Pseudomonas syringae infectivity correlates to altered transcript and metabolite levels of Arabidopsis mediator mutants.

Rapid metabolic responses to pathogens are essential for plant survival and depend on numerous transcription factors. Mediator is the major transcriptional co-regulator for integration and transmission of signals from transcriptional regulators to RNA polymerase II. Using four Arabidopsis Mediator mutants, med16, med18, med25 and cdk8, we studied how differences in regulation of their transcript and metabolite levels correlate to their responses to Pseudomonas syringae infection. We found that med16 and cdk8 were susceptible, while med25 showed increased resistance. Glucosinolate, phytoalexin and carbohydrate levels were reduced already before infection in med16 and cdk8, but increased in med25, which also displayed increased benzenoids levels. Early after infection, wild type plants showed reduced glucosinolate and nucleoside levels, but increases in amino acids, benzenoids, oxylipins and the phytoalexin camalexin. The Mediator mutants showed altered levels of these metabolites and in regulation of genes encoding key enzymes for their metabolism. At later stage, mutants displayed defective levels of specific amino acids, carbohydrates, lipids and jasmonates which correlated to their infection response phenotypes. Our results reveal that MED16, MED25 and CDK8 are required for a proper, coordinated transcriptional response of genes which encode enzymes involved in important metabolic pathways for Arabidopsis responses to Pseudomonas syringae infections.

Co-PATHOgenex web application for assessing complex stress responses in pathogenic bacteria.

IMPORTANCE: Unveiling gene co-expression networks in bacterial pathogens has the potential for gaining insights into their adaptive strategies within the host environment. Here, we developed Co-PATHOgenex, an interactive and user-friendly web application that enables users to construct networks from gene co-expressions using custom-defined thresholds (https://avicanlab.shinyapps.io/copathogenex/). The incorporated search functions and visualizations within the tool simplify the usage and facilitate the interpretation of the analysis output. Co-PATHOgenex also includes stress stimulons for various bacterial species, which can help identify gene products not previously associated with a particular stress condition.

Mapping nonlocal relationships between metadata and network structure with metadata-dependent encoding of random walks.

Integrating structural information and metadata, such as gender, social status, or interests, enriches networks and enables a better understanding of the large-scale structure of complex systems. However, existing approaches to augment networks with metadata for community detection only consider immediately adjacent nodes and cannot exploit the nonlocal relationships between metadata and large-scale network structure present in many spatial and social systems. Here, we develop a flow-based community detection framework based on the map equation that integrates network information and metadata of distant nodes and reveals more complex relationships. We analyze social and spatial networks and find that our methodology can detect functional metadata-informed communities distinct from those derived solely from network information or metadata. For example, in a mobility network of London, we identify communities that reflect the heterogeneity of income distribution, and in a European power grid network, we identify communities that capture relationships between geography and energy prices beyond country borders.

Socioeconomic and environmental patterns behind H1N1 spreading in Sweden.

Identifying the critical factors related to influenza spreading is crucial in predicting and mitigating epidemics. Specifically, uncovering the relationship between epidemic onset and various risk indicators such as socioeconomic, mobility and climate factors can reveal locations and travel patterns that play critical roles in furthering an outbreak. We study the 2009 A(H1N1) influenza outbreaks in Sweden's municipalities between 2009 and 2015 and use the Generalized Inverse Infection Method (GIIM) to assess the most significant contributing risk factors. GIIM represents an epidemic spreading process on a network: nodes correspond to geographical objects, links indicate travel routes, and transmission probabilities assigned to the links guide the infection process. Our results reinforce existing observations that the influenza outbreaks considered in this study were driven by the country's largest population centers, while meteorological factors also contributed significantly. Travel and other socioeconomic indicators have a negligible effect. We also demonstrate that by training our model on the 2009 outbreak, we can predict the epidemic onsets in the following five seasons with high accuracy.

A multiscale view of the Phanerozoic fossil record reveals the three major biotic transitions.

