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1.
J Math Biol ; 88(2): 24, 2024 02 02.
Artigo em Inglês | MEDLINE | ID: mdl-38308102

RESUMO

The evolution of mutualism between host and symbiont communities plays an essential role in maintaining ecosystem function and should therefore have a profound effect on their range expansion dynamics. In particular, the presence of mutualistic symbionts at the leading edge of a host-symbiont community should enhance its propagation in space. We develop a theoretical framework that captures the eco-evolutionary dynamics of host-symbiont communities, to investigate how the evolution of resource exchange may shape community structure during range expansion. We consider a community with symbionts that are mutualistic or parasitic to various degrees, where parasitic symbionts receive the same amount of resource from the host as mutualistic symbionts, but at a lower cost. The selective advantage of parasitic symbionts over mutualistic ones is increased with resource availability (i.e. with host density), promoting mutualism at the range edges, where host density is low, and parasitism at the population core, where host density is higher. This spatial selection also influences the speed of spread. We find that the host growth rate (which depends on the average benefit provided by the symbionts) is maximal at the range edges, where symbionts are more mutualistic, and that host-symbiont communities with high symbiont density at their core (e.g. resulting from more mutualistic hosts) spread faster into new territories. These results indicate that the expansion of host-symbiont communities is pulled by the hosts but pushed by the symbionts, in a unique push-pull dynamic where both the host and symbionts are active and tightly-linked players.


Assuntos
Ecossistema , Parasitos , Animais , Simbiose , Reprodução
2.
Glob Chang Biol ; 29(12): 3347-3363, 2023 06.
Artigo em Inglês | MEDLINE | ID: mdl-37021593

RESUMO

Human activity is leading to changes in the mean and variability of climatic parameters in most locations around the world. The changing mean has received considerable attention from scientists and climate policy makers. However, recent work indicates that the changing variability, that is, the amplitude and the temporal autocorrelation of deviations from the mean, may have greater and more imminent impact on ecosystems. In this paper, we demonstrate that changes in climate variability alone could drive cyclic predator-prey ecosystems to extinction via so-called phase-tipping (P-tipping), a new type of instability that occurs only from certain phases of the predator-prey cycle. We construct a mathematical model of a variable climate and couple it to two self-oscillating paradigmatic predator-prey models. Most importantly, we combine realistic parameter values for the Canada lynx and snowshoe hare with actual climate data from the boreal forest. In this way, we demonstrate that critically important species in the boreal forest have increased likelihood of P-tipping to extinction under predicted changes in climate variability, and are most vulnerable during stages of the cycle when the predator population is near its maximum. Furthermore, our analysis reveals that stochastic resonance is the underlying mechanism for the increased likelihood of P-tipping to extinction.


Assuntos
Lebres , Lynx , Animais , Humanos , Ecossistema , Dinâmica Populacional , Modelos Teóricos , Comportamento Predatório
3.
J Theor Biol ; 534: 110958, 2022 02 07.
Artigo em Inglês | MEDLINE | ID: mdl-34748733

RESUMO

Many studies examine how the landscape affects memory-informed movement patterns, but very few examine how memory-informed foragers influence the landscape. This reverse relationship is an important factor in preventing the continued decline of many ecosystem services. We investigate this question in the context of crop pollination services by wild bees, a critical ecosystem service that is in steep decline. Many studies suggest that adding wild flower patches near crops can result in higher crop pollination services, but specific advice pertaining to the optimal location and density of these wild flower patches is lacking, as well as any estimate of the expected change in crop pollination services. In this work, we seek to understand what is the optimal placement of a flower patch relative to a single crop field, during crop bloom and considering spatial factors alone. We develop an individual based model of memory-based foraging by bumble bees to simulate bee movement from a single nest while the crop is in bloom, and measure the resulting crop pollination services. We consider a single crop field enhanced with a wild flower patch in a variable location, and measure crop flower visitation over the course of a single day. We analyze the pollination intensity and spatial distribution of flower visits to determine optimal wild flower patch placement for an isolated crop field. We find that the spatial arrangement of crop and wild flower patch have a significant effect on the number of crop flower visits, and that these effects arise from the memory-informed foraging pattern. The most effective planting locations are either in the centre of the crop field or on the far side of the crop field, away from the single bumble bee nest.


