Evolutionary bet hedging encapsulates the counterintuitive idea that organisms evolve traits that reduce short-term reproductive success in favor of longer-term risk reduction. It has been widely investigated theoretically, and many putative examples have been cited including practical ones such as the dormancy involved in microbe and weed persistence. However, long-term data on demographic variation from the actual evolutionarily relevant environments have been unavailable to test for its mechanistic relationship to alleged bet hedging traits. I report an association between delayed germination (a bet hedging trait) and risk using a 22-year data set on demographic variation for 10 species of desert annual plants. Species with greater variation in reproductive success (per capita survival from germination to reproduction x per capita fecundity of survivors) were found to have lower average germination fractions. This provides a definitive test using realistic data on demographic variance that confirms the life history prediction for bet hedging. I also showed that the species with greater long-term demographic variation tended to be the ones with greater sensitivity of reproductive success to variation among years in growing-season precipitation.

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Bet hedging in a guild of desert annuals

Uncertainty is a problem not only in human decision-making, but is a prevalent quality of natural environments and thus requires evolutionary response. Unpredictable natural selection is expected to result in the evolution of bet-hedging strategies, which are adaptations to long-term fluctuating selection. Despite a recent surge of interest in bet hedging, its study remains mired in conceptual and practical difficulties, compounded by confusion over what constitutes evidence for its existence. Here, I attempt to resolve misunderstandings about bet hedging and its relationship with other modes of response to environmental change, identify the challenges inherent to its study and assess the state of existing empirical evidence. The variety and distribution of plausible bet-hedging traits found across 16 phyla in over 100 studies suggest their ubiquity. Thus, bet hedging should be considered a specific mode of response to environmental change. However, the distribution of bet-hedging studies across evidence categories—defined according to potential strength—is heavily skewed towards weaker categories, underscoring the need for direct appraisals of the adaptive significance of putative bet-hedging traits in nature.

https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2011.0176

Modes of response to environmental change and the elusive empirical evidence for bet hedging

Introduction. Ecologically Meaningful Germination Studies. Types of Seed Dormancy. Germination Ecology of Seeds with Nondeep Physiological Dormancy. Germination Ecology of Seeds with Morphophysiological Dormancy. Germination Ecology of Seeds with Physical Dormancy. Germination Ecology of Seeds in the Persistent Seed Bank. Causes of Within-Species Variations in Seed Dormancy and Germination Characteristics. A Geographical Perspective on Germination Ecology: Tropical and Subtropical Zones. A Geographical Perspective on Germination Ecology: Temperate and Arctic Zones. Germination Ecology of Plants with Specialized Life Cycles and/or Habitats. Biogeographical and Evolutionary Aspects of Seed Dormancy. Subject Index.

A Definitive Book on Seed Ecology@@@Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination

Genetically identical cells grown in the same culture display striking cell-to-cell heterogeneity in gene expression and other traits. A crucial challenge is to understand how much of this heterogeneity reflects the noise tolerance of a robust system and how much serves a biological function. In bacteria, stochastic gene expression results in cell-to-cell heterogeneity that might serve as a bet-hedging mechanism, allowing a few cells to survive through an antimicrobial treatment while others perish. Despite its clinical importance, the molecular mechanisms underlying bet hedging remain unclear. Here, we investigate the mechanisms of bet hedging in Saccharomyces cerevisiae using a new high-throughput microscopy assay that monitors variable protein expression, morphology, growth rate, and survival outcomes of tens of thousands of yeast microcolonies simultaneously. We find that clonal populations display broad distributions of growth rates and that slow growth predicts resistance to heat killing in a probabalistic manner. We identify several gene products that are likely to play a role in bet hedging and confirm that Tsl1, a trehalose-synthesis regulator, is an important component of this resistance. Tsl1 abundance correlates with growth rate and replicative age and predicts survival. Our results suggest that yeast bet hedging results from multiple epigenetic growth states determined by a combination of stochastic and deterministic factors.

/pdf/bet-hedging-in-yeast-by-heterogeneous-age-correlated-cd4ldfkkzz.pdf

Bet Hedging in Yeast by Heterogeneous, Age-Correlated Expression of a Stress Protectant

For a short-lived organism, such as a freshwater zooplankter, the ways of coping with years of local recruitment failure are either to disperse between habitats and recolonise or to disperse in time through diapause. Diapause is common among freshwater zooplankton and is generally seen as a way to escape periods of harsh environmental conditions. The egg-bank or pool of diapausing copepodites in lake sediments resulting from the production of diapausing stages has several implications for zooplankton ecology, genetics, and evolution which we outline in this review. The presence of a benthic dormant stage also creates a coupling between the benthic habitat and the pelagic, and we argue that zooplankton phenology is a result of selective forces in both habitats. The spatial distribution of diapausing eggs appears to be governed by random resuspension dynamics coupled with higher hatching rates in shallow waters. For diapausing copepodites, however, an active choice of where and how deep to enter the sediment may affect their distribution. In a reanalysis of published data, we found a size-dependent bathymetric distribution and vertical distribution in the sediment of diapausing cyclopoid copepodites. Our review of published laboratory studies showed that predictors of seasonal change such as photoperiod and temperature were the only type of cues used for the termination of diapause. We also found a relation between generation length and the type of cue used for diapause induction: copepods mainly used seasonal cues from the abiota, rotifers mainly used cues from the biotic environment, and cladocerans used a mix of both types. We describe patterns in emergence timing and contribution to population dynamics from studies using in situ estimation of emergence, and conclude that hatching from dormant stages may qualitatively and quantitatively affect zooplankton population dynamics and seasonal succession.

Dormancy in freshwater zooplankton: Induction, termination and the importance of benthic-pelagic coupling

Temporal variability in survivorship and reproduction is predicted to affect the evolution of life‐history characters. Desert annual plants experience temporal variation in reproductive success that is largely caused by precipitation variability. We studied several populations of the desert annual Plantago insularis along a precipitation gradient. Whereas models of bet hedging in unpredictable environments generally predict one optimal germination fraction for a population, empirical studies have shown that environmental conditions during germination can cause a range of germination fractions to be expressed. In a 4‐yr field study, we found that populations in historically more xeric environments had lower mean germination fractions, as is predicted by bet‐hedging models. However, populations exhibited significant variation in germination among years. Two experimental studies measuring germination under several environment conditions were conducted to elucidate the source of this in situ variati...

https://www.jstor.org/stable/10.1086/303314

M. J. Clauss

Papers

Seed Germination in Desert Annuals: An Empirical Test of Adaptive Bet Hedging.