The hypothesis of the Great Evolutionary Faunas is a foundational concept of macroevolutionary research postulating that three global mega-assemblages have dominated Phanerozoic oceans following abrupt biotic transitions. Empirical estimates of this large-scale pattern depend on several methodological decisions and are based on approaches unable to capture multiscale dynamics of the underlying Earth-Life System. Combining a multilayer network representation of fossil data with a multilevel clustering that eliminates the subjectivity inherent to distance-based approaches, we demonstrate that Phanerozoic oceans sequentially harbored four global benthic mega-assemblages. Shifts in dominance patterns among these global marine mega-assemblages were abrupt (end-Cambrian 494 Ma; end-Permian 252 Ma) or protracted (mid-Cretaceous 129 Ma), and represent the three major biotic transitions in Earth's history. Our findings suggest that gradual ecological changes associated with the Mesozoic Marine Revolution triggered a protracted biotic transition comparable in magnitude to the end-Permian transition initiated by the most severe biotic crisis of the past 500 million years. Overall, our study supports the notion that both long-term ecological changes and major geological events have played crucial roles in shaping the mega-assemblages that dominated Phanerozoic oceans.

Impacts of Oxazepam on Perch (Perca fluviatilis) Behavior: Fish Familiarized to Lake Conditions Do Not Show Predicted Anti-anxiety Response.

A current theory in environmental science states that dissolved anxiolytics (oxazepam) from wastewater effluents can reduce anti-predator behavior in fish with potentially negative impacts on prey fish populations. Here, we hypothesize that European perch (Perca fluviatilis) populations being exposed to oxazepam in situ show reduced anti-predator behavior, which has previously been observed for exposed isolated fish in laboratory studies. We tested our hypothesis by exposing a whole-lake ecosystem, containing both perch (prey) and northern pike (Esox lucius; predator), to oxazepam while tracking fish behavior before and after exposure in the exposed lake as well as in an unexposed nearby lake (control). Oxazepam concentrations in the exposed lake ranged between 11 and 24 µg L-1, which is >200 times higher than concentrations reported for European rivers. In contrast to our hypothesis, we did not observe an oxazepam-induced reduction in anti-predator behavior, inferred from perch swimming activity, distance to predators, distance to conspecifics, home-range size, and habitat use. In fact, exposure to oxazepam instead stimulated anti-predator behavior (decreased activity, decreased distance to conspecifics, and increased littoral habitat use) when using behavior in the control lake as a reference. Shoal dynamics and temperature changes may have masked modest reductions in anti-predator behavior due to oxazepam. Although we cannot fully resolve the mechanism(s) behind our observations, our results indicate that the effects of oxazepam on perch behavior in a familiar natural ecosystem are negligible in comparison to the effects of other environmental conditions.

Regularities in species' niches reveal the world's climate regions.

Climate regions form the basis of many ecological, evolutionary, and conservation studies. However, our understanding of climate regions is limited to how they shape vegetation: they do not account for the distribution of animals. Here, we develop a network-based framework to identify important climates worldwide based on regularities in realized niches of about 26,000 tetrapods. We show that high-energy climates, including deserts, tropical savannas, and steppes, are consistent across animal- and plant-derived classifications, indicating similar underlying climatic determinants. Conversely, temperate climates differ across all groups, suggesting that these climates allow for idiosyncratic adaptations. Finally, we show how the integration of niche classifications with geographical information enables the detection of climatic transition zones and the signal of geographic and historical processes. Our results identify the climates shaping the distribution of tetrapods and call for caution when using general climate classifications to study the ecology, evolution, or conservation of specific taxa.