Assuntos
Ecossistema , Polinização , Animais , Abelhas , Produtos Agrícolas , Flores
4.
J Theor Biol ; 531: 110859, 2021 12 21.
Artigo em Inglês | MEDLINE | ID: mdl-34389360

RESUMO

Arbuscular mycorrhizal (AM) fungi play a key role in determining ecosystem functionality. Understanding how diversity in the fungal community affects plant productivity is therefore an important question in ecology. Current research has focused on understanding the role of functional complementarity in the fungal community when the host plant faces multiple stress factors. Fewer studies, however, have investigated how variation in traits affecting nutrient exchange can impact the plant growth dynamics, even in the absence of environmental stressors. Combining experimental data and a mathematical model based on ordinary differential equations, we investigate the role played by carbon sink strength on plant productivity. We simulate and measure plant growth over time when the plant is associated with two fungal isolates with different carbon sink strength, and when the plant is in pairwise association with each of the isolates alone. Overall, our theoretical as well as our experimental results show that co-inoculation with fungi with different carbon sink strength can induce positive non-additive effects (or synergistic effects) in plant productivity. Fungi with high carbon sink strength are able to quickly establish a fungal community and increase the nutrient supply to the plant, with a consequent positive impact on plant growth rate. On the other side, fungi with low carbon sink strength inflict lower carbon costs to the host plant, and support maximal plant productivity once plant biomass is large. As AM fungi are widely used as organic fertilizers worldwide, our findings have important implications for restoration ecology and agricultural management.


Assuntos
Micorrizas , Biomassa , Carbono , Sequestro de Carbono , Ecossistema , Fungos , Raízes de Plantas , Solo , Microbiologia do Solo
5.
Bull Math Biol ; 83(10): 107, 2021 09 04.
Artigo em Inglês | MEDLINE | ID: mdl-34482488

RESUMO

As many ecosystems worldwide are in peril, efforts to manage them sustainably require scientific advice. While numerous researchers around the world use a great variety of models to understand ecological dynamics and their responses to disturbances, only a small fraction of these models are ever used to inform ecosystem management. There seems to be a perception that ecological models are not useful for management, even though mathematical models are indispensable in many other fields. We were curious about this mismatch, its roots, and potential ways to overcome it. We searched the literature on recommendations and best practices for how to make ecological models useful to the management of ecosystems and we searched for 'success stories' from the past. We selected and examined several cases where models were instrumental in ecosystem management. We documented their success and asked whether and to what extent they followed recommended best practices. We found that there is not a unique way to conduct a research project that is useful in management decisions. While research is more likely to have impact when conducted with many stakeholders involved and specific to a situation for which data are available, there are great examples of small groups or individuals conducting highly influential research even in the absence of detailed data. We put the question of modelling for ecosystem management into a socio-economic and national context and give our perspectives on how the discipline could move forward.


Assuntos
Ecossistema , Conceitos Matemáticos , Humanos , Modelos Teóricos
6.
J Math Biol ; 83(5): 50, 2021 10 12.
Artigo em Inglês | MEDLINE | ID: mdl-34636970

RESUMO

Bumble bees provide valuable pollination services to crops around the world. However, their populations are declining in intensively farmed landscapes. Understanding the dispersal behaviour of these bees is a key step in determining how agricultural landscapes can best be enhanced for bumble bee survival. Here we develop a partial integro-differential equation model to predict the spatial distribution of foraging bumble bees in dynamic heterogeneous landscapes. In our model, the foraging population is divided into two subpopulations, one engaged in an intensive search mode (modeled by diffusion) and the other engaged in an extensive search mode (modeled by advection). Our model considers the effects of resource-dependent switching rates between movement modes, resource depletion, central-place foraging behaviour, and memory. We use our model to investigate how crop pollination services are affected by wildflower enhancements. We find that planting wildflowers such that the crop is located in between the wildflowers and the nest site can benefit crop pollination in two different scenarios. If the bees do not have a strong preference for wildflowers, a small or low density wildflower patch is beneficial. If, on the other hand, the bees strongly prefer the wildflowers, then a large or high density wildflower patch is beneficial. The increase of the crop pollination services in the later scenario is of remarkable magnitude.


