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

2019 ◽  
Vol 286 (1916) ◽  
pp. 20192070 ◽  
Author(s):  
Thomas R. Haaland ◽  
Jonathan Wright ◽  
Irja I. Ratikainen

In order to understand how organisms cope with ongoing changes in environmental variability, it is necessary to consider multiple adaptations to environmental uncertainty on different time scales. Conservative bet-hedging (CBH) represents a long-term genotype-level strategy maximizing 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 evolutionary simulation models, we investigate whether selection for such variance-prone strategies is counteracted by selection for bet-hedging that works to adaptively reduce fitness variance. In our model, variance proneness evolves in fine-grained environments (lower correlations among individuals in energetic state and/or payoffs), and with larger numbers of independent decision events over which resources accumulate prior to selection. Conversely, multiplicative fitness accumulation, caused by coarser environmental grain and fewer decision events selection, favours CBH via greater variance aversion. We discuss examples of variance-sensitive strategies in optimal foraging, migration, life histories and cooperative breeding using this bet-hedging perspective. By linking disparate fields of research studying adaptations to variable environments, we should be better able to understand effects of human-induced rapid environmental change.

2019 ◽  
Author(s):  
Thomas R. Haaland ◽  
Jonathan Wright ◽  
Irja I. Ratikainen

AbstractIn 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.


2017 ◽  

AbstractIn evolutionary biology, bet-hedging refers to a strategy that reduces the variance of reproductive success at the cost of reduced mean reproductive success. In unpredictably fluctuating environments, bet-hedgers benefit from higher geometric mean fitness despite having lower arithmetic mean fitness than their specialist competitors. We examine the extent to which sexual reproduction can be considered a type of bet-hedging, by clarifying past arguments, examining parallels and differences to evolutionary games, and by presenting a simple model examining geometric and arithmetic mean payoffs of sexual and asexual reproduction. Sex typically has lower arithmetic mean fitness than asex, while the geometric mean fitness can be higher if sexually produced offspring are not identical. However, asexual individuals that are heterozygotes can gain conservative bet-hedging benefits of similar magnitude while avoiding the costs of sex. This highlights that bet-hedging always has to be specified relative to the payoff structure of relevant competitors. It also makes it unlikely that sex, at least when associated with significant male production, evolves solely based on bet-hedging in the context of frequently and repeatedly occupied environmental states. Future work could usefully consider bet-hedging in open-ended evolutionary scenarios with de novo mutations.


2019 ◽  
Author(s):  
Thomas Ray Haaland ◽  
Jonathan Wright ◽  
Irja Ida Ratikainen

AbstractBet-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.Impact 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 “don’t 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.


2015 ◽  
Vol 282 (1798) ◽  
pp. 20141525 ◽  
Author(s):  
Francisco Garcia-Gonzalez ◽  
Yukio Yasui ◽  
Jonathan P. Evans

Polyandry (female multiple mating) has profound evolutionary and ecological implications. Despite considerable work devoted to understanding why females mate multiply, we currently lack convincing empirical evidence to explain the adaptive value of polyandry. Here, we provide a direct test of the controversial idea that bet-hedging functions as a risk-spreading strategy that yields multi-generational fitness benefits to polyandrous females. Unfortunately, testing this hypothesis is far from trivial, and the empirical comparison of the across-generations fitness payoffs of a polyandrous (bet hedger) versus a monandrous (non-bet hedger) strategy has never been accomplished because of numerous experimental constraints presented by most ‘model’ species. In this study, we take advantage of the extraordinary tractability and versatility of a marine broadcast spawning invertebrate to overcome these challenges. We are able to simulate multi-generational (geometric mean) fitness among individual females assigned simultaneously to a polyandrous and monandrous mating strategy. Our approaches, which separate and account for the effects of sexual selection and pure bet-hedging scenarios, reveal that bet-hedging, in addition to sexual selection, can enhance evolutionary fitness in multiply mated females. In addition to offering a tractable experimental approach for addressing bet-hedging theory, our study provides key insights into the evolutionary ecology of sexual interactions.