There are many distinct climates on Earth, from tropical savannas and temperate forests to dry deserts. Historically, each region has been defined by how its annual weather patterns shape the type of vegetation present. For example, hot and humid environments support the growth of evergreen forests that would not survive in drier places. Identifying the boundaries between climate regions is key to understanding how life is organized on Earth and predicting how climate change will affect different species. Current climate classifications, however, do not account for where animals can be found or how local conditions, such as precipitation and average temperatures, shape the distribution of different animal species. To bridge this gap, Calatayud et al. analyzed the preferred climate of about 26,000 animal species, including amphibians, birds, mammals and reptiles. For each species, Calatayud et al. calculated the annual rainfall and temperature of its local environment, or 'niche', using previously collected data. They then used a computer algorithm to group together climates that had similar species. This identified 16 climate regions which govern the distribution of the animals studied. Calatayud et al. found that these newly defined climatic regions resembled some of the regions classified using plants. This was particularly true for high-energy climates that had lower levels of rainfall and hot temperatures, such as deserts and the tropical savanna. The animals and plant species living in high-energy regions were found to be fairly consistent across both classification systems. Whereas the species present in milder and colder climates, such as temperate forests or Mediterranean climates, were found to be much more varied. This suggests that temperate climates are harder to classify and may affect the distribution of plants and animals differently. It also implies that less extreme conditions support a larger range of species than harsher climates in which only species with certain adaptations are able to survive. These findings build the basis for a better understanding of how climates shape ecosystems. More specific climate classifications, based on such analyses, could be used to inform conservation strategies for animal species in the face of climate change.

Mapping flows on bipartite networks.

Mapping network flows provides insight into the organization of networks, but even though many real networks are bipartite, no method for mapping flows takes advantage of the bipartite structure. What do we miss by discarding this information and how can we use it to understand the structure of bipartite networks better? The map equation models network flows with a random walk and exploits the information-theoretic duality between compression and finding regularities to detect communities in networks. However, it does not use the fact that random walks in bipartite networks alternate between node types, information worth 1 bit. To make some or all of this information available to the map equation, we developed a coding scheme that remembers node types at different rates. We explored the community landscape of bipartite real-world networks from no node-type information to full node-type information and found that using node types at a higher rate generally leads to deeper community hierarchies and a higher resolution. The corresponding compression of network flows exceeds the amount of extra information provided. Consequently, taking advantage of the bipartite structure increases the resolution and reveals more network regularities.

Mapping flows on sparse networks with missing links.

Unreliable network data can cause community-detection methods to overfit and highlight spurious structures with misleading information about the organization and function of complex systems. Here we show how to detect significant flow-based communities in sparse networks with missing links using the map equation. Since the map equation builds on Shannon entropy estimation, it assumes complete data such that analyzing undersampled networks can lead to overfitting. To overcome this problem, we incorporate a Bayesian approach with assumptions about network uncertainties into the map equation framework. Results in both synthetic and real-world networks show that the Bayesian estimate of the map equation provides a principled approach to revealing significant structures in undersampled networks.

Positive associations among rare species and their persistence in ecological assemblages.

According to the competitive exclusion principle, species with low competitive abilities should be excluded by more efficient competitors; yet, they generally remain as rare species. Here, we describe the positive and negative spatial association networks of 326 disparate assemblages, showing a general organization pattern that simultaneously supports the primacy of competition and the persistence of rare species. Abundant species monopolize negative associations in about 90% of the assemblages. On the other hand, rare species are mostly involved in positive associations, forming small network modules. Simulations suggest that positive interactions among rare species and microhabitat preferences are the most probable mechanisms underpinning this pattern and rare species persistence. The consistent results across taxa and geography suggest a general explanation for the maintenance of biodiversity in competitive environments.

Exploring the solution landscape enables more reliable network community detection.

To understand how a complex system is organized and functions, researchers often identify communities in the system's network of interactions. Because it is practically impossible to explore all solutions to guarantee the best one, many community-detection algorithms rely on multiple stochastic searches. But for a given combination of network and stochastic algorithms, how many searches are sufficient to find a solution that is good enough? The standard approach is to pick a reasonably large number of searches and select the network partition with the highest quality or derive a consensus solution based on all network partitions. However, if different partitions have similar qualities such that the solution landscape is degenerate, the single best partition may miss relevant information, and a consensus solution may blur complementary communities. Here we address this degeneracy problem with coarse-grained descriptions of the solution landscape. We cluster network partitions based on their similarity and suggest an approach to determine the minimum number of searches required to describe the solution landscape adequately. To make good use of all partitions, we also propose different ways to explore the solution landscape, including a significance clustering procedure. We test these approaches on synthetic networks and a real-world network using two contrasting community-detection algorithms: The algorithm that can identify more general structures requires more searches, and networks with clearer community structures require fewer searches. We also find that exploring the coarse-grained solution landscape can reveal complementary solutions and enable more reliable community detection.

