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1.
Many workers have demonstrated a genetic basis for variation in inflorescence traits, but this variation can also have an environmental component. Because flowering can incur significant water costs, I estimated plasticity of inflorescence traits of three populations of Lobelia siphilitica in response to drought. I manipulated soil water availability in the greenhouse and measured seven inflorescence traits. Under drought conditions, plants from one population flowered later and produced fewer flowers with shorter corollas and narrower landing pads. In contrast, the height of the flowering stalk decreased in response to drought in all three populations. Consequently, pollinator-mediated natural selection on these plastic traits may depend on soil water availability. Plastic responses differed between genotypes only for the height of the flowering stalk and the length of the corolla tube and only in one or two populations. This suggests that genotype × environment interactions would not limit the evolution of inflorescence traits in L. siphilitica. The strength and sign of phenotypic correlations among inflorescence traits did not respond plastically to drought, suggesting that indirect selection on inflorescence traits of L. siphilitica will not vary strongly with water availability. My results suggest that plasticity of inflorescence traits may influence their evolution, but the effects are population- and trait-specific.  相似文献   

2.
Genetic variation in life history traits has important consequences for life-history evolution. Here we report the results of a greenhouse experiment investigating the broad sense genetic basis of variation in life history traits within and among five populations of Campanula americana distributed along a latitudinal gradient. The populations exhibit differentiation for a number of morphological traits (seed weight, number of branches, final plant size, number of capsules) and the phenological traits, days to emergence, days to bolting, the onset of flowering, and the duration of flowering. Families within populations differed only in days to emergence and seed weight. These results suggest that the life history differences among populations are genetically based. In addition, two life history types—winter annuals and biennials—have previously been reported from natural populations of Campanula americana. This experiment identified a third type—summer annuals from the Florida population.  相似文献   

3.
Aims Foliar herbivory and water stress may affect floral traits attractive to pollinators. Plant genotypes may differ in their responses to the interplay between these factors, and evolution of phenotypic plasticity could be expected, particularly in heterogeneous environments. We aimed at evaluating the effects of simulated herbivory and experimental drought on floral traits attractive to pollinators in genetic families of the annual tarweed Madia sativa, which inhabits heterogeneous environments in terms of water availability, herbivore abundance and pollinator abundance.Methods In a greenhouse experiment with 15 inbred lines from a M. sativa population located in central Chile (Mediterranean-type climate), we measured the effects of apical bud damage and reduced water availability on: number of ray florets per flower head, length of ray florets, flower head diameter, number of open flower heads per plant, flowering plant height and flowering time.Important findings Apical damage and water shortage reduced phenotypic expression of floral traits attractive to pollinators via additive and non-additive effects. Plants in low water showed decreased height and had fewer and shorter ray florets, and fewer and smaller flower heads. Damaged plants showed delayed flowering, were less tall, and showed shorter ray florets and smaller flower heads. The number of ray florets was reduced by damage only in the low water treatment. Plant height, flowering time and number of flower heads showed among-family variation. These traits also showed genetic variation for plasticity to water availability. Ray floret length, flower head size and time to flowering showed genetic variation for plastic responses to apical damage. Plasticity in flowering time may allow M. sativa to adjust to the increased aridity foreseen for its habitat. Because genetic variation for plastic responses was detected, conditions are given for evolutionary responses to selective forces acting on plastic traits. We suggest that the evolution of adaptive floral plasticity in M. sativa in this ecological scenario (heterogeneous environments) would result from selective forces that include not only pollinators but also resource availability and herbivore damage.  相似文献   

4.
How much of the variation seen in life histories is consistent with adaptive hypotheses, and how much requires other kinds of explanation? Differences in flowering time between Sonoran (earlier flowering) and Chihuahuan Desert (later flowering) populations of the desert annual Eriogonum abertianum Torr. (Polygonaceae) are significant, repeatable between greenhouse experiments, and persist into a second greenhouse generation. These apparent genetic differences are consistent with a hypothesis of local adaptation: field demographic studies (Fox, 1989b) show that many fewer Sonoran than Chihuahuan Desert plants survive to the summer rainy season, suggesting selection for earlier flowering in the Sonoran Desert. Within natural populations there is considerable phenological complexity: time of first flowering varies by up to six months, and individuals may have zero, one, or several reproductive episodes. Greenhouse sib analyses revealed only marginal among-family genetic variation for flowering size. The resemblance between parents and offspring for size and time of flowering varied with growth conditions, suggesting that this marginal variation among families may be at least partly due to factors other than additive genetic variance. On the other hand, moisture limitation significantly delayed the onset of flowering in two independent experiments. Variation in moisture availability in both time and space is characteristic of desert environments. The phenological complexity in natural populations may thus be generated by random variation in moisture availability, possibly in conjunction with variation in germination date and plant size. The results call into question the claim that drought generally induces flowering in desert annuals.  相似文献   

