Modeling the Heart as a Communication System

Electrical communication between cardiomyocytes can be perturbed during arrhythmia, but these perturbations are not captured by conventional electrocardiographic metrics. We developed a theoretical framework to quantify electrical communication using information theory metrics in 2-dimensional cell lattice models of cardiac excitation propagation. The time series generated by each cell was coarse-grained to 1 when excited or 0 when resting. The Shannon entropy for each cell was calculated from the time series during four clinically important heart rhythms: normal heartbeat, anatomical reentry, spiral reentry, and multiple reentry. We also used mutual information to perform spatial profiling of communication during these cardiac arrhythmias. We found that information sharing between cells was spatially heterogeneous. In addition, cardiac arrhythmia significantly impacted information sharing within the heart. Entropy localized the path of the drifting core of spiral reentry, which could be an optimal target of therapeutic ablation. We conclude that information theory metrics can quantitatively assess electrical communication among cardiomyocytes. The traditional concept of the heart as a functional syncytium sharing electrical information cannot predict altered entropy and information sharing during complex arrhythmia. Information theory metrics may find clinical application in the identification of rhythm-specific treatments which are currently unmet by traditional electrocardiographic techniques.Comment: 26 pages (including Appendix), 6 figures, 8 videos (not uploaded due to size limitation

The Missing Part of Seed Dispersal Networks: Structure and Robustness of Bat-Fruit Interactions

Mutualistic networks are crucial to the maintenance of ecosystem services. Unfortunately, what we know about seed dispersal networks is based only on bird-fruit interactions. Therefore, we aimed at filling part of this gap by investigating bat-fruit networks. It is known from population studies that: (i) some bat species depend more on fruits than others, and (ii) that some specialized frugivorous bats prefer particular plant genera. We tested whether those preferences affected the structure and robustness of the whole network and the functional roles of species. Nine bat-fruit datasets from the literature were analyzed and all networks showed lower complementary specialization (H2' = 0.37±0.10, mean ± SD) and similar nestedness (NODF = 0.56±0.12) than pollination networks. All networks were modular (M = 0.32±0.07), and had on average four cohesive subgroups (modules) of tightly connected bats and plants. The composition of those modules followed the genus-genus associations observed at population level (Artibeus-Ficus, Carollia-Piper, and Sturnira-Solanum), although a few of those plant genera were dispersed also by other bats. Bat-fruit networks showed high robustness to simulated cumulative removals of both bats (R = 0.55±0.10) and plants (R = 0.68±0.09). Primary frugivores interacted with a larger proportion of the plants available and also occupied more central positions; furthermore, their extinction caused larger changes in network structure. We conclude that bat-fruit networks are highly cohesive and robust mutualistic systems, in which redundancy is high within modules, although modules are complementary to each other. Dietary specialization seems to be an important structuring factor that affects the topology, the guild structure and functional roles in bat-fruit networks

