In his recent review of the developing theory of insect-plant interactions, Gilbert (52) identified four major thrusts in research: insect-plant coevolution, host plants as islands, plant apparency and chemical defense, and resource predictability versus evolutionary strategies of insects. From his review it is evident that, with a few exceptions (23, 45, 53, 64), devdoping theory is addressing primarily a two trophie levd system. In reality, of course, all terrestrial communities based on living plants are composed of at least three interacting trophic levels: plants, herbivores, and natural enemies of herbivores. We argue that theory on insect-plant interactions cannot progress realistically without consideration of the third trophic level. A closer look at the mechanisms of interactions reveals a paradox, and plants have many effects, direct and indirect, positive and negative, not only on herbivores but also on the enemies of herbivores. The third trophic level must be considered as part of a plant's battery of defenses against herbivores.
Ongoing climate change has affected the ecological dynamics of many species and is expected to impose natural selection on ecologically important traits. Droughts and other anticipated changes in precipitation may be particularly potent selective factors, especially in arid regions. Here we demonstrate the evolutionary response of an annual plant, Brassica rapa, to a recent climate fluctuation resulting in a multiyear drought. Ancestral (predrought) genotypes were recovered from stored seed and raised under a set of common environments with descendant (postdrought) genotypes and with ancestor؋descendant hybrids. As predicted, the abbreviated growing seasons caused by drought led to the evolution of earlier onset of flowering. Descendants bloomed earlier than ancestors, advancing first flowering by 1.9 days in one study population and 8.6 days in another. The intermediate flowering time of ancestor؋descendant hybrids supports an additive genetic basis for divergence. Experiments confirmed that summer drought selected for early flowering, that flowering time was heritable, and that selection intensities in the field were more than sufficient to account for the observed evolutionary change. Natural selection for drought escape thus appears to have caused adaptive evolution in just a few generations. A systematic effort to collect and store propagules from suitable species would provide biologists with materials to detect and elucidate the genetic basis of further evolutionary shifts driven by climate change.contemporary evolution ͉ global climate change ͉ life history theory ͉ local adaptation ͉ plant phenology M any species have shifted phenology (the seasonal timing of reproduction and other life history events) in response to ongoing climate change (1-3). For example, a recent study reviewing flowering times (FT) in 461 plant species showed a trend of earlier flowering with climate warming (1), and another study showed shifts in plant flowering and bird and butterfly arrival dates in Mediterranean habitats (4). These shifts are largely attributed to rising temperatures, but anticipated changes in precipitation (5) may also affect phenology, especially in arid regions. Observed shifts in phenology are due in part to direct effects of climate on physiological and developmental rates (phenotypic plasticity). However, climate change can impose natural selection on phenology and thereby cause genetically based evolutionary shifts. These shifts may occur rapidly, providing important opportunities for the study of adaptive evolution in natural populations.Abundant evidence has accumulated over the past several decades showing that natural selection can cause evolutionary change in just a few generations (6, 7). Several cases of contemporary evolution implicate climate change as a selective agent, using two general protocols. The first compares contemporary and previous data on natural populations. This approach has shown shifts over the past few decades in the frequencies of climate-associated isozyme alleles and chrom...
The role of biotic interactions in shaping plant flowering phenology has long been controversial; plastic responses to the abiotic environment, limited precision of biological clocks and inconsistency of selection pressures have generally been emphasized to explain phenological variation. However, part of this variation is heritable and selection analyses show that biotic interactions can modulate selection on flowering phenology. Our review of the literature indicates that pollinators tend to favour peak or earlier flowering, whereas pre-dispersal seed predators tend to favour off-peak or later flowering. However, effects strongly vary among study systems. To understand such variation, future studies should address the impact of mutualist and antagonist dispersal ability, ecological specialization, and habitat and plant population characteristics. Here, we outline future directions to study how such interactions shape flowering phenology. IntroductionFor plant reproduction, timing is everything. An individual plant that flowers too early, before it has had time to accumulate sufficient material resources, will have a limited capacity for seed production. One that delays flowering might gain higher capacity, but might also run out of time to use it before the end of the season. Flowering phenology is affected by many environmental factors, among which temperature and photoperiod, which are reliable signals of seasons, are probably the best studied. Accurate detection of such environmental cues and the resulting plastic response of plants enable flowering to occur when climatic conditions are most suitable for reproduction. Thus, resources and conditions impose bottom-up selective forces on phenology.By contrast, top-down forces act on reproductive timing, particularly those imposed by mutualists (pollinators and seed dispersers) and antagonists (floral pathogens and predispersal seed predators). Here, we review recent progress in understanding some of the top-down selective forces that act on reproductive timing. We highlight what is known,
It has been argued from first principles that plants mate assortatively by flowering time. However, there have been very few studies of phenological assortative mating, perhaps because current methods to infer paternal phenotype are difficult to apply to natural populations. Two methods are presented to estimate the phenotypic correlation between mates-the quantitative genetic metric for assortative mating-for phenological traits. The first method uses individual flowering schedules to estimate mating probabilities for every potential pairing in a sample. These probabilities are then incorporated into a weighted phenotypic correlation between all potential mates and thus yield a prospective estimate based on mating opportunities. The correlation between mates can also be estimated retrospectively by comparing the regression of offspring phenotype over one parent, which is inflated by assortative mating, to the regression over mid-parent, which is not. In a demonstration experiment with Brassica rapa, the prospective correlation between flowering times (days from germination to anthesis) of pollen recipients and their potential donors was 0.58. The retrospective estimate of this correlation strongly agreed with the prospective estimate. The prospective method is easily employed in field studies that explore the effect of phenological assortative mating on selection response and population differentiation.
