Collective Information Processing and Criticality, Evolution and Limited Attention.

Im ersten Teil analysiere ich die Selbstorganisation zur Kritikalität (hier ein Phasenübergang von Ordnung zu Unordnung) und untersuche, ob Evolution ein möglicher Organisationsmechanismus ist. Die Kernfrage ist, ob sich ein simulierter kohäsiver Schwarm, der versucht, einem Raubtier auszuweichen, durch Evolution selbst zum kritischen Punkt entwickelt, um das Ausweichen zu optimieren? Es stellt sich heraus, dass (i) die Gruppe den Jäger am besten am kritischen Punkt vermeidet, aber (ii) nicht durch einer verstärkten Reaktion, sondern durch strukturelle Veränderungen, (iii) das Gruppenoptimum ist evolutionär unstabiler aufgrund einer maximalen räumlichen Selbstsortierung der Individuen. Im zweiten Teil modelliere ich experimentell beobachtete Unterschiede im kollektiven Verhalten von Fischgruppen, die über mehrere Generationen verschiedenen Arten von größenabhängiger Selektion ausgesetzt waren. Diese Größenselektion soll Freizeitfischerei (kleine Fische werden freigelassen, große werden konsumiert) und die kommerzielle Fischerei mit großen Netzbreiten (kleine/junge Individuen können entkommen) nachahmen. Die zeigt sich, dass das Fangen großer Fische den Zusammenhalt und die Risikobereitschaft der Individuen reduziert. Beide Befunde lassen sich mechanistisch durch einen Aufmerksamkeits-Kompromiss zwischen Sozial- und Umweltinformationen erklären. Im letzten Teil der Arbeit quantifiziere ich die kollektive Informationsverarbeitung im Feld. Das Studiensystem ist eine an sulfidische Wasserbedingungen angepasste Fischart mit einem kollektiven Fluchtverhalten vor Vögeln (wiederholte kollektive Fluchttauchgängen). Die Fische sind etwa 2 Zentimeter groß, aber die kollektive Welle breitet sich über Meter in dichten Schwärmen an der Oberfläche aus. Es zeigt sich, dass die Wellengeschwindigkeit schwach mit der Polarisation zunimmt, bei einer optimalen Dichte am schnellsten ist und von ihrer Richtung relativ zur Schwarmorientierung abhängt.In the first part, I focus on the self-organization to criticality (here an order-disorder phase transition) and investigate if evolution is a possible self-tuning mechanism. Does a simulated cohesive swarm that tries to avoid a pursuing predator self-tunes itself by evolution to the critical point to optimize avoidance? It turns out that (i) the best group avoidance is at criticality but (ii) not due to an enhanced response but because of structural changes (fundamentally linked to criticality), (iii) the group optimum is not an evolutionary stable state, in fact (iv) it is an evolutionary accelerator due to a maximal spatial self-sorting of individuals causing spatial selection. In the second part, I model experimentally observed differences in collective behavior of fish groups subject to multiple generation of different types of size-dependent selection. The real world analog to this experimental evolution is recreational fishery (small fish are released, large are consumed) and commercial fishing with large net widths (small/young individuals can escape). The results suggest that large harvesting reduces cohesion and risk taking of individuals. I show that both findings can be mechanistically explained based on an attention trade-off between social and environmental information. Furthermore, I numerically analyze how differently size-harvested groups perform in a natural predator and fishing scenario. In the last part of the thesis, I quantify the collective information processing in the field. The study system is a fish species adapted to sulfidic water conditions with a collective escape behavior from aerial predators which manifests in repeated collective escape dives. These fish measure about 2 centimeters, but the collective wave spreads across meters in dense shoals at the surface. I find that wave speed increases weakly with polarization, is fastest at an optimal density and depends on its direction relative to shoal orientation

