Biology subjects

Ratikainen, I. I.

Publications and source records attributed to Ratikainen, I. I..

8 recordsLinked to original sources

bioRxiv2023

Eco-evolutionary dynamics of partially migratory metapopulations in spatially and seasonally varying environments

Predicting population responses to environmental changes requires understanding interactions among environmentally induced phenotypic variation, selection, demography and genetic variation, and thereby predicting eco-evolutionary dynamics emerging across diverse temporal and spatial scales. Partially migratory metapopulations (PMMPs), featuring seasonal coexistence of resident and migrant individuals across multiple spatially distinct subpopulations, have clear potential for complex spatio-seasonal eco-evolutionary dynamics through impacts of selection on migration on spatial population dynamics, and feedbacks resulting from ongoing micro-evolution. However, the key genetic and environmental conditions that maintain migratory polymorphisms, and eco-evolutionary dynamics of PMMPs under stochastic environmental variation and strong seasonal perturbations, have not yet been resolved. Accordingly, we present a general individual-based model that tracks eco-evolutionary dynamics in PMMPs inhabiting spatially structured, seasonally varying landscapes, with migration formulated as a quantitative genetic threshold trait. Our simulations show that such genetic and landscape structures, which commonly occur in nature, can readily produce a variety of stable partially migratory systems given diverse regimes of spatio-seasonal environmental variation. Typically, partial migration is maintained whenever sites differ in non-breeding season suitability resulting from variation in density-dependence, causing ideal free non-breeding distributions where residents and migrants occur with frequencies that generate similar survival probabilities. Yet, stable partial migration can also arise without any fixed differences in non-breeding season density-dependence among sites, and even without density-dependence at all, through risk-spreading given sufficiently large stochastic environmental fluctuations among sites and years. Finally, we show that local non-breeding season mortality events, as could result from extreme climatic events, can generate eco-evolutionary dynamics that ripple out to affect breeding and non-breeding season space use of subpopulations throughout the PMMP, on both short and longer timeframes. Such effects result from spatially divergent selection on both the occurrence and destinations of migration. Our model thus shows how facultative seasonal migration can act as a key mediator of eco-evolutionary dynamics in spatially and seasonally structured environments, providing key steps towards predicting responses of natural partially migratory populations to ongoing changes in spatio-seasonal patterns of environmental variation.

evolutionary biology↗

bioRxiv2023

Evolution during uncorrelated environmental fluctuations: bet-hedging or phenotypic plasticity?

Evolution in fluctuating environments is predicted to disfavor specialization and instead select for alternative strategies, such as phenotypic plasticity or possibly bet-hedging, depending on the accuracy of environmental cues and type of fluctuations. While these two alternatives are often contrasted in theoretical studies, their evolution are seldom studied together in empirical work. We used experimental evolution in the nematode worm Caenorhabditis remanei to simultaneously study the evolution of plasticity and bet-hedging in environments differing only in their temperature variability. We exposed worms for 30 generations to either fluctuating or slowly increasing temperature, these two environments had the same average temperature over evolutionary time. After experimental evolution, we scored size at sexual maturity and fitness in full siblings reared in two different temperatures, optimal 20{degrees}C and mildly stressful 25{degrees}C. Experimental evolution in the fluctuating environment resulted in the evolution of increased body size plasticity but not increased bet-hedging, compared to evolution in the slowly changing environment. Plasticity followed the temperature size rule as size decreased with increasing temperature and this plastic response was adaptive. In addition, we documented substantial standing genetic variation in body size, which represents a potential for further evolutionary change.

evolutionary biology↗

bioRxiv2022

The small-bat-in-summer paradigm: energetics and adaptive behavioural routines of bats investigated through a stochastic dynamic model

Strong seasonality at high latitudes represents a major challenge for many endotherms as they must balance survival and reproduction in an environment that varies widely in food availability and temperature. Being heterotherms, bats spend long cold winters in hibernation, avoiding the challenges faced by many animals. To avoid energetic mismatches caused by limited foraging time and stochastic weather conditions, bats can also employ this energy-saving state of torpor during summer to save accumulated energy reserves. However, at high latitudes small-bats-in-summer face a particular challenge: as nocturnal foragers they rely on the darkness of the night to avoid predators and/or interspecific competition, but for many the summer involves short nights of mostly twilight, and even a lack of true night at the northernmost distributions of some bat species. To investigate optimal individual behaviour across diurnal cycles, we constructed a stochastic dynamic model of bats living at high latitudes. Using a detailed parameterized model framework with values that are representative for our study system, we show that individual energetic reserves are a strong driver of day-time use of torpor and night-time foraging behaviour alike, with these linked effects being both temperature and photoperiod dependent. We further used the model framework to predict survival probabilities at five locations across a latitudinal gradient (60.1{degrees}N to 70.9{degrees}N), finding that photoperiod is the main limiting factor to bat species distributions. To verify the accuracy of our model results, we compared predictions for optimal decisions with our own empirical data collected on northern bats (Eptesicus nilssonii) from two latitudes in Norway. The similarities between our predictions and observations provide strong confirmation that this model framework incorporates the most important drivers of diurnal decision-making in bat physiology and behaviour. Our model findings regarding state-dependent decisions in bats should therefore contribute to the understanding of how bats cope with the summer challenges at high latitudes.

