Induction of embryonic dormancy in the Calanoid copepod Acartia hudsonica: proximal cues and variation among individuals

In this study of Acartia hudsonica, the change in egg types from subitaneous eggs to dormant eggs was investigated. Temperature and photoperiod were hypothesized to act as the proximal environmental cues inducing this population to produce dormant eggs, and the variation in the dormant egg responses of individual females was quantified. Some field-collected females produced some dormant eggs early in the spring (March and April), but most of the switch to dormant eggs occurred in late June. Short-term experiments showed that increased temperatures increased the percentage of dormant eggs produced, whereas increased length of day did not. Long-term experiments showed a different pattern from short-term experiments, indicating that duration of exposure to particular temperatures or photoperiods may influence dormant egg production. Under certain conditions, it appeared that nearly all copepods were capable of producing dormant eggs, although identical experimental conditions did not affect all individuals similarly. In several experiments that revealed a significant treatment effect, between 42% and 85% of the variance detected could be attributed to variation among individuals. Also, the fraction of individuals producing dormant eggs generally increased with increasing water temperature. In Narragansett Bay, A. hudsonica appears to produce dormant eggs primarily in response to increased water temperatures although an effect of photoperiod could not be ruled out. Furthermore, individual responses vary greatly. Appreciation of such variation in an important life history trait, such as embryonic dormancy, is important in understanding how this population may be adapted to its environment.     

Spatially heterogeneous stochasticity and the adaptive diversification of dormancy

Diversified bet‐hedging, a strategy that leads several individuals with the same genotype to express distinct phenotypes in a given generation, is now well established as a common evolutionary response to environmental stochasticity. Life‐history traits defined as diversified bet‐hedging (e.g. germination or diapause strategies) display marked differences between populations in spatial proximity. In order to find out whether such differences can be explained by local adaptations to spatially heterogeneous environmental stochasticity, we explored the evolution of bet‐hedging dormancy strategies in a metapopulation using a two‐patch model with patch differences in stochastic juvenile survival. We found that spatial differences in the level of environmental stochasticity, restricted dispersal, increased fragmentation and intermediate survival during dormancy all favour the adaptive diversification of bet‐hedging dormancy strategies. Density dependency also plays a major role in the diversification of dormancy strategies because: (i) it may interact locally with environmental stochasticity and amplify its effects; however, (ii) it can also generate chaotic population dynamics that may impede diversification. Our work proposes new hypotheses to explain the spatial patterns of bet‐hedging strategies that we hope will encourage new empirical studies of this topic.     

Impact of temperature shifts on the joint evolution of seed dormancy and size

Seed dormancy and size are two important life‐history traits that interplay as adaptation to varying environmental settings. As evolution of both traits involves correlated selective pressures, it is of interest to comparatively investigate the evolution of the two traits jointly as well as independently. We explore evolutionary trajectories of seed dormancy and size using adaptive dynamics in scenarios of deterministic or stochastic temperature variations. Ecological dynamics usually result in unbalanced population structures, and temperature shifts or fluctuations of high magnitude give rise to more balanced ecological structures. When only seed dormancy evolves, it is counter‐selected and temperature shifts hasten this evolution. Evolution of seed size results in the fixation of a given strategy and evolved seed size decreases when seed dormancy is lowered. When coevolution is allowed, evolutionary variations are reduced while the speed of evolution becomes faster given temperature shifts. Such coevolution scenarios systematically result in reduced seed dormancy and size and similar unbalanced population structures. We discuss how this may be linked to the system stability. Dormancy is counter‐selected because population dynamics lead to stable equilibrium, while small seeds are selected as the outcome of size‐number trade‐offs. Our results suggest that unlike random temperature variation between generations, temperature shifts with high magnitude can considerably alter population structures and accelerate life‐history evolution. This study increases our understanding of plant evolution and persistence in the context of climate changes.     