Collective predator evasion: Putting the criticality hypothesis to the test

According to the criticality hypothesis, collective biological systems should operate in a special parameter region, close to so-called critical points, where the collective behavior undergoes a qualitative change between different dynamical regimes. Critical systems exhibit unique properties, which may benefit collective information processing such as maximal responsiveness to external stimuli. Besides neuronal and gene-regulatory networks, recent empirical data suggests that also animal collectives may be examples of self-organized critical systems. However, open questions about self-organization mechanisms in animal groups remain: Evolutionary adaptation towards a group-level optimum (group-level selection), implicitly assumed in the "criticality hypothesis", appears in general not reasonable for fission-fusion groups composed of non-related individuals. Furthermore, previous theoretical work relies on non-spatial models, which ignore potentially important self-organization and spatial sorting effects. Using a generic, spatially-explicit model of schooling prey being attacked by a predator, we show first that schools operating at criticality perform best. However, this is not due to optimal response of the prey to the predator, as suggested by the "criticality hypothesis", but rather due to the spatial structure of the prey school at criticality. Secondly, by investigating individual-level evolution, we show that strong spatial self-sorting effects at the critical point lead to strong selection gradients, and make it an evolutionary unstable state. Our results demonstrate the decisive role of spatio-temporal phenomena in collective behavior, and that individual-level selection is in general not a viable mechanism for self-tuning of unrelated animal groups towards criticality

Acoustic and visual stimuli combined promote stronger responses to aerial predation in fish

Bird predation poses a strong selection pressure on fish. Since birds must enter the water to catch fish, a combination of visual and mechano-acoustic cues (multimodal) characterize an immediate attack, while single cues (unimodal) may represent less dangerous disturbances. We investigated whether fish could use this information to distinguish between non-threatening and dangerous events and adjust their antipredator response to the perceived level of risk. To do so, we investigated the antipredator behavior of the sulphur molly (Poecilia sulphuraria), a small freshwater fish which is almost exclusively preyed on by piscivorous birds in its endemic sulfide spring habitat. In a field survey, we confirmed that these fish frequently have to distinguish between disturbances stemming from attacking birds (multimodal) and those which pose no (immediate) threat such as bird overflights (unimodal). In a laboratory experiment, we then exposed fish to artificial visual and/or acoustic stimuli presented separately or combined. Sensitivity was high regardless of stimulus type and number (more than 96% of fish initiated diving), but fish dove deeper, faster, and for longer when both stimuli were available simultaneously. Based on the system’s high rates of bird activity, we argue that such an unselective dive initiation with subsequent fine-tuning of diving parameters in accordance to cue modality represents an optimal strategy for these fish to save energy necessary to respond to future attacks. Ultimately, our study shows that fish anticipate the imminent risk posed by disturbances linked to bird predation through integrating information from both visual and acoustic cues.Peer Reviewe

Diurnal Changes in Hypoxia Shape Predator-Prey Interaction in a Bird-Fish System

Animals often face changing environments, and behavioral flexibility allows them to rapidly and adaptively respond to abiotic factors that vary more or less regularly. However, abiotic factors that affect prey species do not necessarily affect their predators. Still, the prey’s response might affect the predator indirectly, yet evidence from the wild for such a classical bottom-up effect of abiotic factors shaping several trophic levels remains sparse. In many aquatic environments, daily changes in oxygen concentrations occur frequently. When oxygen levels drop to hypoxic levels, many fishes respond with aquatic surface respiration (ASR), during which they obtain oxygen by skimming the upper, oxygenated surface layer. By increasing time at the surface, fish become more vulnerable to fish-eating birds. We explored these cascading effects in a sulfidic spring system that harbors the endemic sulphur molly (Poecilia sulphuraria) as prey species and several fish-eating bird species. Sulfide-rich springs pose harsh conditions as hydrogen sulfide (H2S) is lethal to most metazoans and reduces dissolved oxygen (DO). Field sampling during three daytimes indicated that water temperatures rose from morning to (after)noon, resulting in the already low DO levels to decrease further, while H2S levels showed no diurnal changes. The drop in DO levels was associated with a decrease in time spent diving in sulphur mollies, which corresponded with an increase in ASR. Interestingly, the laboratory-estimated threshold at which the majority of sulphur mollies initiate ASR (ASR50: <1.7 mg/L DO) was independent of temperature and this value was exceeded daily when hypoxic stress became more severe toward noon. As fish performed ASR, large aggregations built up at the water surface over the course of the day. As a possible consequence of fish spending more time at the surface, we found high activity levels of fish-eating birds at noon and in the afternoon. Our study reveals that daily fluctuations in water’s oxygen levels have the potential to alter predator-prey interactions profoundly and thus highlights the joined actions of abiotic and biotic factors shaping the evolution of a prey species.Peer Reviewe