Interaction parasites and the coevolution of mutualisms

Mutualismos são interações em que os parceiros se exploram reciprocamente com benefícios líquidos para ambos os indivíduos que interagem. Sistemas mutualistas multiespecíficos podem ser descritos como redes de interação, tais como aquelas formadas por sistemas de polinização, dispersão de sementes, estações de limpeza em ambientes recifais, formigas defensoras de plantas, mimetismo mülleriano e bactérias fixadoras de nitrogênio em raízes de plantas. As interações mutualísticas estão sujeitas à trapaça por indivíduos que, por meio de algum comportamento, alcançam o benefício oferecido pelo parceiro sem oferecer nada ou oferecer muito pouco em troca. No entanto, interações mutualísticas persistem apesar da existência de trapaceiros. Neste trabalho, mostro que os parasitas de interações mutualísticas, os trapaceiros, aumentam a resiliência das redes mutualísticas às perturbações mais rapidamente em redes aninhadas, redes tipicamente encontradas em mutualismos ricos em espécies. Portanto os efeitos combinados de trapaceiros, estrutura e dinâmica das redes mutualísticas podem ter implicações para a forma como a biodiversidade é mantida. Em seguida, estudo as condições em que flores tubulares, que sofrem maiores danos ao interagirem com ladrões de néctar, conseguem coexistir com flores planares, polinizadores e pilhadores por meio de efeitos indiretos da trapaça em seu sucesso reprodutivo. O roubo do néctar pode aumentar o sucesso de uma planta se as interações com pilhadores gerarem maior quantidade de polinização cruzada, aumentando assim o sucesso reprodutivo das plantas que interagem com ambos os visitantes florais. Tal resultado sugere uma nova fonte de manutenção da cooperação e da diversidade de estratégias por meio de efeitos não lineares das interações entre diferentes estratégias. Finalmente, estudo como as interações locais promovem a prevalência de mímicos (trapaceiros) em uma certa população na ausência de seus modelos. Mostro que presas que interagem localmente podem favorecer a predominância de mímicos e predadores que os evitam após algumas gerações e que uma distribuição não aleatória de indivíduos no espaço pode reforçar ainda mais este efeito inesperado de alopatria de modelo e mímicoMutualisms are interactions in which organisms of different species exploit each other with net benefits for both interacting individuals. Multispecific mutualistic system can be depicted as interaction networks, such as those formed by plant-pollinator interactions, dispersal systems, species interacting in cleaning stations in reef environments, protective ants in plants, müllerian mimicry, and nitrogen fixing bacteria on the roots of plants. Mutualistic interaction is subject to cheating by individuals who, by means of a diversity of behavioral strategies, achieve the benefit provided by the partner offering nothing or few in return. However, the mutualistic interactions persist despite the existence of cheaters. In this work I show that the parasites of mutualistic interactions increase the resilience of mutualistic networks to disturbances in nested networks, typically found in species-rich mutualisms. Therefore the joint effect of cheating, structure and dynamics of mutualistic networks have implications for how biodiversity is maintained. I subsequently study the conditions under which tubular flowers, which suffer stronger damages when interacting with nectar robbers, can coexist with planar flowers, pollinators, and robbers through indirect effects of cheating on their reproductive success. The theft of nectar may increase the success of a plant if its interactions with robbers generate higher degrees of cross-pollination, thus increasing the reproductive success of plants that interact with both floral visitors. This study suggests a new source of continued cooperation and diversity strategies through non-linear effects of the interactions between different strategies. Finally, I study how local interactions can promote the prevalence of mimic (the cheaters) in a given population in the absence of their models. I found that prey interacting locally may favor the predominance of mimic preys and avoid predators that, after a few generations and under a non-random distribution of individuals in space, can further strengthen this unexpected effect allopatry of the mimic and its mode

Interaction networks between frugivorous bats and plants : geographical variation and niche conservatism