Assuntos
Agricultura , Polinização , Animais , Abelhas , Plantas
7.
JAAPA ; 34(4): 38-39, 2021 Apr 01.
Artigo em Inglês | MEDLINE | ID: mdl-33735139

RESUMO

ABSTRACT: Purple urine bag syndrome most commonly occurs in chronically catheterized patients who develop urinary tract infections. Although this syndrome can cause concern because of the appearance of abnormally colored urine, it is relatively benign. However, if left untreated, purple urine bag syndrome can lead to complications associated with progressive lower urinary tract infections. This article discusses how to identify, diagnose, and treat purple urine bag syndrome.


Assuntos
Infecções Urinárias , Humanos , Síndrome , Infecções Urinárias/diagnóstico , Infecções Urinárias/tratamento farmacológico
8.
Proc Biol Sci ; 287(1923): 20192312, 2020 03 25.
Artigo em Inglês | MEDLINE | ID: mdl-32208836

RESUMO

Biodiversity is an important component of healthy ecosystems, and thus understanding the mechanisms behind species coexistence is critical in ecology and conservation biology. In particular, few studies have focused on the dynamics resulting from the co-occurrence of mutualistic and competitive interactions within a group of species. Here we build a mathematical model to study the dynamics of a guild of competitors who are also engaged in mutualistic interactions with a common partner. We show that coexistence as well as competitive exclusion can occur depending on the competition strength and on strength of the mutualistic interactions, and we formulate concrete criteria for predicting invasion success of an alien mutualist based on propagule pressure, alien traits (such as its resource exchange ability) and composition of the recipient community. We find that intra guild diversity promotes the coexistence of species that would otherwise competitively exclude each other, and makes a guild less vulnerable to invasion. Our results can serve as a useful framework to predict the consequences of species manipulation in mutualistic communities.


Assuntos
Biodiversidade , Espécies Introduzidas , Simbiose , Ecologia , Ecossistema , Dinâmica Populacional
9.
Bull Math Biol ; 82(1): 14, 2020 01 14.
Artigo em Inglês | MEDLINE | ID: mdl-31932981

RESUMO

During an epidemic, the interplay of disease and opinion dynamics can lead to outcomes that are different from those predicted based on disease dynamics alone. Opinions and the behaviours they elicit are complex, so modelling them requires a measure of abstraction and simplification. Here, we develop a differential equation model that couples SIR-type disease dynamics with opinion dynamics. We assume a spectrum of opinions that change based on current levels of infection as well as interactions that to some extent amplify the opinions of like-minded individuals. Susceptibility to infection is based on the level of prophylaxis (disease avoidance) that an opinion engenders. In this setting, we observe how the severity of an epidemic is influenced by the distribution of opinions at disease introduction, the relative rates of opinion and disease dynamics, and the amount of opinion amplification. Some insight is gained by considering how the effective reproduction number is influenced by the combination of opinion and disease dynamics.


Assuntos
Epidemias , Comportamentos Relacionados com a Saúde , Modelos Biológicos , Atitude , Atitude Frente a Saúde , Número Básico de Reprodução/estatística & dados numéricos , Doenças Transmissíveis/epidemiologia , Doenças Transmissíveis/psicologia , Doenças Transmissíveis/transmissão , Simulação por Computador , Suscetibilidade a Doenças , Epidemias/prevenção & controle , Epidemias/estatística & dados numéricos , Humanos , Conceitos Matemáticos , Modelos Psicológicos
10.
Bull Math Biol ; 81(11): 4484-4517, 2019 11.
Artigo em Inglês | MEDLINE | ID: mdl-29541997

RESUMO

We present an analysis of an avian flu model that yields insight into the roles of different transmission routes in the recurrence of avian influenza epidemics. Recent modelling work suggests that the outbreak periodicity of the disease is mainly determined by the environmental transmission rate. This conclusion, however, is based on a modelling study that only considers a weak between-host transmission rate. We develop an approximate model for stochastic avian flu epidemics, which allows us to determine the relative contribution of environmental and direct transmission routes to the periodicity and intensity of outbreaks over the full range of plausible parameter values for transmission. Our approximate model reveals that epidemic recurrence is chiefly governed by the product of a rotation and a slowly varying standard Ornstein-Uhlenbeck process (i.e. mean-reverting process). The intrinsic frequency of the damped deterministic version of the system predicts the dominant period of outbreaks. We show that the typical periodicity of major avian flu outbreaks can be explained in terms of either or both types of transmission and that the typical amplitude of epidemics is highly sensitive to the direct transmission rate.