The relationship between pyridine nucleotides and seed dormancy

To investigate the proposed role for NAD metabolism in regulating seed dormancy, NAD metabolites and associated enzyme activities were analysed in seed of Arabidopsis thaliana ecotypes ranging from Col-0, which has low seed dormancy, to Cvi, which is highly dormant. Seed poly(ADP-ribosyl)ation levels did not correlate well with the depth of seed dormancy but did correlate with the sensitivity of germination to the DNA damaging agent MMS. Cvi seed had relatively high NAD and low NADP levels compared with the less dormant ecotypes and the NAD : NADP ratios correlated well with dormancy. The activity of NAD kinase was relatively low, and NADP phosphatase was relatively high in dormant Cvi seed, indicating that these enzymes may be involved in controlling the NAD : NADP ratio. Dormant fresh Cvi and nondormant after-ripened Cvi seeds were used to investigate further. Measurement of reduced and oxidised pyridine nucleotides indicated that breaking of dormancy was associated with a reduction in NAD levels but not with an increase in NADP levels. It is proposed that poly(ADP-ribose) polymerase is involved in protecting the seed from genotoxic stress, whereas the level of NAD affects the depth of dormancy, perhaps by enhancing abscisic acid (ABA) synthesis.     

Habitat preferences as related to the prolonged dormancy of perennial herbs and ferns

Prolonged dormancy (hereafter dormancy), a phenomenon in which a perennial herbaceous plant does not sprout for one or more years, is examined. The phenomenon may be more frequent than stated so far and discovery of its role in plant life history and performance is still underway. Data from published papers was reviewed and all known species exhibiting dormancy were analysed from the aspect of species ecological values. Adaptation to environmental factors (light, moisture, pH and nitrogen as estimated by Ellenberg indicator values) influences the maximum duration of dormancy. A higher proportion of plants are dormant in species that prefer to grow in good light conditions, dry sites and infertile soil. The duration of dormant period is longer in species that prefer to grow in dry sites and/or calcareous soils. A range of factors, we believe, control plant behaviour, however, it is suggested that environmental stress is the principal factor inducing dormancy.     

Prolonged diapause: a trait increasing invasion speed?

Invasive species are considered to be the second cause of biodiversity erosion, and one challenge is to determine the life history traits that cause an increased invasion capacity. Prolonged diapause is a major trait in evolution and insect population dynamics, but its effects on invasion speed remain unknown. From a recently developed mathematical approach (integro-difference equations) applied to the insect dormancy, we show that despite a dispersal cost, bet-hedging diapause strategies with low (0.1-0.2) prolonged diapause frequency (emergence after 1 or 2 years) can have a higher invasion speed than a simple diapause strategy (emergence after 1 year) when the environmental stochasticity is sufficiently high. In such conditions, prolonged diapause is a trait supporting invasion capacity by increasing population stochastic growth rate. This conclusion, which applies to a large range of demographic parameters, is in opposition to the usual view that prolonged dormancy is an alternative strategy to dispersal. However, prolonged diapause does not support invasion if the level of environmental stochasticity is low. Therefore, conclusion about its influence on invasion ability needs a good knowledge of environmental stochasticity in the introduction area of considered species.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

A QTL located on chromosome 4A associated with dormancy in white- and red-grained wheats of diverse origin

Improved resistance to preharvest sprouting in modern bread wheat (Triticum aestivum. L.) can be achieved via the introgression of grain dormancy and would reduce both the incidence and severity of damage due to unfavourable weather at harvest. The dormancy phenotype is strongly influenced by environmental factors making selection difficult and time consuming and this trait an obvious candidate for marker assisted selection. A highly significant Quantitative Trait Locus (QTL) associated with grain dormancy and located on chromosome 4A was identified in three bread wheat genotypes, two white- and one red-grained, of diverse origin. Flanking SSR markers on either side of the putative dormancy gene were identified and validated in an additional population involving one of the dormant genotypes. Genotypes containing the 4A QTL varied in dormancy phenotype from dormant to intermediate dormant. Based on a comparison between dormant red- and white-grained genotypes, together with a white-grained mutant derived from the red-grained genotype, it is concluded that the 4A QTL is a critical component of dormancy; associated with at least an intermediate dormancy on its own and a dormant phenotype when combined with the R gene in the red-grained genotype and as yet unidentified gene(s) in the white-grained genotypes. These additional genes appeared to be different in AUS1408 and SW95-50213.     

Microbial seed banks: the ecological and evolutionary implications of dormancy

Dormancy is a bet-hedging strategy used by a wide range of taxa, including microorganisms. It refers to an organism's ability to enter a reversible state of low metabolic activity when faced with unfavourable environmental conditions. Dormant microorganisms generate a seed bank, which comprises individuals that are capable of being resuscitated following environmental change. In this Review, we highlight mechanisms that have evolved in microorganisms to allow them to successfully enter and exit a dormant state, and discuss the implications of microbial seed banks for evolutionary dynamics, population persistence, maintenance of biodiversity, and the stability of ecosystem processes.     