Orientador: Paulo Roberto Guimarães JúniorDissertação (mestrado) - Universidade Estadual de Campinas, Instituto de BiologiaResumo: Os mutualismos são interações importantes do ponto de vista ecológico e evolutivo, uma vez que influenciam a sobrevivência de diversos organismos, podendo determinar a organização da comunidade. Dada a importância ecológica e evolutiva das interações mutualísticas, é importante estudar quais fatores influenciam a sua organização. Como organismos aparentados compartilham características fenotípicas similares, é comum que espécies filogeneticamente próximas interajam com um conjunto de espécies similar, levando a um padrão conhecido por conservação filogenética do nicho. Apesar de este padrão ocorrer com frequencia, condições bióticas e abióticas locais podem levar à variação geográfica nas interações. A presença ou ausência de potenciais parceiros mutualísticos também podem alterar o padrão local de interações de uma espécie. Estudando interações entre morcegos frugívoros e plantas, investiguei como os padrões de interação de espécies em dez redes mutualísticas variavam conforme a distribuição geográfica. Nesse sentido, duas hipóteses principais guiaram este trabalho: a hipótese da conservação de nicho das interações e a hipótese da variação geográfica das interações. Caracterizei o padrão de interação das espécies de morcegos nas redes por meio de diferentes métricas usadas no estudo de redes ecológicas. Analisei a conservação de nicho das interações e a variação geográfica de espécies de morcego que ocorreram em um maior número de redes. Espécies filogeneticamente próximas não tenderam a apresentar padrões de interação mais parecidos entre si. Algumas das espécies mais frequentes apresentaram características dos padrões de interação em gradiente ou mosaico ao longo de sua distribuição geográfica. Estes resultados evidenciam que algumas características do padrão de interação de morcegos frugívoros, como grau de generalismo e o grau padronizado dentro do módulo são conservadas pela filogenia. No entanto algumas espécies apresentaram o grau de generalismo e o papel em relação aos módulos da rede variando ao longo da geografia, seguindo previsões da teoria do mosaico geográfico coevolutivoAbstract: Mutualisms are important interactions to ecology and evolution because they affect the survivorship of many organisms and may shape community organization. Therefore, it is important to study which factors affect their organization. As related organisms share similar phenotypic traits, it is common that close phylogenetic species interact with a similar set of species, resulting in a pattern known as phylogenetic niche conservatism. Although this pattern frequently occurs, biotic and abiotic local conditions may lead to geographic variation of the interactions. The presence or absence of potential mutualistic partners may also change local interactions of a species. I studied interactions between frugivorous bats and plants and I investigated how species interactions patterns varied geographically across ten mutualistic networks. Two main hypotheses guided this work: the interaction niche conservatism hypothesis and the geographical variation hypothesis. I delineated the interaction patterns of bat species in the networks through different metrics used in ecological network studies. I analyzed the niche conservatism and geographical variation of bat species which occurred frequently in the networks. Related species did not tend to have similar patterns of interaction. Within their geographical range, some species vary markedly in their patterns of interaction and this variation often occurs as mosaics or, less frequently, as gradients. These results point out that some patterns of interaction of frugivorous bats, as the specialization degree and the standardized within-module degree, are conserved by phylogeny. However, some species showed variations through geography both in the specialization degree and in their role regarding the network modules, leading to a geographic mosaic of coevolution with potential implications to ecology and coevolutionMestradoEcologiaMestre em Ecologi

The Missing Part of Seed Dispersal Networks: Structure and Robustness of Bat-Fruit Interactions

Mutualistic networks are crucial to the maintenance of ecosystem services. Unfortunately, what we know about seed dispersal networks is based only on bird-fruit interactions. Therefore, we aimed at filling part of this gap by investigating bat-fruit networks. It is known from population studies that: (i) some bat species depend more on fruits than others, and (ii) that some specialized frugivorous bats prefer particular plant genera. We tested whether those preferences affected the structure and robustness of the whole network and the functional roles of species. Nine bat-fruit datasets from the literature were analyzed and all networks showed lower complementary specialization (H2\u27 = 0.37±0.10, mean ± SD) and similar nestedness (NODF = 0.56±0.12) than pollination networks. All networks were modular (M = 0.32±0.07), and had on average four cohesive subgroups (modules) of tightly connected bats and plants. The composition of those modules followed the genus-genus associations observed at population level (Artibeus-Ficus, Carollia-Piper, and Sturnira-Solanum), although a few of those plant genera were dispersed also by other bats. Bat-fruit networks showed high robustness to simulated cumulative removals of both bats (R = 0.55±0.10) and plants (R = 0.68±0.09). Primary frugivores interacted with a larger proportion of the plants available and also occupied more central positions; furthermore, their extinction caused larger changes in network structure. We conclude that bat-fruit networks are highly cohesive and robust mutualistic systems, in which redundancy is high within modules, although modules are complementary to each other. Dietary specialization seems to be an important structuring factor that affects the topology, the guild structure and functional roles in bat-fruit networks