Plant phenology, the annually recurring sequence of plant developmental stages, is important for plant functioning and ecosystem services and their biophysical and biogeochemical feedbacks to the climate system. Plant phenology depends on temperature, and the current rapid climate change has revived interest in understanding and modeling the responses of plant phenology to the warming trend and the consequences thereof for ecosystems. Here, we review recent progresses in plant phenology and its interactions with climate change. Focusing on the start (leaf unfolding) and end (leaf coloring) of plant growing seasons, we show that the recent rapid expansion in ground- and remote sensing- based phenology data acquisition has been highly beneficial and has supported major advances in plant phenology research. Studies using multiple data sources and methods generally agree on the trends of advanced leaf unfolding and delayed leaf coloring due to climate change, yet these trends appear to have decelerated or even reversed in recent years. Our understanding of the mechanisms underlying the plant phenology responses to climate warming is still limited. The interactions between multiple drivers complicate the modeling and prediction of plant phenology changes. Furthermore, changes in plant phenology have important implications for ecosystem carbon cycles and ecosystem feedbacks to climate, yet the quantification of such impacts remains challenging. We suggest that future studies should primarily focus on using new observation tools to improve the understanding of tropical plant phenology, on improving process-based phenology modeling, and on the scaling of phenology from species to landscape-level.

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Plant phenology and global climate change: Current progresses and challenges

Contents Summary 32 I. The importance of plant carbon metabolism for climate change 32 II. Rising atmospheric CO2 and carbon metabolism 33 III. Rising temperatures and carbon metabolism 37 IV. Thermal acclimation responses of carbon metabolic processes can be best understood when studied together 38 V. Will elevated CO2 offset warming-induced changes in carbon metabolism? 40 VI. No plant is an island: water and nutrient limitations define plant responses to climate drivers 41 VII. Conclusions 42 Acknowledgements 42 References 42 Appendix A1 48 SUMMARY: Plant carbon metabolism is impacted by rising CO2 concentrations and temperatures, but also feeds back onto the climate system to help determine the trajectory of future climate change. Here we review how photosynthesis, photorespiration and respiration are affected by increasing atmospheric CO2 concentrations and climate warming, both separately and in combination. We also compile data from the literature on plants grown at multiple temperatures, focusing on net CO2 assimilation rates and leaf dark respiration rates measured at the growth temperature (Agrowth and Rgrowth , respectively). Our analyses show that the ratio of Agrowth to Rgrowth is generally homeostatic across a wide range of species and growth temperatures, and that species that have reduced Agrowth at higher growth temperatures also tend to have reduced Rgrowth , while species that show stimulations in Agrowth under warming tend to have higher Rgrowth in the hotter environment. These results highlight the need to study these physiological processes together to better predict how vegetation carbon metabolism will respond to climate change.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.15283

Plant carbon metabolism and climate change: elevated CO2 and temperature impacts on photosynthesis, photorespiration and respiration.

To realistically simulate climate feedbacks from the land surface to the atmosphere, models must replicate the responses of plants to environmental changes. Several processes, operating at various scales, cause the responses of photosynthesis and plant respiration to temperature and CO2 to change over time of exposure to new or changing environmental conditions. Here, we review the latest empirical evidence that short-term responses of plant carbon exchange rates to temperature and CO2 are modified by plant photosynthetic and respiratory acclimation as well as biogeochemical feedbacks. We assess the frequency with which these responses have been incorporated into vegetation models, and highlight recently designed algorithms that can facilitate their incorporation. Few models currently include representations of the long-term plant responses that have been recorded by empirical studies, likely because these responses are still poorly understood at scales relevant for models. Studies show that, at a regional scale, simulated carbon flux between the atmosphere and vegetation can dramatically differ between versions of models that do and do not include acclimation. However, the realism of these results is difficult to evaluate, as algorithm development is still in an early stage, and a limited number of data are available. We provide a series of recommendations that suggest how a combination of empirical and modeling studies can produce mechanistic algorithms that will realistically simulate longer term responses within global-scale models.