ecology↗

bioRxiv2020

Individual reversible plasticity as a genotype-level bet-hedging strategy

Reversible plasticity in phenotypic traits allows organisms to cope with environmental variation within lifetimes, but costs of plasticity may limit just how well the phenotype matches the environmental optimum. An additional adaptive advantage of plasticity might be to reduce fitness variance, or bet-hedging to maximize geometric (rather than simply arithmetic) mean fitness. Here we model the evolution of reaction norm slopes, with increasing costs as the slope or degree of plasticity increases. We find that greater investment in plasticity (i.e. steeper reaction norm slopes) is favoured in scenarios promoting bet-hedging as a response to multiplicative fitness accumulation (i.e. coarser environmental grains and fewer time steps prior to reproduction), because plasticity lowers fitness variance across environmental conditions. In contrast, in scenarios with finer environmental grain and many time steps prior to reproduction, bet-hedging plays less of a role and individual-level optimization favours evolution of shallower reaction norm slopes. We discuss contrasting predictions from this partitioning of the different adaptive causes of plasticity into short-term individual benefits versus long-term genotypic (bet-hedging) benefits under different costs of plasticity scenarios, thereby enhancing our understanding of the evolution of optimum levels of plasticity in examples from thermal physiology to advances in avian lay dates. Impact summaryPhenotypic plasticity is a key mechanism by which organisms cope with environmental change. Plasticity relies on the existence of some reliable environmental cue that allows organisms to infer current or future conditions, and adjust their traits in response to better match the environment. In contrast, when environmental fluctuations are unpredictable, bet-hedging favours lineages that persist by lowering their fitness variance, either among or within individuals. Plasticity and bet-hedging are therefore often considered to be alternative modes of adaptation to environmental change. However, we here make the point that plasticity also has the capacity to change an organisms variance in fitness across different environmental conditions, and could thus itself be part of - and not an alternative to - a bet-hedging strategy. We show that bet-hedging at the genotype level affects the optimal degree of plasticity that individuals use to track environmental fluctuations, because despite a reduction in expected fitness at the individual level, costly investment in the ability to be plastic also lowers variance in fitness. We also discuss alternative predictions that arise from scenarios with different types of costs of plasticity. Evolutionary bet-hedging and phenotypic plasticity are both topics experiencing a renewed surge of interest as researchers seek to better integrate different adaptations to ongoing rapid environmental change in a range of areas of literature within ecology and evolution, including behavioural ecology, evolutionary physiology and life-history theory. We believe that demonstrating an important novel link between these two mechanisms is of interest to research in many different fields, and opens new avenues for understanding organismal adaptation to environmental change.

evolutionary biology↗

bioRxiv2020

Variable environments select for short lifespan

The role of environmental variation in the evolution of lifespan is contested. Classic theory predicts that variable environments favour the evolution of long life, but recent theoretical work suggests that environmental variation can instead select for shorter lifespan when the changes are rapid relative to generation time. Here, we used experimental evolution to study how lifespan evolves in response to temperature variation. Genetically diverse populations of the outcrossing nematode Caenorhabditis remanei, were exposed to a novel, stressful temperature for 30 generations, under stable, gradually warming or fluctuating thermal regimes. We found the evolution of shorter lifespan in populations exposed to environments that fluctuated within the individuals lifetime, compared to populations evolving in stable warm temperature. While climate warming is predicted to increase environmental stochasticity, our results show that fast temperature cycles can rapidly select for short lifespan.

evolutionary biology↗

bioRxiv2019

Bet-hedging across generations can affect the evolution of variance-sensitive strategies within generations

