigorous assessment of the effectiveness of different restoration methods, especially when complemented by statistical modelling approaches that capitalise on existing data sets to improve causal understandings and prioritise research strategies where appropriate. In addition to recovering belowground microbiota, restoration strategies that include soil microbiota can improve the resilience of whole ecosystems. Fundamentally, restoration planning should identify appropriate reference target ecosystem attributes and - from the perspective of soil microbiota - comprehensibly consider potential physical, chemical and biological influences on recovery. We identify that inoculating ecologically appropriate soil microbiota into degraded environments can support a range of restoration interventions (e.g. targeted, broad-spectrum and cultured inoculations) with promising results. Such inoculations however are currently underutilised and knowledge gaps persist surrounding successful establishment in light of community dynamics, including priority effects and community coalescence. We show how the ecological trajectories of restoration sites can be assessed by characterising microbial diversity, composition, and functions in the soil. Ultimately, we highlight practical ways to apply the soil microbiota toolbox across the planning, intervention, and monitoring stages of ecosystem restoration and address persistent open questions at each stage. With continued collaborations between researchers and practitioners to address knowledge gaps, these approaches can improve current restoration practices and ecological outcomes.