5.
Sex-specific interactions with antagonists may explain female maintenance in gynodioecious populations if seeds produced by hermaphroditic plants are preferred over seeds produced by female plants. Among antagonistic interactions, pre-dispersal seed predators have received relatively little attention even though they may exert sex-specific selective pressures on the evolution of floral and flowering traits. In this work, I investigate temporal variation in seed predation in gynodioecious Geranium sylvaticum, where in addition to female and hermaphrodite individuals, plants with an intermediate sexual expression are also present in most populations. Specifically, I examined whether seed predation is linked to flowering phenology, plant gender, and sexual dimorphism in floral and seed traits over the flowering season using an experimental field population. Within the population, I selected female, intermediate, and hermaphrodite plants with different timing of flowering onset (early, mid, or late), and collected seeds across the fruiting period. Seeds were weighed and examined for seed predator damage. The results show that the three genders experienced similar levels of seed predation attack regardless of their flowering phenology, and that overall seed predation was not related to changes in seed production or seed mass. These results suggest that sexual dimorphism in seed predation cannot be responsible for female maintenance in this species.  相似文献   

6.
We examined patterns of genetic variance and covariance in two traits (i) carbon stable isotope ratio delta13C (dehydration avoidance) and (ii) time to flowering (drought escape), both of which are putative adaptations to local water availability. Greenhouse screening of 39 genotypes of Arabidopsis thaliana native to habitats spanning a wide range of climatic conditions, revealed a highly significant positive genetic correlation between delta13C and flowering time. Studies in a range of C3 annuals have also reported large positive correlations, suggesting the presence of a genetically based trade-off between mechanisms of dehydration avoidance (delta13C) and drought escape (early flowering). We examined the contribution of pleiotropy by using a combination of mutant and near-isogenic lines to test for positive mutational covariance between delta13C and flowering time. Ecophysiological mutants generally showed variation in delta13C but not flowering time. However, flowering time mutants generally demonstrated pleiotropic effects consistent with natural variation. Mutations that caused later flowering also typically resulted in less negative delta13C and thus probably higher water use efficiency. We found strong evidence for pleiotropy using near-isogenic lines of Frigida and Flowering locus C, cloned loci known to be responsible for natural variation in flowering time. These data suggest the correlated evolution of delta13C and flowering time is explained in part by the fixation of pleiotropic alleles that alter both delta13C and time to flowering.  相似文献   

7.
Increases in nitrogen (N) deposition and variation in precipitation have been occurring in temperate deserts; however, little information is available regarding plant phenological responses to environmental cues and their relationships with plant growth pattern in desert ecosystems. In this study, plant phenology and growth of six annuals in response to N and water addition were monitored throughout two consecutive growing seasons in 2011 and 2012 in a temperate desert in northwestern China. The effects of N and water addition on reproductive phenology differed among plant species. N and water addition consistently advanced the flowering onset time and fruiting time of four spring ephemerals; however, their effects on two spring‐summer annuals were inconsistent, with advances being noted in one species and delays in another. N and water addition alone increased plant height, relative growth rate, leaf number, flower number, and individual biomass, while their combinative effects on plant growth and reproductive phenology were dependent on species. Multiple regression analysis showed that flowering onset time was negatively correlated with relative growth rate of two species, and negatively correlated with maximum plant height of the other four species. Our study demonstrates that phenological responses to increasing precipitation and N deposition varied in annuals with different life histories, whereby the effects of climate change on plant growth rate were related to reproductive phenology. Desert annuals that were able to accelerate growth rate under increasing soil resource availability tended to advance their flowering onset time to escape drought later in the growing season. This study promotes our understanding of the responses of temperate desert annuals to increasing precipitation and N deposition in this desert.  相似文献   