Human activity is altering the world's zoogeographical regions.

Zoogeographical regions, or zooregions, are areas of the Earth defined by species pools that reflect ecological, historical and evolutionary processes acting over millions of years. Consequently, researchers have assumed that zooregions are robust and unlikely to change on a human timescale. However, the increasing number of human-mediated introductions and extinctions can challenge this assumption. By delineating zooregions with a network-based algorithm, here we show that introductions and extinctions are altering the zooregions we know today. Introductions are homogenising the Eurasian and African mammal zooregions and also triggering less intuitive effects in birds and amphibians, such as dividing and redefining zooregions representing the Old and New World. Furthermore, these Old and New World amphibian zooregions are no longer detected when considering introductions plus extinctions of the most threatened species. Our findings highlight the profound and far-reaching impact of human activity and call for identifying and protecting the uniqueness of biotic assemblages.

From networks to optimal higher-order models of complex systems.

Rich data is revealing that complex dependencies between the nodes of a network may escape models based on pairwise interactions. Higher-order network models go beyond these limitations, offering new perspectives for understanding complex systems.

Constrained information flows in temporal networks reveal intermittent communities.

Many real-world networks represent dynamic systems with interactions that change over time, often in uncoordinated ways and at irregular intervals. For example, university students connect in intermittent groups that repeatedly form and dissolve based on multiple factors, including their lectures, interests, and friends. Such dynamic systems can be represented as multilayer networks where each layer represents a snapshot of the temporal network. In this representation, it is crucial that the links between layers accurately capture real dependencies between those layers. Often, however, these dependencies are unknown. Therefore, current methods connect layers based on simplistic assumptions that do not capture node-level layer dependencies. For example, connecting every node to itself in other layers with the same weight can wipe out dependencies between intermittent groups, making it difficult or even impossible to identify them. In this paper, we present a principled approach to estimating node-level layer dependencies based on the network structure within each layer. We implement our node-level coupling method in the community detection framework Infomap and demonstrate its performance compared to current methods on synthetic and real temporal networks. We show that our approach more effectively constrains information inside multilayer communities so that Infomap can better recover planted groups in multilayer benchmark networks that represent multiple modes with different groups and better identify intermittent communities in real temporal contact networks. These results suggest that node-level layer coupling can improve the modeling of information spreading in temporal networks and better capture intermittent community structure.

Modelling sequences and temporal networks with dynamic community structures.

In evolving complex systems such as air traffic and social organisations, collective effects emerge from their many components' dynamic interactions. While the dynamic interactions can be represented by temporal networks with nodes and links that change over time, they remain highly complex. It is therefore often necessary to use methods that extract the temporal networks' large-scale dynamic community structure. However, such methods are subject to overfitting or suffer from effects of arbitrary, a priori-imposed timescales, which should instead be extracted from data. Here we simultaneously address both problems and develop a principled data-driven method that determines relevant timescales and identifies patterns of dynamics that take place on networks, as well as shape the networks themselves. We base our method on an arbitrary-order Markov chain model with community structure, and develop a nonparametric Bayesian inference framework that identifies the simplest such model that can explain temporal interaction data.The description of temporal networks is usually simplified in terms of their dynamic community structures, whose identification however relies on a priori assumptions. Here the authors present a data-driven method that determines relevant timescales for the dynamics and uses it to identify communities.

The many facets of community detection in complex networks.