Incidence of diapause varies among populations of <Emphasis Type="Italic">Daphnia pulicaria</Emphasis>

Dormancy is a common way in which organisms survive environmental conditions that would be lethal to the active individual. However, while dormant, individuals forgo reproduction. Hence theory suggests an optimal time in which to enter dormancy, depending on risks associated with both remaining active and entering dormancy. When these relative risks differ among habitats, dormancy strategies are predicted to vary as well. For freshwater zooplankton, it has been suggested that sensitivity to the cues that initiate dormancy should be selected against when females have the opportunity to remain in the water column year round. We tested this prediction with 12 populations of lake-dwelling Daphnia pulicaria (Crustacea: Cladocera). Differences among lakes in basin morphometry, predators and resources create a gradient of risk for Daphnia in the water column. Some populations persist in high numbers year round while others are abundant only in spring. We used this difference in persistence ability as an estimate of risk in the water-column. For 3 years of field sampling we found consistent differences among the lake populations in the incidence of dormancy. In some populations, only a small fraction of females switched to producing dormant eggs each year whereas in others the majority of eggs produced in the late spring were dormant. In general, populations that experienced predictably low abundances in the active form exhibited higher incidence of dormancy than did populations that persisted in high abundance year round, but there were exceptions. Our results confirm that the incidence of dormancy varies considerably among populations in a fashion consistent with general theory, but suggest that persistence in the water column is not the sole predictor of the diapause strategy found in any particular lake.     

Environmental factors for establishment of the dormant state in oocytes

Guaranteeing the sustainability of gametogenesis is a fundamental issue for perpetuating the species. In the mammalian ovary, sustainability is accomplished by keeping a number of oocytes “stocked” in the dormant state. Despite the evident importance of this state, the mechanisms underlying the oocyte dormancy are not fully understood, although it is presumed that both intrinsic and extrinsic factors are involved. Here, we review environmental factors involved in the regulation of oocyte dormancy. Consideration of the environmental factors illustrates the nature of the ovarian compartment, in which primordial follicles reside. This should greatly improve our understanding of the mechanisms and also assist in reconstitution of the dormant state in culture. Accumulating knowledge on the dormant state of oocytes will contribute to a wide range of research in fields such as developmental biology, reproductive biology and regenerative medicine.     

Population dynamics and the colour of environmental noise.

The effect of red, white and blue environmental noise on discrete-time population dynamics is analyzed. The coloured noise is superimposed on Moran-Ricker and Maynard Smith dynamics, the resulting power spectra are less than examined. Time series dominated by short- and long-term fluctuations are said to be blue and red, respectively. In the stable range of the Moran-Ricker dynamics, environmental noise of any colour will make population dynamics red or blue depending the intrinsic growth rate. Thus, telling apart the colour of the noise from the colour of the population dynamics may not be possible. Population dynamics subjected to red and blue environmental noises show, respectively, more red or blue power spectra than those subjected to white noise. The sensitivity to differences in the noise colours decreases with increasing complexity and ultimately disappears in the chaotic range of the population dynamics. These findings are duplicated with the Maynard Smith model for high growth rates when the strength of density dependence changes. However, for low growth rates the power spectra of the population dynamics with noise are red in stable, periodic and aperiodic ranges irrespective of the noise colour. Since chaotic population fluctuations may show blue spectra in the deterministic case, this implies that blue deterministic chaos may become red under any colour of the noise.     

Contrasting bud bank dynamics of two co-occurring grasses in tallgrass prairie: implications for grassland dynamics

Because most shoot recruitment in perennial grasses occurs from belowground axillary buds, bud dynamics determine plant population dynamics and meristem limitation to population growth. Therefore, grassland vegetation responses to environmental change or disturbance may be influenced by interspecific differences in bud banks and the patterns and environmental controls of bud development and demography. We examined bud bank dynamics in Andropogon gerardii and Dichanthelium oligosanthes in tallgrass prairie by enumerating and classifying buds throughout 15?months to determine whether grass buds live for multiple years and accumulate; whether bud natality, dormancy and outgrowth are synchronous or variable; and whether bud bank dynamics differ between these co-occurring species. Andropogon gerardii (a C₄ species) maintained a larger dormant bud bank, showed synchrony in bud development and transition to tiller, and its buds lived for multiple years. Thus, multiple previous years?? bud cohorts contributed to recruitment. By contrast, D. oligosanthes (a C₃ species) maintained a smaller dormant bud bank, had asynchronous bud development with active buds present year-round, and its buds lived for 1?year. Buds played different roles in the dynamics of each species, allowing A. gerardii to over-winter and recruit new spring tillers and D. oligosanthes to survive and recruit new tillers following summer dormancy. These differences in bud bank age structure, phenology, and dynamics between these species suggest greater demographic buffering and time-lag effects in A. gerardii populations. Interspecific differences in bud bank structure and dynamics may explain and help predict grassland responses to environmental change.     