2014 ◽  
Vol 281 (1787) ◽  
pp. 20140706 ◽  
Author(s):  
Jeffrey K. Graham ◽  
Myron L. Smith ◽  
Andrew M. Simons

All organisms are faced with environmental uncertainty. Bet-hedging theory expects unpredictable selection to result in the evolution of traits that maximize the geometric-mean fitness even though such traits appear to be detrimental over the shorter term. Despite the centrality of fitness measures to evolutionary analysis, no direct test of the geometric-mean fitness principle exists. Here, we directly distinguish between predictions of competing fitness maximization principles by testing Cohen's 1966 classic bet-hedging model using the fungus Neurospora crassa . The simple prediction is that propagule dormancy will evolve in proportion to the frequency of ‘bad’ years, whereas the prediction of the alternative arithmetic-mean principle is the evolution of zero dormancy as long as the expectation of a bad year is less than 0.5. Ascospore dormancy fraction in N. crassa was allowed to evolve under five experimental selection regimes that differed in the frequency of unpredictable ‘bad years’. Results were consistent with bet-hedging theory: final dormancy fraction in 12 genetic lineages across 88 independently evolving samples was proportional to the frequency of bad years, and evolved both upwards and downwards as predicted from a range of starting dormancy fractions. These findings suggest that selection results in adaptation to variable rather than to expected environments.


2012 ◽  
Vol E95-B (2) ◽  
pp. 647-650
Author(s):  
Ning WANG ◽  
Julian CHENG ◽  
Chintha TELLAMBURA

Author(s):  
Annie Jonsson

AbstractMost animal species have a complex life cycle (CLC) with metamorphosis. It is thus of interest to examine possible benefits of such life histories. The prevailing view is that CLC represents an adaptation for genetic decoupling of juvenile and adult traits, thereby allowing life stages to respond independently to different selective forces. Here I propose an additional potential advantage of CLCs that is, decreased variance in population growth rate due to habitat separation of life stages. Habitat separation of pre- and post-metamorphic stages means that the stages will experience different regimes of environmental variability. This is in contrast to species with simple life cycles (SLC) whose life stages often occupy one and the same habitat. The correlation in the fluctuations of the vital rates of life stages is therefore likely to be weaker in complex than in simple life cycles. By a theoretical framework using an analytical approach, I have (1) derived the relative advantage, in terms of long-run growth rate, of CLC over SLC phenotypes for a broad spectrum of life histories, and (2) explored which life histories that benefit most by a CLC, that is avoid correlation in vital rates between life stages. The direction and magnitude of gain depended on life history type and fluctuating vital rate. One implication of our study is that species with CLCs should, on average, be more robust to increased environmental variability caused by global warming than species with SLCs.


2003 ◽  
Vol 1 (3) ◽  
pp. 109-115 ◽  
Author(s):  
Thomas F. Clasen ◽  
Andrew Bastable

Paired water samples were collected and analysed for thermotolerant coliforms (TTC) from 20 sources (17 developed or rehabilitated by Oxfam and 3 others) and from the stored household water supplies of 100 households (5 from each source) in 13 towns and villages in the Kailahun District of Sierra Leone. In addition, the female head of the 85 households drawing water from Oxfam improved sources was interviewed and information recorded on demographics, hygiene instruction and practices, sanitation facilities and water collection and storage practices. At the non-improved sources, the arithmetic mean TTC load was 407/100 ml at the point of distribution, rising to a mean count of 882/100 ml at the household level. Water from the improved sources met WHO guidelines, with no faecal contamination. At the household level, however, even this safe water was subject to frequent and extensive faecal contamination; 92.9% of stored household samples contained some level of TTC, 76.5% contained more than the 10 TTC per 100 ml threshold set by the Sphere Project for emergency conditions. The arithmetic mean TTC count for all samples from the sampled households was 244 TTC per 100 ml (geometric mean was 77). These results are consistent with other studies that demonstrate substantial levels of faecal contamination of even safe water during collection, storage and access in the home. They point to the need to extend drinking water quality beyond the point of distribution to the point of consumption. The options for such extended protection, including improved collection and storage methods and household-based water treatment, are discussed.


2015 ◽  
Vol 22 (2) ◽  
pp. 194-209 ◽  
Author(s):  
Dejian YU ◽  
Wenyu ZHANG ◽  
George HUANG

Dual hesitant fuzzy sets (DHFSs) is a generalization of fuzzy sets (FSs) and it is typical of membership and non-membership degrees described by some discrete numerical. In this article we chiefly concerned with introducing the aggregation operators for aggregating dual hesitant fuzzy elements (DHFEs), including the dual hesitant fuzzy arithmetic mean and geometric mean. We laid emphasis on discussion of properties of newly introduced operators, and give a numerical example to describe the function of them. Finally, we used the proposed operators to select human resources outsourcing suppliers in a dual hesitant fuzzy environment.


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