Community detection, the decomposition of a graph into essential building blocks, has been a core research topic in network science over the past years. Since a precise notion of what constitutes a community has remained evasive, community detection algorithms have often been compared on benchmark graphs with a particular form of assortative community structure and classified based on the mathematical techniques they employ. However, this comparison can be misleading because apparent similarities in their mathematical machinery can disguise different goals and reasons for why we want to employ community detection in the first place. Here we provide a focused review of these different motivations that underpin community detection. This problem-driven classification is useful in applied network science, where it is important to select an appropriate algorithm for the given purpose. Moreover, highlighting the different facets of community detection also delineates the many lines of research and points out open directions and avenues for future research.

Infomap Bioregions: Interactive Mapping of Biogeographical Regions from Species Distributions.

Biogeographical regions (bioregions) reveal how different sets of species are spatially grouped and therefore are important units for conservation, historical biogeography, ecology, and evolution. Several methods have been developed to identify bioregions based on species distribution data rather than expert opinion. One approach successfully applies network theory to simplify and highlight the underlying structure in species distributions. However, this method lacks tools for simple and efficient analysis. Here, we present Infomap Bioregions, an interactive web application that inputs species distribution data and generates bioregion maps. Species distributions may be provided as georeferenced point occurrences or range maps, and can be of local, regional, or global scale. The application uses a novel adaptive resolution method to make best use of often incomplete species distribution data. The results can be downloaded as vector graphics, shapefiles, or in table format. We validate the tool by processing large data sets of publicly available species distribution data of the world's amphibians using species ranges, and mammals using point occurrences. We then calculate the fit between the inferred bioregions and WWF ecoregions. As examples of applications, researchers can reconstruct ancestral ranges in historical biogeography or identify indicator species for targeted conservation. [Biogeography; bioregionalization; conservation; mapping].

Efficient community detection of network flows for varying Markov times and bipartite networks.

Community detection of network flows conventionally assumes one-step dynamics on the links. For sparse networks and interest in large-scale structures, longer timescales may be more appropriate. Oppositely, for large networks and interest in small-scale structures, shorter timescales may be better. However, current methods for analyzing networks at different timescales require expensive and often infeasible network reconstructions. To overcome this problem, we introduce a method that takes advantage of the inner workings of the map equation and evades the reconstruction step. This makes it possible to efficiently analyze large networks at different Markov times with no extra overhead cost. The method also evades the costly unipartite projection for identifying flow modules in bipartite networks.

Estimating the resolution limit of the map equation in community detection.

A community detection algorithm is considered to have a resolution limit if the scale of the smallest modules that can be resolved depends on the size of the analyzed subnetwork. The resolution limit is known to prevent some community detection algorithms from accurately identifying the modular structure of a network. In fact, any global objective function for measuring the quality of a two-level assignment of nodes into modules must have some sort of resolution limit or an external resolution parameter. However, it is yet unknown how the resolution limit affects the so-called map equation, which is known to be an efficient objective function for community detection. We derive an analytical estimate and conclude that the resolution limit of the map equation is set by the total number of links between modules instead of the total number of links in the full network as for modularity. This mechanism makes the resolution limit much less restrictive for the map equation than for modularity; in practice, it is orders of magnitudes smaller. Furthermore, we argue that the effect of the resolution limit often results from shoehorning multilevel modular structures into two-level descriptions. As we show, the hierarchical map equation effectively eliminates the resolution limit for networks with nested multilevel modular structures.

Memory in network flows and its effects on spreading dynamics and community detection.

Random walks on networks is the standard tool for modelling spreading processes in social and biological systems. This first-order Markov approach is used in conventional community detection, ranking and spreading analysis, although it ignores a potentially important feature of the dynamics: where flow moves to may depend on where it comes from. Here we analyse pathways from different systems, and although we only observe marginal consequences for disease spreading, we show that ignoring the effects of second-order Markov dynamics has important consequences for community detection, ranking and information spreading. For example, capturing dynamics with a second-order Markov model allows us to reveal actual travel patterns in air traffic and to uncover multidisciplinary journals in scientific communication. These findings were achieved only by using more available data and making no additional assumptions, and therefore suggest that accounting for higher-order memory in network flows can help us better understand how real systems are organized and function.