Assuntos
Epidemias/veterinária , Influenza Aviária/epidemiologia , Influenza Aviária/transmissão , Modelos Biológicos , Animais , Aves/virologia , Simulação por Computador , Surtos de Doenças/estatística & dados numéricos , Surtos de Doenças/veterinária , Reservatórios de Doenças , Microbiologia Ambiental , Epidemias/estatística & dados numéricos , Interações entre Hospedeiro e Microrganismos , Especificidade de Hospedeiro , Humanos , Influenza Humana/epidemiologia , Influenza Humana/transmissão , Modelos Lineares , Conceitos Matemáticos , Recidiva , Processos Estocásticos
11.
J Math Biol ; 79(6-7): 2133-2155, 2019 12.
Artigo em Inglês | MEDLINE | ID: mdl-31520107

RESUMO

Many real populations exhibit stochastic behaviour that appears to have some periodicity. In terms of populations, these time series can occur as limit cycles that arise through seasonal variation of parameters such as, e.g., disease transmission rate. The general mathematical context is that of a stochastic differential system with periodic parametric forcing whose solution is a stochastically perturbed limit cycle. Earlier work identified the power spectral density (PSD) features of these fluctuations by computation of the autocorrelation function of the stochastic process and its transform. Here, we present an alternative analysis which shows that the structure of the fluctuations around the limit cycle is analogous to that of fluctuations about a fixed point. Furthermore, we show that these fluctuations can be expressed, approximately, as a factorization which reveals the combined frequencies of the limit cycle and the stochastic perturbation. This result, based on a new limit theorem near a Hopf point, yields an understanding of the previously found features of the PSD. Further insights are obtained from the corresponding stochastic equations for phase and amplitude.


Assuntos
Modelos Biológicos , Estações do Ano , Processos Estocásticos , Distribuição Animal , Animais , Simulação por Computador , Transmissão de Doença Infecciosa/estatística & dados numéricos , Cadeia Alimentar , Humanos , Dinâmica não Linear , Comportamento Predatório
12.
Ecol Lett ; 20(8): 1074-1092, 2017 08.
Artigo em Inglês | MEDLINE | ID: mdl-28633194

RESUMO

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer-resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.


Assuntos
Biodiversidade , Evolução Biológica , Animais , Ecossistema , Densidade Demográfica , Dinâmica Populacional , Comportamento Predatório
13.
Am Nat ; 188(5): 539-553, 2016 Nov.
Artigo em Inglês | MEDLINE | ID: mdl-27788349

RESUMO

The functional response of some predator species changes from a pattern characteristic for a generalist to that for a specialist according to seasonally varying prey availability. Current theory does not address the dynamic consequences of this phenomenon. Since season length correlates strongly with altitude and latitude and is predicted to change under future climate scenarios, including this phenomenon in theoretical models seems essential for correct prediction of future ecosystem dynamics. We develop and analyze a two-season model for the great horned owl (Bubo virginialis) and snowshoe hare (Lepus americanus). These species form a predator-prey system in which the generalist to specialist shift in predation pattern has been documented empirically. We study the qualitative behavior of this predator-prey model community as summer season length changes. We find that relatively small changes in summer season length can have a profound impact on the system. In particular, when the predator has sufficient alternative resources available during the summer season, it can drive the prey to extinction, there can be coexisting stable states, and there can be stable large-amplitude limit cycles coexisting with a stable steady state. Our results illustrate that the impacts of global change on local ecosystems can be driven by internal system dynamics and can potentially have catastrophic consequences.


