Slow expanders invade by forming dented fronts in microbial colonies.

Most organisms grow in space, whether they are viruses spreading within a host tissue or invasive species colonizing a new continent. Evolution typically selects for higher expansion rates during spatial growth, but it has been suggested that slower expanders can take over under certain conditions. Here, we report an experimental observation of such population dynamics. We demonstrate that mutants that grow slower in isolation nevertheless win in competition, not only when the two types are intermixed, but also when they are spatially segregated into sectors. The latter was thought to be impossible because previous studies focused exclusively on the global competitions mediated by expansion velocities, but overlooked the local competitions at sector boundaries. Local competition, however, can enhance the velocity of either type at the sector boundary and thus alter expansion dynamics. We developed a theory that accounts for both local and global competitions and describes all possible sector shapes. In particular, the theory predicted that a slower on its own, but more competitive, mutant forms a dented V-shaped sector as it takes over the expansion front. Such sectors were indeed observed experimentally, and their shapes matched quantitatively with the theory. In simulations, we further explored several mechanisms that could provide slow expanders with a local competitive advantage and showed that they are all well-described by our theory. Taken together, our results shed light on previously unexplored outcomes of spatial competition and establish a universal framework to understand evolutionary and ecological dynamics in expanding populations.

National polarization and international agreements.

The network of international environmental agreements (IEAs) has been characterized as a complex adaptive system (CAS) in which the uncoordinated responses of nation states to changes in the conditions addressed by particular agreements may generate seemingly coordinated patterns of behavior at the level of the system. Unfortunately, since the rules governing national responses are ill understood, it is not currently possible to implement a CAS approach. Polarization of both political parties and the electorate has been implicated in a secular decline in national commitment to some IEAs, but the causal mechanisms are not clear. In this paper, we explore the impact of polarization on the rules underpinning national responses. We identify the degree to which responsibility for national decisions is shared across political parties and calculate the electoral cost of party positions as national obligations under an agreement change. We find that polarization typically affects the degree but not the direction of national responses. Whether national commitment to IEAs strengthens or weakens as national obligations increase depends more on the change in national obligations than on polarization per se. Where the rules governing national responses are conditioned by the current political environment, so are the dynamic consequences both for the agreement itself and for the network to which it belongs. Any CAS analysis requires an understanding of such conditioning effects on the rules governing national responses.

Exploiting spatial dimensions to enable parallelized continuous directed evolution.

Current strategies to improve the throughput of continuous directed evolution technologies often involve complex mechanical fluid-controlling system or robotic platforms, which limits their popularization and application in general laboratories. Inspired by our previous study on bacterial range expansion, in this study, we report a system termed SPACE for rapid and extensively parallelizable evolution of biomolecules by introducing spatial dimensions into the landmark phage-assisted continuous evolution system. Specifically, M13 phages and chemotactic Escherichia coli cells were closely inoculated onto a semisolid agar. The phages came into contact with the expanding front of the bacterial range, and then comigrated with the bacteria. This system leverages competition over space, wherein evolutionary progress is closely associated with the production of spatial patterns, allowing the emergence of improved or new protein functions. In a prototypical problem, SPACE remarkably simplified the process and evolved the promoter recognition of T7 RNA polymerase (RNAP) to a library of 96 random sequences in parallel. These results establish SPACE as a simple, easy to implement, and massively parallelizable platform for continuous directed evolution in general laboratories.

Geographic accessibility and hospital competition for emergency blood transfusion services in Bungoma, Western Kenya.

BACKGROUND: Estimating accessibility gaps to essential health interventions helps to allocate and prioritize health resources. Access to blood transfusion represents an important emergency health requirement. Here, we develop geo-spatial models of accessibility and competition to blood transfusion services in Bungoma County, Western Kenya. METHODS: Hospitals providing blood transfusion services in Bungoma were identified from an up-dated geo-coded facility database. AccessMod was used to define care-seeker's travel times to the nearest blood transfusion service. A spatial accessibility index for each enumeration area (EA) was defined using modelled travel time, population demand, and supply available at the hospital, assuming a uniform risk of emergency occurrence in the county. To identify populations marginalized from transfusion services, the number of people outside 1-h travel time and those residing in EAs with low accessibility indexes were computed at the sub-county level. Competition between the transfusing hospitals was estimated using a spatial competition index which provided a measure of the level of attractiveness of each hospital. To understand whether highly competitive facilities had better capacity for blood transfusion services, a correlation test between the computed competition metric and the blood units received and transfused at the hospital was done. RESULTS: 15 hospitals in Bungoma county provide transfusion services, however these are unevenly distributed across the sub-counties. Average travel time to a blood transfusion centre in the county was 33 min and 5% of the population resided outside 1-h travel time. Based on the accessibility index, 38% of the EAs were classified to have low accessibility, representing 34% of the population, with one sub-county having the highest marginalized population. The computed competition index showed that hospitals in the urban areas had a spatial competitive advantage over those in rural areas. CONCLUSION: The modelled spatial accessibility has provided an improved understanding of health care gaps essential for health planning. Hospital competition has been illustrated to have some degree of influence in provision of health services hence should be considered as a significant external factor impacting the delivery, and re-design of available services.