Impact of Variable Speed on Collective Movement of Animal Groups

The collective dynamics and structure of animal groups has attracted the attention of scientists across a broad range of fields. A variety of agent-based models have been developed to help understand the emergence of coordinated collective behavior from simple interaction rules. A common, simplifying assumption of such collective movement models, is that individual agents move with a constant speed. In this work we critically re-asses this assumption. First, we discuss experimental data showcasing the omnipresent speed variability observed in different species of live fish and artificial agents (RoboFish). Based on theoretical considerations accounting for inertia and rotational friction, we derive a functional dependence of the turning response of individuals on their instantaneous speed, which is confirmed by experimental data. We then investigate the interplay of variable speed and speed-dependent turning on self-organized collective behavior by implementing an agent-based model which accounts for both these effects. We show that, besides the average speed of individuals, the variability in individual speed can have a dramatic impact on the emergent collective dynamics: a group which differs to another only in a lower speed variability of its individuals (groups being identical in all other behavioral parameters), can be in the polarized state while the other group is disordered. We find that the local coupling between group polarization and individual speed is strongest at the order-disorder transition, and that, in contrast to fixed speed models, the group’s spatial extent does not have a maximum at the transition. Furthermore, we demonstrate a decrease in polarization with group size for groups of individuals with variable speed, and a sudden decrease in mean individual speed at a critical group size (N = 4 for Voronoi interactions) linked to a topological transition from an all-to-all to a distributed spatial interaction network. Overall, our work highlights the importance to account for fundamental kinematic constraints in general, and variable speed in particular, when modeling self-organized collective dynamics.Peer Reviewe

Evolutionary Impact of Size-Selective Harvesting on Shoaling Behavior: Individual-Level Mechanisms and Possible Consequences for Natural and Fishing Mortality

16 pages, 5 figures, 1 table, supplementary data https://doi.org/10.5061/dryad.x95x69pkf; supplementary material https://www.journals.uchicago.edu/doi/suppl/10.1086/718591.-- Data and Code Availability: Experimental data and the R code have been deposited in the Dryad Digital Repository (https://doi.org/10.5061/dryad.x95x69pkf; Sbragaglia 2021). The code to run the burst-and-coast model with the fishing/predator scenarios is available at GitHub (https://doi.org/10.5281/zenodo.5651937; https://github.com/PaPeK/sde_burst_coast). The optimization was done with a Python implementation of the CMA-ES (https://github.com/CMA-ES/pycma)Intensive and size-selective harvesting is an evolutionary driver of life history as well as individual behavioral traits. Yet whether and to what degree harvesting modifies the collective behavior of exploited species are largely unknown. We present a multigeneration harvest selection experiment with zebrafish, Danio rerio, as a model species to understand the effects of size-selective harvesting on shoaling behavior. The experimental system is based on a large-harvested (typical of most wild-capture fisheries targeting larger size classes) and small-harvested (typical of specialized fisheries and gape-limited predators targeting smaller size classes) selection lines. By combining high-resolution tracking of fish behavior with computational agent-based modeling, we show that shoal cohesion changed in the direction expected by a trade-off between individual vigilance and the use of social cues. In particular, we document a decrease of individual vigilance in the small-harvested line, which was linked to an increase in the attention to social cues, favoring more cohesive shoals. Opposing outcomes were found for the large-harvested line, which formed less cohesive shoals. Using the agent-based model, we outline possible consequences of changes in shoaling behavior for both fishing and natural mortality. The changes in shoaling induced by large size-selective harvesting may decrease fishing mortality but increase mortality by natural predators. Our work suggests an insofar overlooked evolutionary mechanism by which size-selective harvesting can affect fishing and natural mortality of exploited fishV.S. was supported by a Leibniz-DAAD (German Academic Exchange Service) postdoctoral research fellowship (91632699), and he is now supported by the Spanish Ministry of Science and Innovation (Juan de la Cierva Incorporación Research Fellowship, IJC2018-035389-I). V.S. also acknowledges the Spanish government through the Severo Ochoa Centre of Excellence accreditation to the Department of Marine Renewable Resources, Institute of Marine Sciences (ICM-CSIC; CEX2019-000928-S). P.P.K. and P.R. received financial support from the German Research Foundation (DFG) under RO 4766/2-1 and under Germany’s Excellence Strategy (EXC 2002/1 Science of Intelligence, project 390523135)Peer reviewe