In order to understand how organisms cope with ongoing changes in environmental variability it is important to consider all types of adaptations to environmental uncertainty on different time-scales. Conservative bet-hedging represents a long-term genotype-level strategy that maximizes lineage geometric mean fitness in stochastic environments by decreasing individual fitness variance, despite also lowering arithmetic mean fitness. Meanwhile, variance-prone (aka risk-prone) strategies produce greater variance in short-term payoffs because this increases expected arithmetic mean fitness if the relationship between payoffs and fitness is accelerating. Using two evolutionary simulation models, we investigate whether selection for such variance-prone strategies are counteracted by selection for bet-hedging that works to adaptively reduce fitness variance. We predict that variance-prone strategies will be favored in scenarios with more decision events per lifetime and when fitness accumulates additively rather than multiplicatively. In our model variance-proneness evolved in fine-grained environments (with lower correlations among individuals in energetic state and/or in payoffs when choosing the variable decision), and with larger numbers of independent decision events over which resources accumulate prior to selection. In contrast, geometric fitness accumulation caused by coarser environmental grain and fewer decision events prior to selection favors conservative bet-hedging via greater variance-aversion. We discuss examples of variance-sensitive strategies in optimal foraging, migration, life histories and cooperative breeding in light of these results concerning bet-hedging. By linking disparate fields of research studying adaptations to variable environments we should be more able to understand the effects in nature of human-induced rapid environmental change. Data depositionR code is available upon request.

evolutionary biology↗

bioRxiv2019

Generalists versus specialists in fluctuating environments: a bet-hedging perspective

Bet-hedging evolves in fluctuating environments because long-term genotype success is determined by geometric mean fitness across generations. However, specialist versus generalist strategies are usually considered in terms of arithmetic mean fitness benefits to individuals, as in habitat or foraging preferences. We model how environmental variability affects phenotypic variation within and among individuals to maximize either long-term arithmetic versus geometric mean fitness. For traits with additive fitness effects within lifetimes (e.g. foraging-related traits), genotypes of similar generalists or diversified specialists perform equally well. However, if fitness effects are multiplicative within lifetimes (e.g. sequential survival probabilities), generalist individuals are always favored, since geometric mean fitness favors greater within-individual phenotypic variation than arithmetic mean fitness does. Interestingly, this conservative bet-hedging effect outcompetes diversifying bet-hedging. These results link behavioral and ecological specialization and earlier models of bet-hedging, and thus apply to a range of natural phenomena from habitat choice to host specificity in parasites.\n\nImpact summaryWhich factors determine whether it is better to be a specialist or a generalist? Environmental fluctuations are becoming larger and more unpredictable across the globe as a result of human-induced rapid environmental change. A key challenge of evolutionary biology is therefore to understand how organisms adapt to such variation within and among generations, and currently represents a knowledge gap in evolutionary theory. Here we focus on how traits evolve when the (changing) environment determines the optimal value of a trait, so that the optimal trait value changes unpredictably over time. Our mathematical model investigates how much variation is optimal in a trait. We expect specialists (low within-individual trait variation) to be favored in stable environments, with generalists (high trait variation) favored in more variable environments. We show that the answer depends on whether we look from the point of view of the individual or all individuals of the same genotype. If an individual does well in the short term, but its offspring all experience a different environment and therefore do badly, the genotype as a whole is in trouble, and will not be favored in the long term. One solution to this problem could be to produce offspring with different trait values, to ensure that at least some of the offspring do well no matter the environmental conditions they grow up in. This \"dont put all your eggs in one basket\" diversification strategy is well-known in some organisms, but how helpful is it if there is also some within-individual (i.e. generalist) trait variation? By answering these questions under various environmental scenarios, we link together many different concepts in evolutionary ecology and animal behavior, increasing our understanding about how organisms may cope with the current changes in environmental conditions around the world.

evolutionary biology↗

bioRxiv2018

Short-term insurance versus long-term bet-hedging strategies as adaptations to variable environments

Understanding how organisms adapt to environmental variation is a key challenge of biology. Central to this are bet-hedging strategies that maximize geometric mean fitness across generations, either by being conservative or diversifying phenotypes. Theoretical models of bet-hedging and the multiplicative fitness effects of environmental variation across generations have traditionally assumed that environmental conditions are constant within lifetimes. However, behavioral ecology has revealed adaptive responses to additive fitness effects of environmental variation within lifetimes, either through insurance or risk-sensitive strategies. Here we explore whether the effects of adaptive insurance interact with the evolution of bet-hedging by varying the position and skew of fitness functions within and between lifetimes. When insurance causes the optimal phenotype to shift from the peak to down the less steeply decreasing side of the fitness function, then conservative bet-hedging does not generally evolve on top of this, even if diversifying bet-hedging can. Canalization to reduce phenotypic variation within a lifetime is almost always favored, except when the tails of the fitness function are steeply convex and produce a novel risk-sensitive increase in phenotypic variance akin to diversifying bet-hedging. Importantly, using skewed fitness functions, we provide the first example of how conservative and diversifying bet-hedging strategies might coexist.

evolutionary biology↗