Habitat heterogeneity mediates effects of individual variation on spatial species coexistence.

Numerous studies have documented the importance of individual variation (IV) in determining the outcome of competition between species. However, little is known about how the interplay between IV and habitat heterogeneity (i.e. variation and spatial autocorrelation in habitat quality) affects species coexistence at the landscape scale. Here, we incorporate habitat heterogeneity into a competition model with IV, in order to explore the mechanism of spatial species coexistence. We find that individual-level variation and habitat heterogeneity interact to promote species coexistence, more obviously at lower dispersal rates. This is in stark contrast to early non-spatial models, which predicted that IV reinforces competitive hierarchies and therefore speeds up species exclusion. In essence, increasing variation in patch quality and/or spatial habitat autocorrelation moderates differences in the competitive ability of species, thereby allowing species to coexist both locally and globally. Overall, our theoretical study offers a mechanistic explanation for emerging empirical evidence that both habitat heterogeneity and IV promote species coexistence and therefore biodiversity maintenance.

Interspecific interactions through 2 million years: are competitive outcomes predictable?

Ecological interactions affect the survival and reproduction of individuals. However, ecological interactions are notoriously difficult to measure in extinct populations, hindering our understanding of how the outcomes of interactions such as competition vary in time and influence long-term evolutionary changes. Here, the outcomes of spatial competition in a temporally continuous community over evolutionary timescales are presented for the first time. Our research domain is encrusting cheilostome bryozoans from the Wanganui Basin of New Zealand over a ca 2 Myr time period (Pleistocene to Recent). We find that a subset of species can be identified as consistent winners, and others as consistent losers, in the sense that they win or lose interspecific competitive encounters statistically more often than the null hypothesis of 50%. Most species do not improve or worsen in their competitive abilities through the 2 Myr period, but a minority of species are winners in some intervals and losers in others. We found that conspecifics tend to cluster spatially and interact more often than expected under a null hypothesis: most of these are stand-off interactions where the two colonies involved stopped growing at edges of encounter. Counterintuitively, competitive ability has no bearing on ecological dominance.

Mating Systems in True Morels (Morchella).

True morels (Morchella spp., Morchellaceae, Ascomycota) are widely regarded as a highly prized delicacy and are of great economic and scientific value. Recently, the rapid development of cultivation technology and expansion of areas for artificial morel cultivation have propelled morel research into a hot topic. Many studies have been conducted in various aspects of morel biology, but despite this, cultivation sites still frequently report failure to fruit or only low production of fruiting bodies. Key problems include the gap between cultivation practices and basic knowledge of morel biology. In this review, in an effort to highlight the mating systems, evolution, and life cycle of morels, we summarize the current state of knowledge of morel sexual reproduction, the structure and evolution of mating-type genes, the sexual process itself, and the influence of mating-type genes on the asexual stages and conidium production. Understanding of these processes is critical for improving technology for the cultivation of morels and for scaling up their commercial production. Morel species may well be good candidates as model species for improving sexual development research in ascomycetes in the future.

Indirect effects of overfishing on Caribbean reefs: sponges overgrow reef-building corals.