Die 4. COVID-19-Welle wurde durch fehlenden Impfschutz angestoßen: Was ist zu tun?

Neue Berechnungen auf der Basis eines mathematischen Ansteckungsmodells (es werden zwei verschiedene Szenarien mit unterschiedlichen Impfeffektivitäten angenommen) zeigen, dass eine effektive Erhöhung der Impfquote dringend notwendig ist: Bei 8 – 9 von 10 SARS-CoV-2-Ansteckungen ist mindestens eine ungeimpfte Person beteiligt – als Ansteckender, Angesteckter oder beides. Geimpfte spielen nur bei 5 – 6 von 10 Ansteckungen eine Rolle. Die röffentlichten Beispiele illustrieren, dass der Großteil der Neuinfektionen durch fehlenden Impfschutz verursacht wird, obwohl ungeimpfte Personen nur ca. 32 % der Bevölkerung ausmachen

Inferring country-specific import risk of diseases from the world air transportation network

Disease propagation between countries strongly depends on their effective distance, a measure derived from the world air transportation network (WAN). It reduces the complex spreading patterns of a pandemic to a wave-like propagation from the outbreak country, establishing a linear relationship to the arrival time of the unmitigated spread of a disease. However, in the early stages of an outbreak, what concerns decision-makers in countries is understanding the relative risk of active cases arriving in their country-essentially, the likelihood that an active case boarding an airplane at the outbreak location will reach them. While there are data-fitted models available to estimate these risks, accurate mechanistic, parameter-free models are still lacking. Therefore, we introduce the 'import risk' model in this study, which defines import probabilities using the effective-distance framework. The model assumes that airline passengers are distributed along the shortest path tree that starts at the outbreak's origin. In combination with a random walk, we account for all possible paths, thus inferring predominant connecting flights. Our model outperforms other mobility models, such as the radiation and gravity model with varying distance types, and it improves further if additional geographic information is included. The import risk model's precision increases for countries with stronger connections within the WAN, and it reveals a geographic distance dependence that implies a pull- rather than a push-dynamic in the distribution process.</p

Rank and relative performance of import risk estimation models.

The different import probability models are compared via their rank (A) and relative performance (B), with the highest values representing the best approach. The rank and relative performance are shown for each (black dots) of the six comparison measures (corr, logcorr, RMSE, logRMSE, cpc, τKendall) the box illustrates the interquartile range, the horizontal line the median and the red triangle the mean. The colors of the boxes illustrate the different distance measures in use. The outlier measure of the import risk models (I.R.) is the logRMSE, where the gravity models with effective distance are performing best. See Material and methods for definitions of comparison measures and Figs E, F in S1 Text for absolute and detailed relative performance.</p