The resurrection approach of reviving ancestors from stored propagules and comparing them with descendants under common conditions has emerged as a powerful method of detecting and characterizing contemporary evolution. As climatic and other environmental conditions continue to change at a rapid pace, this approach is becoming particularly useful for predicting and monitoring evolutionary responses. We evaluate this approach, explain the advantages and limitations, suggest best practices for implementation, review studies in which this approach has been used, and explore how it can be incorporated into conservation and management efforts. We find that although the approach has thus far been used in a limited number of cases, these studies have provided strong evidence for rapid contemporary adaptive evolution in a variety of systems, particularly in response to anthropogenic environmental change, although it is far from clear that evolution will be able to rescue many populations from extinction given current rates of global changes. We also highlight one effort, known as Project Baseline, to create a collection of stored seeds that can take advantage of the resurrection approach to examine evolutionary responses to environmental change over the coming decades. We conclude that the resurrection approach is a useful tool that could be more widely employed to examine basic questions about evolution in natural populations and to assist in the conservation and management of these populations as they face continued environmental change.
Climate change is likely to spur rapid evolution, potentially altering integrated suites of life‐history traits. We examined evolutionary change in multiple life‐history traits of the annual plant Brassica rapa collected before and after a recent 5‐year drought in southern California. We used a direct approach to examining evolutionary change by comparing ancestors and descendants. Collections were made from two populations varying in average soil moisture levels, and lines propagated from the collected seeds were grown in a greenhouse and experimentally subjected to conditions simulating either drought (short growing season) or high precipitation (long growing season) years. Comparing ancestors and descendants, we found that the drought caused many changes in life‐history traits, including a shift to earlier flowering, longer duration of flowering, reduced peak flowering and greater skew of the flowering schedule. Descendants had thinner stems and fewer leaf nodes at the time of flowering than ancestors, indicating that the drought selected for plants that flowered at a smaller size and earlier ontogenetic stage rather than selecting for plants to develop more rapidly. Thus, there was not evidence for absolute developmental constraints to flowering time evolution. Common principal component analyses showed substantial differences in the matrix of trait covariances both between short and long growing season treatments and between populations. Although the covariances matrices were generally similar between ancestors and descendants, there was evidence for complex evolutionary changes in the relationships among the traits, and these changes depended on the population and treatment. These results show that a full appreciation of the impacts of global change on phenotypic evolution will entail an understanding of how changes in climatic conditions affect trait values and the structure of relationships among traits.
As climate changes at unprecedented rates, understanding population responses is a major challenge. Resurrection studies can provide crucial insights into the contemporary evolution of species to climate change. We used a seed collection of two Californian populations of the annual plant Brassica rapa made over two decades of dramatic precipitation fluctuations, including increasingly severe droughts. We compared flowering phenology, other drought response traits, and seed production among four generations, grown under drought and control conditions, to test for evolutionary change and to characterize the strength and direction of selection. Postdrought generations flowered earlier, with a reduced stem diameter, and lower water-use efficiency (WUE), while intervening wet seasons reversed these adaptations. There was selection for earlier flowering, which was adaptive, but delayed flowering after wet years resulted in reduced total seed mass, indicating a maladaptive response caused by brief wet periods. Furthermore, evolutionary changes and plastic responses often differed in magnitude between populations and drought periods, suggesting independent adaptive pathways. While B. rapa rapidly evolved a drought escape strategy, plant fitness was reduced in contemporary generations, suggesting that rapid shifts in flowering time may no longer keep up with the increasing severity of drought periods, especially when drought adaptation is slowed by occasional wet seasons.
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