Consumer-mediated indirect effects at the community level are difficult to demonstrate empirically. Here, we show an explicit indirect effect of overfishing on competition between sponges and reef-building corals from surveys of 69 sites across the Caribbean. Leveraging the large-scale, long-term removal of sponge predators, we selected overfished sites where intensive methods, primarily fish-trapping, have been employed for decades or more, and compared them to sites in remote or marine protected areas (MPAs) with variable levels of enforcement. Sponge-eating fishes (angelfishes and parrotfishes) were counted at each site, and the benthos surveyed, with coral colonies scored for interaction with sponges. Overfished sites had >3 fold more overgrowth of corals by sponges, and mean coral contact with sponges was 25.6%, compared with 12.0% at less-fished sites. Greater contact with corals by sponges at overfished sites was mostly by sponge species palatable to sponge predators. Palatable species have faster rates of growth or reproduction than defended sponge species, which instead make metabolically expensive chemical defenses. These results validate the top-down conceptual model of sponge community ecology for Caribbean reefs, as well as provide an unambiguous justification for MPAs to protect threatened reef-building corals. An unanticipated outcome of the benthic survey component of this study was that overfished sites had lower mean macroalgal cover (23.1% vs. 38.1% for less-fished sites), a result that is contrary to prevailing assumptions about seaweed control by herbivorous fishes. Because we did not quantify herbivores for this study, we interpret this result with caution, but suggest that additional large-scale studies comparing intensively overfished and MPA sites are warranted to examine the relative impacts of herbivorous fishes and urchins on Caribbean reefs.

Detecting regular spacing in patchy environments and estimating its density using nearest-neighbour graphical analysis.

Organisms can aggregate to form patches or clusters interspersed with interstices of unoccupied space for behavioural or ecological reasons. Territorial spacing in the clusters could follow, resulting in regularity. In these circumstances an analysis of spatial pattern in a sample area defined by a boundary enclosing the clusters might reveal the aggregation but not the regularity. To demonstrate the occurrence of regularity, spacing trends and an estimate of density based on a nearest-neighbour graphical analysis can be used. The method is independent of boundaries and the presence of interstices. The basis of the method, and its strengths and limitations, are examined. A re-analysis of acorn woodpecker data, which has generated some controversy in the past, illustrates its potential for revealing "hidden" regularity.

Small-scale association measures in epibenthic communities as a clue for allelochemical interactions.

The small-scale associations in a rocky subtidal community in the northwestern Mediterranean were studied by a development of the continuous line transect method. This method allowed the overall measurement of non-randomness in interspecific contacts and the assignment of an association index to each species-pair, whose, significance was tested by Monte Carlo procedures. At the same time, the continuous recording allowed the study of the weakening of the interactions with increasing distances. Our purpose was to uncover evidence for allelochemical mechanisms of space occupation and maintenance. A strong non-randomness was found in the interspecific associations. This was mostly due to the interactions of the poecilosclerid sponge Crambe crambe (Schmidt) with its neighbours, especially its negative associations with other sponge species. The strength of the relationships fell drastically over the first few centimeters from the contact borders of the different species. The results pointed strongly to an allelochemical mechanism. The extracts of this sponge featured high bioactivity in laboratory assays, and field experiments demonstrated that the sponge can inhibit the growth of species in the community studied. Standard sampling techniques would have overlooked the spatial structure present in the data. The study emphasizes the need for both contact data and distance data in order to identify the underlying processes reliably. The line transect method provides both types of information easily and allows testing of models and identification of organisms likely to use chemical defenses in space competition. Its use as a preliminary step in studies of chemical ecology might help to detect presumptive allelochemical processes prior to experimental work on the potentially active species.

Associations between spatial position, stress and anxiety in forest baboons Papio anubis.

Spatial position within a group affects the value of group-living benefits such as reduced predation risk and improved foraging. The threat of predation, poor nutrition or increased competition from conspecifics can all cause stress. In many species, central positions are known to be more beneficial than peripheral positions in terms of reduced predation, vigilance and foraging. In this study, we examine whether spatial position within a group is associated with stress and anxiety in a troop of olive baboons (Papio anubis). We predicted that the benefits of occupying central positions would be reflected by a reduction in stress and anxiety for animals who spent the most time in the centre of the group. The study subjects appeared to compete actively for the centre of the group. Physiological stress measures (faecal glucocorticoid concentrations) were positively correlated with time spent in central positions. Time spent in central positions was positively correlated with proximity but negatively correlated with vigilance behaviours (alarm barks). Vigilance rates were positively correlated with measures of anxiety (self-scratch frequency). It is suggested that individuals experience chronic stress due to proximity to conspecifics in central positions, whilst perceived predation risk causes anxiety, with perceived predation risk experienced more by individuals on the periphery.