8.
In heterogeneous environments, selection on life-history traits and flowering time may vary considerably among populations because of differences in the extent to which mortality is related to age or size, and because of differences in the seasonal patterns of resource availability and intensity of biotic interactions. Spatial variation in optimal reproductive effort and flowering time may result in the evolution of genetic differences in life-history traits, but also in the evolution of adaptive phenotypic plasticity. The perennial herb Primula farinosa occurs at sites that differ widely in soil depth and therefore in water-holding capacity, vegetation cover, and frost-induced soil movement in winter. We used data from eight natural populations and a common-garden experiment to test the predictions that reproductive allocation is negatively correlated with soil depth while age at first reproduction and first flowering date among reproductive individuals are positively correlated with soil depth. In the common-garden experiment, maternal families collected in the field were grown from seed and monitored for 5 years. In the field, reproductive effort (number of flowers in relation to rosette area) varied among populations and was negatively related to soil depth. In the common-garden experiment, among-population differences in age at first reproduction, and reproductive effort were statistically significant, but relatively small and not correlated with soil depth at the site of origin. Flowering time varied considerably among populations, but was not related to soil depth at the site of origin. Taken together, the results suggest that among-population variation in reproductive effort observed in the field largely reflects phenotypic plasticity. They further suggest that among-population differentiation in flowering time cannot be attributed to variation in environmental factors correlated with soil depth.  相似文献   

9.
Local adaptation of plants along environmental gradients provides strong evidence for clinal evolution mediated by natural selection. Plants have developed diverse strategies to mitigate stress, for example, drought escape is a phenological strategy to avoid drought stress, while polyploidy was proposed as a genomic adaptation to stress. Polyploidy as an adaptation to aridity (an environmental parameter integrating temperature and precipitation) was previously documented in annual Brachypodium spp. (Poaceae) in the Western Mediterranean. Here, we examined whether polyploidy or phenology are associated with aridity in annual Brachypodium spp. along the aridity gradient in the Eastern Mediterranean. Using flow cytometry, we determined ploidy levels of plants from natural populations along the Israeli gradient, spanning ∼424 km from mesic Mediterranean to extreme desert climates. In a common garden we recorded time of seedling emergence, flowering and senescence. We tested whether the proportion of allotetraploids in the populations and phenological traits were associated with aridity. Contrary to a previous study in the Western Mediterranean, we found no effect of aridity on the proportion of allotetraploids and diploids within populations. Interestingly, phenology was associated with aridity: time of emergence was later, while flowering and senescence were earlier in desert plants. Our results indicate that in the Eastern Mediterranean, adaptation of Brachypodium to aridity is mediated mainly by phenology, rather than ploidy level. Therefore, we suggest that genome duplication is not the main driver of adaptation to environmental stress; rather, phenological change as a drought escape mechanism may be the major adaptation.  相似文献   

10.
Reproductive timing is a key life‐history trait that impacts the pool of available mates, the environment experienced during flowering, and the expression of other traits through genetic covariation. Selection on phenology, and its consequences on other life‐history traits, has considerable implications in the context of ongoing climate change and shifting growing seasons. To test this, we grew field‐collected seed from the wildflower Mimulus guttatus in a greenhouse to assess the standing genetic variation for flowering time and covariation with other traits. We then created full‐sib families through phenological assortative mating and grew offspring in three photoperiod treatments representing seasonal variation in daylength. We find substantial quantitative genetic variation for the onset of flowering time, which covaried with vegetative traits. The assortatively‐mated offspring varied in their critical photoperiod by over two hours, so that families differed in their probability of flowering across treatments Allocation to flowering and vegetative growth changed across the daylength treatments, with consistent direction and magnitude of covariation among flowering time and other traits. Our results suggest that future studies of flowering time evolution should consider the joint evolution of correlated traits and shifting seasonal selection to understand how environmental variation influences life histories.  相似文献   