/pdf/plant-respiration-and-photosynthesis-in-global-scale-models-1zh9rkekus.pdf

Plant respiration and photosynthesis in global‐scale models: incorporating acclimation to temperature and CO2

The phenological state of vegetation significantly affects exchanges of heat, mass, and momentum between the Earth's surface and the atmosphere. Although current patterns can be estimated from satellites, we lack the ability to predict future trends in response to climate change. We searched the literature for a common set of variables that might be combined into an index to quantify the greenness of vegetation throughout the year. We selected as variables: daylength (photoperiod), evaporative demand (vapor pressure deficit), and suboptimal (minimum) temperatures. For each variable we set threshold limits, within which the relative phenological performance of the vegetation was assumed to vary from inactive (0) to unconstrained (1). A combined Growing Season Index (GSI) was derived as the product of the three indices. Ten-day mean GSI values for nine widely dispersed ecosystems showed good agreement (r>0.8) with the satellite-derived Normalized Difference Vegetation Index (NDVI). We also tested the model at a temperate deciduous forest by comparing model estimates with average field observations of leaf flush and leaf coloration. The mean absolute error of predictions at this site was 3 days for average leaf flush dates and 2 days for leaf coloration dates. Finally, we used this model to produce a global map that distinguishes major differences in regional phenological controls. The model appears sufficiently robust to reconstruct historical variation as well as to forecast future phenological responses to changing climatic conditions.

A generalized, bioclimatic index to predict foliar phenology in response to climate

Autumn remains a relatively neglected season in climate change research in temperate and arctic ecosystems. This neglect occurs despite the importance of autumn events, including leaf senescence, fruit ripening, bird and insect migration, and induction of hibernation and diapause. Changes in autumn phenology alter the reproductive capacity of individuals, exacerbate invasions, allow pathogen amplification and higher disease-transmission rates, reshuffle natural enemy‐prey dynamics, shift the ecological dynamics among interacting species, and affect the net productivity of ecosystems. We synthesize some of our existing understanding of autumn phenology and identify five areas ripe for future climate change research. We provide recommendations to address common pitfalls in autumnal research as well as to support the conservation and management of vulnerable ecosystems and taxa.

http://www.concordmuseum.org/assets/Autumn,%20the%20neglected%20season%20in%20climate%20change%20research.pdf

Autumn, the neglected season in climate change research

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open-top chambers supplied three levels of warming (+0, +2, and +4 C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO2 assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17 to 34 were observed. Across species, acclimation potentials varied from 0.55 C to 1.07 C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responsesmore » should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.« less

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

Predicting forest responses to warming climates relies on assumptions about niche and temperature sensitivity that remain largely untested. Observational studies have related current and historical temperatures to phenological shifts, but experimental evidence is sparse, particularly for autumn responses. A 4 year field experiment exposed four deciduous forest species from contrasting climates (Liquidambar styraciflua, Quercus rubra, Populus grandidentata, and Betula alleghaniensis) to air temperatures 2 and 4 °C above ambient controls, using temperature-controlled open top chambers. Impacts of year-round warming on bud burst (BB), senescence, and abscission were evaluated in relation to thermal provenance. Leaves emerged earlier in all species by an average of 4–9 days at +2 °C and 6–14 days at +4 °C. Magnitude of advance varied with species and year, but was larger for the first 2 °C increment than for the second. Effect of warming increased with early BB, favoring Liquidambar, but even BB of northern species advanced, despite temperatures exceeding those of the realized niche. Treatment differences in BB were inadequately explained by temperature sums alone. In autumn, chlorophyll was retained an average of 4 and 7 days longer in +2 and +4 °C treatments, respectively, and abscission delayed by 8 and 13 days. Growing seasons in the warmer atmospheres averaged 5–18 days (E2) and 6–28 days (E4) longer, according to species, with the least impact in Quercus. Results are compared with a 16 years record of canopy onset and offset in a nearby upland deciduous forest, where BB showed similar responsiveness to spring temperatures (2–4 days °C−1). Offset dates in the stand tracked August–September temperatures, except when late summer drought caused premature senescence. The common garden-like experiment provides evidence that warming alone extends the growing season, at both ends, even if stand-level impacts may be complicated by variation in other environmental factors.

Ashley V. Walker

Papers

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

Forest phenology and a warmer climate – growing season extension in relation to climatic provenance