Warming experiments underpredict plant phenological responses to climate change.

Warming experiments are increasingly relied on to estimate plant responses to global climate change. For experiments to provide meaningful predictions of future responses, they should reflect the empirical record of responses to temperature variability and recent warming, including advances in the timing of flowering and leafing. We compared phenology (the timing of recurring life history events) in observational studies and warming experiments spanning four continents and 1,634 plant species using a common measure of temperature sensitivity (change in days per degree Celsius). We show that warming experiments underpredict advances in the timing of flowering and leafing by 8.5-fold and 4.0-fold, respectively, compared with long-term observations. For species that were common to both study types, the experimental results did not match the observational data in sign or magnitude. The observational data also showed that species that flower earliest in the spring have the highest temperature sensitivities, but this trend was not reflected in the experimental data. These significant mismatches seem to be unrelated to the study length or to the degree of manipulated warming in experiments. The discrepancy between experiments and observations, however, could arise from complex interactions among multiple drivers in the observational data, or it could arise from remediable artefacts in the experiments that result in lower irradiance and drier soils, thus dampening the phenological responses to manipulated warming. Our results introduce uncertainty into ecosystem models that are informed solely by experiments and suggest that responses to climate change that are predicted using such models should be re-evaluated.

Relatedness of Macrophomina phaseolina isolates from tallgrass prairie, maize, soybean and sorghum.

Agricultural and wild ecosystems may interact through shared pathogens such as Macrophomina phaseolina, a generalist clonal fungus with more than 284 plant hosts that is likely to become more important under climate change scenarios of increased heat and drought stress. To evaluate the degree of subdivision in populations of M. phaseolina in Kansas agriculture and wildlands, we compared 143 isolates from maize fields adjacent to tallgrass prairie, nearby sorghum fields, widely dispersed soybean fields and isolates from eight plant species in tallgrass prairie. Isolate growth phenotypes were evaluated on a medium containing chlorate. Genetic characteristics were analysed based on amplified fragment length polymorphisms and the sequence of the rDNA-internal transcribed spacer (ITS) region. The average genetic similarity was 58% among isolates in the tallgrass prairie, 71% in the maize fields, 75% in the sorghum fields and 80% in the dispersed soybean fields. The isolates were divided into four clusters: one containing most of the isolates from maize and soybean, two others containing isolates from wild plants and sorghum, and a fourth containing a single isolate recovered from Solidago canadensis in the tallgrass prairie. Most of the sorghum isolates had the dense phenotype on media containing chlorate, while those from other hosts had either feathery or restricted phenotypes. These results suggest that the tallgrass prairie supports a more diverse population of M. phaseolina per area than do any of the crop species. Subpopulations show incomplete specialization by host. These results also suggest that inoculum produced in agriculture may influence tallgrass prairie communities, and conversely that different pathogen subpopulations in tallgrass prairie can interact there to generate 'hybrids' with novel genetic profiles and pathogenic capabilities.

Climate change effects on plant disease: genomes to ecosystems.

Research in the effects of climate change on plant disease continues to be limited, but some striking progress has been made. At the genomic level, advances in technologies for the high-throughput analysis of gene expression have made it possible to begin discriminating responses to different biotic and abiotic stressors and potential trade-offs in responses. At the scale of the individual plant, enough experiments have been performed to begin synthesizing the effects of climate variables on infection rates, though pathosystem-specific characteristics make synthesis challenging. Models of plant disease have now been developed to incorporate more sophisticated climate predictions. At the population level, the adaptive potential of plant and pathogen populations may prove to be one of the most important predictors of the magnitude of climate change effects. Ecosystem ecologists are now addressing the role of plant disease in ecosystem processes and the challenge of scaling up from individual infection probabilities to epidemics and broader impacts.

Trade-offs between male and female reproduction associated with allozyme variation in phosphoglucoisomerase in an annual plant (Clarkia unguiculata: Onagraceae).

The genotype of an individual for allozymes such as phosphoglucoisomerase (Pgi) is often not neutral with regard to fitness. Studies of several taxa have found consistent fitness differences among Pgi genotypes expressing different allozymes. We conducted a greenhouse experiment with Clarkia unguiculata to determine whether allelic variation at the Pgi-C1 locus may affect components of male and female function. We found significant differences in siring success between pollen donors homozygous for different Pgi alleles. When a mixture of pollen was applied to stigmas under conditions of gametophytic competition (more pollen deposited on stigmas than there are ovules available to fertilize), donors homozygous for the C allele of Pgi sired more seeds per fruit than B-allele donors. Differences between genotypes with respect to female fertility per fruit contrasted with the male advantage associated with the C allele. Recipients homozygous for the C allele produced fruits with more aborted seeds and fewer viable seeds than recipients homozygous for the B allele. These results suggest that allelic variation at a single locus may have opposing effects on male and female reproductive success in C. unguiculata, and that trade-offs between the two types of reproductive success could contribute to the maintenance of variation at the Pgi-C1 locus.

Differential siring success of Pgi genotypes in Clarkia Unguiculata (Onagraceae).

Competition among pollen grains for the fertilization of ovules can play an important role in determining the male and female reproductive success of flowering plants. To examine the influence of pollen-donor genotype on male reproductive success, hand-pollinations were conducted on Clarkia unguiculata and the siring success of pollen-donor plants was compared between donors homozygous for different allelomorphs of the allozyme PGI (phosphoglucoisomerase). Donors homozygous for the B allele sired more seeds than C-allele donors. Single-donor crosses indicated that C-donor-sired seeds are aborted more often than are B-donor-sired seeds, suggesting that the B-allele donor's advantage in mixed pollinations was a result of differential abortion. A negative relationship between pollen load and the siring success of B-allele donors implies that pollen from B-allele donors has reduced performance relative to C-allele donors when pollen loads are high. These data demonstrate consistent differences in siring success between individuals homozygous for different alleles at a single locus and suggest that variation at the Pgi locus may be maintained by a post-pollination trade-off.

The absence of cryptic self-incompatibility in Clarkia unguiculata (Onagraceae).

Many species exhibit reduced siring success of self-relative to outcross-pollen donors. This can be attributed either to postfertilization abortion of selfed ovules or to cryptic self-incompatibility (CSI). CSI is a form of self-incompatibility whereby the advantage to outcross pollen is expressed only following pollinations where there is gametophytic competition between self and outcross pollen. Under the definition of CSI, this differential success is due to the superior prefertilization performance (pollen germination rate and pollen tube growth rate) of outcross pollen relative to self pollen. Although CSI has been demonstrated in several plant species, no studies have assessed among-population variation in the expression of CSI. We conducted a greenhouse study on Clarkia unguiculata (an annual species with a mixed-mating system) to detect CSI, and we compare our observations to previous reports of CSI in C. gracilis and another population of C. unguiculata. In contrast to these previous studies of CSI in Clarkia, we used genetic rather than phenotypic markers to measure the relative performance of selfed vs. outcross pollen. In this study, we measured the intensity of CSI in C. unguiculata from a large population in southern California and we determined whether the magnitude of pollen competition (manipulated by controlling the number of pollen grains deposited on a stigma) influenced the outcome of competition between self and outcross pollen. In contrast to previous investigations of Clarkia, we found no evidence for CSI. The mean number of seeds sired per fruit did not differ between self and outcross pollen following either single-donor or mixed pollinations. In addition, the relative success of selfed vs. outcross pollen was independent of the magnitude of pollen competition. These results suggest that: (1) one of the few nonheterostylous species previously thought to be cryptically self-incompatible is completely self-compatible (at least in the population studied here) or (2) phenotypic markers may be problematic for the detection of CSI.