11.
Flowering time and water-use efficiency (WUE) are two ecological traits that are important for plant drought response. To understand the evolutionary significance of natural genetic variation in flowering time, WUE, and WUE plasticity to drought in Arabidopsis thaliana, we addressed the following questions: (1) How are ecophysiological traits genetically correlated within and between different soil moisture environments? (2) Does terminal drought select for early flowering and drought escape? (3) Is WUE plasticity to drought adaptive and/or costly? We measured a suite of ecophysiological and reproductive traits on 234 spring flowering accessions of A. thaliana grown in well-watered and season-ending soil drying treatments, and quantified patterns of genetic variation, correlation, and selection within each treatment. WUE and flowering time were consistently positively genetically correlated. WUE was correlated with WUE plasticity, but the direction changed between treatments. Selection generally favored early flowering and low WUE, with drought favoring earlier flowering significantly more than well-watered conditions. Selection for lower WUE was marginally stronger under drought. There were no net fitness costs of WUE plasticity. WUE plasticity (per se) was globally neutral, but locally favored under drought. Strong genetic correlation between WUE and flowering time may facilitate the evolution of drought escape, or constrain independent evolution of these traits. Terminal drought favored drought escape in these spring flowering accessions of A. thaliana. WUE plasticity may be favored over completely fixed development in environments with periodic drought.  相似文献   

12.
We document phenotypic and genetic variation within and among populations of the seed heteromorphic species Heterosperma pinnatum Cav. (Compositae) in the production of seed morphs and in a variety of life-history and morphological characteristics that might be correlated with seed and head traits. Each trait is found to have significant genetic variance in most or, usually, all populations. Significant among-population genetic variation exists for all traits except number of achenes per head and seedling shape, although some traits have much less genetic variation among than within populations. Number and percentage of intermediate achenes per head, total number of achenes per head, and lengths of central and peripheral achenes had little among-population genetic variation compared to within-population variation. Most traits had slightly less genetic variation among than within populations; however, some traits (percentage of central achenes, length of awns, date that the first flowering head opened, date that the first fruiting head opened, and death date) had more among-population genetic variation. The proportions of achene morphs produced had high broad-sense heritabilities and high among-population genetic variance, except in the case of intermediate achenes. All phenological variables had high among-population genetic variation. Within-population heritabilities were high for dates of first flowering head and fruiting head but low for death date and reproductive interval. Family and population means measured in the greenhouse for traits having high broad-sense heritability or among-population genetic variance were closely correlated with field means for the corresponding families or populations. The amounts of phenotypic variation were similar for traits that were measured in both the field and the greenhouse. These lines of evidence suggest that greenhouse results provide reasonable estimates of genetic variation in the field for this species. Numerous studies have reported variation in the proportion of seed morphs for different heteromorphic-seeded species and have discussed adaptive scenarios for the evolution of seed proportions; however, our investigation is one of only a few that have documented the amount of phenotypic and genetic variation within and among populations.  相似文献   

13.
Premise of the studyAs global climate change alters drought regimes, rapid evolution of traits that facilitate adaptation to drought can rescue populations in decline. The evolution of phenological advancement can allow plant populations to escape drought, but evolutionary responses in phenology can vary across a species'' range due to differences in drought intensity and standing genetic variation.Methods Mimulus cardinalis, a perennial herb spanning a broad climatic gradient, recently experienced a period of record drought. Here, we used a resurrection study comparing flowering time and stem height at first flower of pre‐drought ancestors and post‐drought descendants from northern‐edge, central, and southern‐edge populations in a common environment to examine the evolution of drought escape across the latitudinal range.Key resultsContrary to the hypothesis of the evolution of advanced phenology in response to recent drought, flowering time did not advance between ancestors and descendants in any population, though storage condition and maternal effects could have impacted these results. Stem height was positively correlated with flowering time, such that plants that flowered earlier were shorter at first flower. This correlation could constrain the evolution of earlier flowering time if selection favors flowering early at a large size.ConclusionsThese findings suggest that rapid evolution of phenology will not rescue these populations from recent climate change. Future work is needed to examine the potential for the evolution of alternative drought strategies and phenotypic plasticity to buffer M. cardinalis populations from changing climate.  相似文献   

14.
Ivey CT  Carr DE 《Annals of botany》2012,109(3):583-598

Background and Aims

Self-fertilizing taxa are often found at the range margins of their progenitors, where sub-optimal habitats may select for alternative physiological strategies. The extent to which self-fertilization is favoured directly vs. arising indirectly through correlations with other adaptive life history traits is unclear. Trait responses to selection depend on genetic variation and covariation, as well as phenotypic and genetic responses to altered environmental conditions. We tested predictions of the hypothesis that self-fertilization in Mimulus arises through direct selection on physiological and developmental traits that allow seasonal drought escape.