Mapping of QTL for seed dormancy in a winter oilseed rape doubled haploid population

Following winter oilseed rape cultivation, considerable numbers of volunteer oilseed rape plants may occur in subsequent years in following crops. The appearance of volunteer oilseed rape plants is based on the capability of the seeds to become secondary dormant and to survive in this stage for many years in the soil. Genetic reduction of secondary seed dormancy in oilseed rape could provide a means to reduce the frequency of volunteer plants and especially the dispersal of transgenic oilseed rape. The objective of the present study was to analyse the inheritance of primary and secondary seed dormancy in a winter oilseed rape doubled haploid population derived from the cross Express 617 × R53 and to study correlations to other seed traits. Field experiments were performed in Germany for 2 years at two locations with two replicates. Seeds harvested from open pollinated plants were used for all analyses, including a laboratory test for seed dormancy. A previously developed molecular marker map of the doubled haploid population was used to map QTL of the relevant traits. For primary, secondary and total seed dormancy, the results showed significant effects of the genotypes and their interactions, with years and locations. Two, four and five QTL were detected for primary, secondary and total seed dormancy which explained 19, 35 and 42 % of the phenotypic variance, respectively. Results show that secondary seed dormancy is a heritable trait and that selection for low secondary seed dormancy is possible.     

种子物理休眠研究进展

种子物理休眠是由种皮不透水层引起的一种休眠类型,是植物在长期系统发育进程中获得的一种适应环境变化的特性。该文简述了种子物理休眠的定义与概念;从不透水层、种皮的特殊水孔器结构以及胚的形态特异性等方面,综述了物理休眠种子的形态特征、物理休眠与综合休眠的解除方法以及物理休眠的可能解除机制;利用Angiosperm Phylogeny Group Ⅲ(APG Ⅲ)系统分析了种子物理休眠的植物在系统发育中的位置;最后提出了今后种子物理休眠有待研究的主要问题。     

Dormancy in the Copepoda — an overview

Dormancy affects copepods in their anatomy, physiology, genetics, population biology, community ecology, evolution and local and geographic distribution. It is known from freeliving representatives of three copepod taxa, namely the Harpacticoida, Cyclopoida and Calanoida. Species showing dormancy occur in various realms and habitats, both freshwater and marine, being benthic, planktic or ice-dwelling. Depending on the taxon, dormancy occurs at various times of the year, prevailing in higher and temperate latitudes. Copepod dormancy is expressed in various ontogenetic stages, such as resting eggs, arrested larval development, juvenile and adult encystment, or arrested development of nonencysted copepodids or adults. Ecologically, dormancy is an energy saving trait, allowing the individual to bridge periods of environmental harshness. Adverse environmental conditions could be abiotic (e.g. desiccation, temperature, oxygen availability) or biotic in nature (e.g. food availability, predation). Diapause s. str. is initiated, maintained and terminated by triggering factors (e.g. photoperiod, temperature, chemical cues, population density/physiological factors). The dormant state and emergence patterns directly affect reproduction, population dynamics, community composition, coexistence and distribution of copepods, as well as the phenology of their predators and living food items. Populations having dormancy, in most cases belong to and affect communities of two realms: the water column and the bottom. Dormant stages may provide means for dispersal as well as for staying in special localities. The variability of dormancy permits flexible and complex life histories. Dormancy is subjected to and on the other hand affects copepod evolution.     

THE JOINT EVOLUTION OF DISPERSAL AND DORMANCY IN A METAPOPULATION WITH LOCAL EXTINCTIONS AND KIN COMPETITION

Dispersal and dormancy are two strategies that allow recolonization of empty patches and escape from kin competition. Because they presumably respond to similar evolutionary forces, it is tempting to consider that these strategies may substitute for each other. Yet in order to predict the outcome of the evolution of dispersal and dormancy, and to characterize the emerging covariation between both traits, it is necessary to consider models where dispersal and dormancy evolve jointly. Here, we analyze the evolution of dispersal and dormancy as a function of direct fitness costs, environmental variation, and competition among relatives. We consider two scenarios depending on whether the rates of dormancy for philopatric and dispersed individuals are constrained to be the same (unconditional dormancy) or allowed to be different (conditional dormancy). We show that only philopatric individuals should enter dormancy, at a rate increasing with increasing rates of local extinction and decreasing population sizes. When dormancy and dispersal evolve jointly, we observe a wide range of evolutionary outcomes. In particular, we find that the pattern of covariation between the evolutionarily stable rates of dispersal and dormancy is molded by the rate of extinction and the local population size.     