Assuntos
Mudança Climática , Lebres , Comportamento Predatório , Estrigiformes , Animais , Ecossistema , Modelos Biológicos , Dinâmica Populacional , Estações do Ano
14.
J Math Sociol ; 40(4): 207-218, 2016.
Artigo em Inglês | MEDLINE | ID: mdl-28529381

RESUMO

Opinions are rarely binary; they can be held with different degrees of conviction, and this expanded attitude spectrum can affect the influence one opinion has on others. Our goal is to understand how different aspects of influence lead to recognizable spatio-temporal patterns of opinions and their strengths. To do this, we introduce a stochastic spatial agent-based model of opinion dynamics that includes a spectrum of opinion strengths and various possible rules for how the opinion strength of one individual affects the influence that this individual has on others. Through simulations, we find that even a small amount of amplification of opinion strength through interaction with like-minded neighbors can tip the scales in favor of polarization and deadlock.

15.
Reprod Biol Endocrinol ; 10: 104, 2012 Dec 05.
Artigo em Inglês | MEDLINE | ID: mdl-23216940

RESUMO

BACKGROUND: Spermatic cord torsion can lead to testis ischemia (I) and subsequent ischemia-reperfusion (I/R) causing germ cell-specific apoptosis. Previously, we demonstrated that the hypoxia-inducible factor-1 (HIF-1) transcription factor, a key regulator of physiological responses to hypoxia, is abundant in Leydig cells in normoxic and ischemic testes. We hypothesize that testicular HIF-1 activates the expression of antiapoptotic target genes to protect Leydig cells from apoptosis. In silico analysis of testis genes containing a consensus hypoxia response element (HRE, 5'-RCGTG-3') identified myeloid cell leukemia-1 (Mcl-1) as a potential HIF-1 target gene. The purpose of this study was to determine whether HIF-1 shows DNA-binding activity in normoxic and ischemic testes and whether Mcl-1 is a target gene of testicular HIF-1. METHODS: The testicular HIF-1 DNA-binding capacity was analyzed in vitro using a quantitative enzyme-linked immunosorbent assay (ELISA) and electrophoretic mobility shift assays (EMSA). MCL-1 protein expression was evaluated by immunoblot analysis and immunohistochemistry. The binding of testicular HIF-1 to the Mcl-1 gene was examined via chromatin immunoprecipitation (ChIP) analysis. RESULTS: The ELISA and EMSA assays demonstrated that testicular HIF-1 from normoxic and ischemic testes binds DNA equally strongly, suggesting physiological roles for HIF-1 in the normoxic testis, unlike most tissues in which HIF-1 is degraded under normoxic conditions and is only activated by hypoxia. MCL-1 protein was determined to be abundant in both normoxic and ischemic testes and expressed in Leydig cells. In a pattern identical to that of HIF-1 expression, the steady-state levels of MCL-1 were not significantly affected by I or I/R and MCL-1 co-localized with HIF-1α in Leydig cells. Chromatin immunoprecipitation (ChIP) analysis using a HIF-1 antibody revealed sequences enriched for the Mcl-1 promoter. CONCLUSIONS: The results demonstrated that, unlike what is observed in most tissues, HIF-1 displays DNA-binding activity in both normoxic and ischemic testes, and Mcl-1 may be a key target gene of testicular HIF-1 with potential roles in the antiapoptotic protection of Leydig cells.


Assuntos
Fator 1 Induzível por Hipóxia/análise , Fator 1 Induzível por Hipóxia/metabolismo , Proteínas Proto-Oncogênicas c-bcl-2/genética , Testículo/química , Animais , Apoptose , Hipóxia Celular , Imunoprecipitação da Cromatina , DNA/metabolismo , Subunidade alfa do Fator 1 Induzível por Hipóxia/análise , Immunoblotting , Imuno-Histoquímica , Isquemia/metabolismo , Células Intersticiais do Testículo/química , Masculino , Proteína de Sequência 1 de Leucemia de Células Mieloides , Proteínas Proto-Oncogênicas c-bcl-2/análise , Ratos , Ratos Sprague-Dawley , Testículo/irrigação sanguínea
16.
Proc Math Phys Eng Sci ; 477(2254): 20210059, 2021 Oct.
Artigo em Inglês | MEDLINE | ID: mdl-35153584