Methods

Phenotypic selection on mating system and drought escape traits was estimated in field populations of M. guttatus. In addition, trait phenotype and phenotypic selection were compared between experimental wet and dry soil in two greenhouse populations each of M. guttatus and M. nasutus. Finally, genetic variation and covariation for traits were compared between wet and dry soil treatments in a greenhouse population of M. guttatus.

Key Results

Consistent with predictions, selection for early flowering was generally stronger than for mating system traits, and selection for early flowering was stronger in dry soil. Inconsistent with predictions, selection for water-use efficiency was largely absent; selection for large flowers was stronger than for drought escape in the field; and most drought escape and mating system traits were not genetically correlated. A positive genetic correlation between flowering time and flower size, which opposed the adaptive contour, emerged only in wet soil, suggesting that variation in water availability may maintain variation in these traits. Plastic responses to soil moisture treatments supported the idea that taxonomic divergence could have been facilitated by plasticity in flowering time and selfing.

Conclusions

The hypothesis that plant mating systems may evolve indirectly via selection on correlated life history characteristics is plausible and warrants increased attention.  相似文献   

15.
Climate change can cause changes in expression of organismal traits that influence fitness. In flowering plants, floral traits can respond to drought, and that phenotypic plasticity has the potential to affect pollination and plant reproductive success. Global climate change is leading to earlier snow melt in snow-dominated ecosystems as well as affecting precipitation during the growing season, but the effects of snow melt timing on floral morphology and rewards remain unknown. We conducted crossed manipulations of spring snow melt timing (early vs. control) and summer monsoon precipitation (addition, control, and reduction) that mimicked recent natural variation, and examined plastic responses in floral traits of Ipomopsis aggregata over 3 years in the Rocky Mountains. We tested whether increased summer precipitation compensated for earlier snow melt, and if plasticity was associated with changes in soil moisture and/or leaf gas exchange. Lower summer precipitation decreased corolla length, style length, corolla width, sepal width, and nectar production, and increased nectar concentration. Earlier snow melt (taking into account natural and experimental variation) had the same effects on those traits and decreased inflorescence height. The effect of reduced summer precipitation was stronger in earlier snow melt years for corolla length and sepal width. Trait reductions were explained by drier soil during the flowering period, but this effect was only partially explained by how drier soils affected plant water stress, as measured by leaf gas exchange. We predicted the effects of plastic trait changes on pollinator visitation rates, pollination success, and seed production using prior studies on I. aggregata. The largest predicted effect of drier soil on relative fitness components via plasticity was a decrease in male fitness caused by reduced pollinator rewards (nectar production). Early snow melt and reduced precipitation are strong drivers of phenotypic plasticity, and both should be considered when predicting effects of climate change on plant traits in snow-dominated ecosystems.  相似文献   

16.
Drought strongly influences plant productivity, suggesting that water limitation has shaped the evolution of many plant physiological traits. One functional strategy that plants employ to cope with decreasing water availability is drought escape. For drought-escaping species, high metabolic activity (gas exchange) and rapid growth are hypothesized to confer a fitness advantage, because this enables a plant to complete its life cycle before the most intense period of drought. By growing an annual grass species (Avena barbata) under well-watered or water-limited conditions in a greenhouse, we directly tested whether high photosynthesis, increased stomatal opening, and early flowering are adaptive under drought. We measured phenotypic selection on instantaneous gas exchange and flowering time as well as the underlying biochemical traits that regulate photosynthesis. We found strong selection for earlier flowering in the dry environment, but no evidence that increased photosynthesis was adaptive under drought. Photosynthetic rate (A) and stomatal conductance (gs) were both adaptively neutral in the dry environment. Increased photosynthetic capacity (Amax) was maladaptive in the dry environment, perhaps because of the respiratory cost associated with maintaining excess enzyme and substrate capacity. There was no correlational selection on the combination of physiology and flowering time in the dry environment, suggesting that accelerated development and high gas exchange may not need to be tightly linked to promote drought escape. In contrast, there was selection for both high photosynthetic function (Amax and A) and early flowering in the well-watered environment. These combinations of traits may have been favored because they maximize both energy and time available for reproduction. Our results suggest that the benefit of increased photosynthesis for plant fitness may be strongest in the absence of drought stress.  相似文献   