Geographical variation in dormancy in a copepod: evidence from population crosses

Populations of the freshwater copepod Mesocyclops edax inhabiting Michigan lakes are dormant during winter, whereas populations inhabiting Florida lakes develop and reproduce continuously throughout the year. A Michigan and a Florida population were exposed to dormancy inducing conditions (low temperature and short photoperiod) in the laboratory and observed for indications of dormancy. All Michigan individuals and a small percentage of the Florida individuals entered dormancy as indicated by prolonged duration of the fourth copepodid instar and cessation of feeding. I suggest that in these population these observations represent diapause, rather than quiescence. The two populations were crossbred to examine the nature of inheritance of dormancy. The F₁ hybrids exhibited an incidence of diapause approximately intermediate between the Florida and Michigan parental stocks. The backcrosses of F₁ individuals to the Michigan and Florida stocks, respectively, exhibited a high and an intermediate incidence of diapause. Survival of the F₂ crosses was very low. The present study presents evidence of genetic differentiation between the Michigan and Florida populations of M. edax with respect to ability to diapause.     

Delayed germination and dispersal in desert annuals: Escape in space and time

Summary A model is developed to consider the interplay between dispersibility and delayed germination in desert annuals. The model explores the effect of low levels of dispersal, considered realistic for annual plants, on optimal germination fraction. The model also demonstrates the effect of the amount and accuracy of predictive (responsive to the environment) dormancy on the optimal innate germination fraction (not responsive to environmental conditions).Optimal germination fraction is found to be very sensitive to changes in despersibility especially at the limited dispersibilities that are realistic for annual plants. As dispersibility increases, optimal germination fraction increases. If plants make two kinds of seeds with differing despersibility, reproduction is maximized if the low dispersal seeds have delayed germination and the high dispersal seeds have quick germination. If dormancy mechanisms permit seeds to germinate when environmental conditions allow successful maturation, and remain dormant when environmental conditions do not permit successful maturation, what fraction of seeds should remain dormant under predicted good conditions as a hedge against inaccurate prediction of the environment? If environmental cues that break dormancy are uncorrelated with environmental conditions that permit successful maturation, predictive dormancy has little or no effect on the optimal innate germination fraction. When predictive dormancy lowers the probability of germinating when environmental conditions preclude successful maturation, the optimal innate germination fraction increases with increasing germination control by predictive dormancy. With a moderate degree of germination control by predictive dormancy, the optimal innate dormancy is still sensitive to changes in dispersal in the low dispersal ranges characteristic of annual plants.Evidence is presented from plant species that have both dispersal and germination dimorphisms to support the predicted correlation of high germination fractions with high dispersal.     

Ethylene in seed dormancy and germination

The role of ethylene in the release of primary and secondary dormancy and the germination of non-dormant seeds under normal and stressed conditions is considered. In many species, exogenous ethylene, or ethephon – an ethylene-releasing compound - stimulates seed germination that may be inhibited because of embryo or coat dormancy, adverse environmental conditions or inhibitors (e.g. abscisic acid, jasmonate). Ethylene can either act alone, or synergistically or additively with other factors. The immediate precursor of ethylene biosynthesis, 1-aminocyclopropane-1-carboxylic acid (ACC), may also improve seed germination, but usually less effectively. Dormant or non-dormant inhibited seeds have a lower ethylene production ability, and ACC and ACC oxidase activity than non-dormant, uninhibited seeds. Aminoethoxyvinyl-glycine (AVG) partially or markedly inhibits ethylene biosynthesis in dormant or non-dormant seeds, but does not affect seed germination. Ethylene binding is required in seeds of many species for dormancy release or germination under optimal or adverse conditions. There are examples where induction of seed germination by some stimulators requires ethylene action. However, the mechanism of ethylene action is almost unknown.
The evidence presented here shows that ethylene performs a relatively vital role in dormancy release and seed germination of most plant species studied.