RESUMO

We identify the phase of a cycle as a new critical factor for tipping points (critical transitions) in cyclic systems subject to time-varying external conditions. As an example, we consider how contemporary climate variability induces tipping from a predator-prey cycle to extinction in two paradigmatic predator-prey models with an Allee effect. Our analysis of these examples uncovers a counterintuitive behaviour, which we call phase tipping or P-tipping, where tipping to extinction occurs only from certain phases of the cycle. To explain this behaviour, we combine global dynamics with set theory and introduce the concept of partial basin instability for attracting limit cycles. This concept provides a general framework to analyse and identify easily testable criteria for the occurrence of phase tipping in externally forced systems, and can be extended to more complicated attractors.

17.
Phys Rev E ; 98(2-1): 022310, 2018 Aug.
Artigo em Inglês | MEDLINE | ID: mdl-30253497

RESUMO

Spatially situated opinions that can be held with different degrees of conviction lead to spatiotemporal patterns such as clustering (homophily), polarization, and deadlock. Our goal is to understand how sensitive these patterns are to changes in the local nature of interactions. We introduce two different mixing mechanisms, spatial relocation and nonlocal interaction ("telephoning"), to an earlier fully spatial model (no mixing). Interestingly, the mechanisms that create deadlock in the fully spatial model have the opposite effect when there is a sufficient amount of mixing. With telephoning, not only is polarization and deadlock broken up, but consensus is hastened. The effects of mixing by relocation are even more pronounced. Further insight into these dynamics is obtained for selected parameter regimes via comparison to the mean-field differential equations.

18.
J R Soc Interface ; 14(126)2017 01.
Artigo em Inglês | MEDLINE | ID: mdl-28123097

RESUMO

The containment of genetically modified (GM) pollen is an issue of significant concern for many countries. For crops that are bee-pollinated, model predictions of outcrossing rates depend on the movement hypothesis used for the pollinators. Previous work studying pollen spread by honeybees, the most important pollinator worldwide, was based on the assumption that honeybee movement can be well approximated by Brownian motion. A number of recent studies, however, suggest that pollinating insects such as bees perform Lévy flights in their search for food. Such flight patterns yield much larger rates of spread, and so the Brownian motion assumption might significantly underestimate the risk associated with GM pollen outcrossing in conventional crops. In this work, we propose a mechanistic model for pollen dispersal in which the bees perform truncated Lévy flights. This assumption leads to a fractional-order diffusion model for pollen that can be tuned to model motion ranging from pure Brownian to pure Lévy. We parametrize our new model by taking the same pollen dispersal dataset used in Brownian motion modelling studies. By numerically solving the model equations, we show that the isolation distances required to keep outcrossing levels below a certain threshold are substantially increased by comparison with the original predictions, suggesting that isolation distances may need to be much larger than originally thought.


Assuntos
Abelhas/fisiologia , Modelos Biológicos , Plantas Geneticamente Modificadas/fisiologia , Pólen/fisiologia , Polinização/fisiologia , Animais
19.
J Homosex ; 45(1): 65-91, 2003.
Artigo em Inglês | MEDLINE | ID: mdl-14567654

RESUMO

Two samples of committed gay and lesbian cohabiting couples and two samples of married couples (couples in which the woman presented the conflict issue to the man, and couples in which the man presented the conflict issue to the woman) engaged in three conversations: (1) an events of the day conversation (after being apart for at least 8 hours), (2) a conflict resolution conversation, and (3) a pleasant topic conversation. The observational data were coded with a system that categorized specific affects displayed. Data were weighted and two time-series created, one for the husband and one of the wife. The time series were modeled with nonlinear difference equations (Cook et al., 1995), and parameters were estimated that indexed uninfluenced steady state, influenced steady state, emotional inertia, repair effectiveness and threshold, and the power of positive and negative affect of one partner to affect the other partner.


Assuntos
Conflito Psicológico , Homossexualidade Feminina/psicologia , Homossexualidade Masculina/psicologia , Relações Interpessoais , Modelos Psicológicos , Cônjuges/psicologia , Adulto , Feminino , Heterossexualidade , Humanos , Masculino , Estado Civil
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