17.
The shrub Encelia farinosa (Asteraceae) exhibits geographic variation in aboveground architecture and leaf traits in parallel with environmental variation in temperature and moisture. Measurements of plants occurring across a natural gradient demonstrated that plants in desert populations produce smaller, more pubescent leaves and are more compact and branched than plants in more mesic coastal environments. This phenotypic variation is interpreted in part as adaptive genetic differentiation; small size and pubescence reduce leaf temperature and thus increase water-use efficiency but at the cost of lower photosynthetic rate, which results in slower growth and more compact growth form. We explored the basis of phenotypic variation by planting seed offspring from coastal and desert populations in common gardens in both environments. Phenotypic differences among populations persisted in both common gardens, suggesting a genetic basis for trait variation. Desert offspring outperformed coastal offspring in the desert garden, suggesting superior adaptation to hot, dry conditions. Herbivore damage was greater for all offspring in the coastal garden. Phenotypic characters also showed plastic responses; all offspring had smaller, more pubescent leaves and more compact growth form in the desert garden. Our results confirm that leaf size and pubescence are heritable characters associated with pronounced variation in plant architecture.  相似文献   

18.
Previous studies revealed significant phenotypic plasticity, genetic variability and population differentiation of flower morphometric traits on dwarf bearded iris Iris pumila. Also, study of I. pumila flowering phenology revealed significant impact of habitat type as well as population differentiation for flowering time. Since the flowering time can influence other flower traits, we performed this analysis of flower morphometric traits in three time points during the flower bud ontogenic development in two habitat types (open vs. shaded). Analysis revealed that for most of the traits greater trait values were recorded for open habitat but only on latter time points. For most of the analyzed traits direction of differences in bud stage was the opposite to the direction of differences in mature flower stage detected in previous studies. However, length of the stem, a trait that showed the greatest variability between habitats and populations and therefore greatest genetic differentiation and phenotypic plasticity, was significantly greater in the samples from the late flowering shaded habitat in all time samples, indicating that in case of this trait different mechanisms were involved. Those findings have implications for design of the future studies on I. pumila.  相似文献   

19.
为了研究植物生长季内开花时间对花特征表型选择的影响,我们以青藏高原东缘高寒草地的毛茛状金莲花Trollius ranunculoides)为实验材料,在生长季内不同开花时间(花前期、花末期)测定花特征,观察访花昆虫的类群和访花频率,生长季结束后收集种子.根据昆虫访花的喜好和季节内类群与访花频率的变化,分析了不同开花时间毛茛状金莲花的花特征与昆虫的选择;并用种子产量表示雌性适合度,估计了毛茛状金莲花的花特征在不同开花时间所受的表型选择.结果表明:不同花期植物的花特征有显著差异,相应的访花昆虫的类群和频率也存在差异,不同类群昆虫访花喜好也不一样.蜂喜好花瓣和花萼较宽、花茎短和花茎数少的个体,这正符合花前期的特征,因而蜂的访花频率在花前期较高;蝇对花特征没有明显的偏好.而通过雌性适合度估计毛茛状金莲花花特征所受的表型选择则是:花前期,花茎较长和花茎数多的植株适合度大;花末期,花茎数多的植株适合度大.我们的研究表明:在植物生长季,花期的分化伴随着传粉昆虫活动的变化.不同花期,访花昆虫的变化可能对植物花特征的分化起了至关重要的作用.但是访花昆虫对花特征的选择与通过雌性适合度估计植物受到的选择不尽相同,这可能是由于其他因素造成的.  相似文献   

20.
Drawing a direct connection between adaptive evolution at the phenotypic level and underlying genetic factors has long been a major goal of evolutionary biologists, but the genetic characterization of adaptive traits in natural populations is notoriously difficult. The study of evolution in experimental populations offers some help — initial conditions are known and changes can be tracked for extended periods under conditions more controlled than wild populations and more realistic than laboratory or greenhouse experiments. In this issue of Molecular Ecology , researchers studying experimental wheat populations over a 12-year period have demonstrated evolution in a major adaptive trait, flowering time, and parallel changes in underlying genetic variation ( Rhoné et al . 2008 ). Their work suggests that cis -regulatory mutations at a single gene may explain most of the flowering time variation in these populations.  相似文献   

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