The carbon economics of vegetative phase change.

Across plant species and biomes, a conserved set of leaf traits govern the economic strategy used to assimilate and invest carbon. As plants age, they face new challenges that may require shifts in this leaf economic strategy. In this study, we investigate the role of the developmental transition, vegetative phase change (VPC), in altering carbon economics as plants age. We used overexpression of microRNA 156 (miR156), the master regulator of VPC, to modulate the timing of VPC in Populus tremula x alba, Arabidopsis thaliana and Zea mays to understand the impact of this transition on leaf economic traits, including construction cost, payback time and return on investment. Here, we find that VPC causes a shift from a low-cost, quick return juvenile strategy to a high-cost, high-return adult strategy. The juvenile strategy is advantageous in light-limited conditions, whereas the adult strategy provides greater returns in high light. The transition between these strategies is correlated with the developmental decline in the level of miR156, suggesting that is regulated by the miR156/SPL pathway. Our results provide an ecophysiological explanation for the existence of juvenile and adult leaf types and suggest that natural selection for these alternative economic strategies could be an important factor in plant evolution.

MicroRNA156-mediated changes in leaf composition lead to altered photosynthetic traits during vegetative phase change.

Plant morphology and physiology change with growth and development. Some of these changes are due to change in plant size and some are the result of genetically programmed developmental transitions. In this study we investigate the role of the developmental transition, vegetative phase change (VPC), on morphological and photosynthetic changes. We used overexpression of microRNA156, the master regulator of VPC, to modulate the timing of VPC in Populus tremula × alba, Zea mays, and Arabidopsis thaliana to determine its role in trait variation independent of changes in size and overall age. Here, we find that juvenile and adult leaves in all three species photosynthesize at different rates and that these differences are due to phase-dependent changes in specific leaf area (SLA) and leaf N but not photosynthetic biochemistry. Further, we found juvenile leaves with high SLA were associated with better photosynthetic performance at low light levels. This study establishes a role for VPC in leaf composition and photosynthetic performance across diverse species and environments. Variation in leaf traits due to VPC are likely to provide distinct benefits under specific environments; as a result, selection on the timing of this transition could be a mechanism for environmental adaptation.

Identification of a major QTL that alters flowering time at elevated [CO(2)] in Arabidopsis thaliana.

BACKGROUND: The transition from vegetative to reproductive stages marks a major milestone in plant development. It is clear that global change factors (e.g., increasing [CO(2)] and temperature) have already had and will continue to have a large impact on plant flowering times in the future. Increasing atmospheric [CO(2)] has recently been shown to affect flowering time, and may produce even greater responses than increasing temperature. Much is known about the genes influencing flowering time, although their relevance to changing [CO(2)] is not well understood. Thus, we present the first study to identify QTL (Quantitative Trait Loci) that affect flowering time at elevated [CO(2)] in Arabidopsis thaliana. METHODOLOGY/PRINCIPAL FINDINGS: We developed our mapping population by crossing a genotype previously selected for high fitness at elevated [CO(2)] (SG, Selection Genotype) to a Cape Verde genotype (Cvi-0). SG exhibits delayed flowering at elevated [CO(2)], whereas Cvi-0 is non-responsive to elevated [CO(2)] for flowering time. We mapped one major QTL to the upper portion of chromosome 1 that explains 1/3 of the difference in flowering time between current and elevated [CO(2)] between the SG and Cvi-0 parents. This QTL also alters the stage at which flowering occurs, as determined from higher rosette leaf number at flowering in RILs (Recombinant Inbred Lines) harboring the SG allele. A follow-up study using Arabidopsis mutants for flowering time genes within the significant QTL suggests MOTHER OF FT AND TFL1 (MFT) as a potential candidate gene for altered flowering time at elevated [CO(2)]. CONCLUSION/SIGNIFICANCE: This work sheds light on the underlying genetic architecture that controls flowering time at elevated [CO(2)]. Prior to this work, very little to nothing was known about these mechanisms at the genomic level. Such a broader understanding will be key for better predicting shifts in plant phenology and for developing successful crops for future environments.

Ecotypic responses of switchgrass to altered precipitation.

Anthropogenic climate change is projected to alter precipitation patterns, resulting in novel environments for plants. The responses of dominant plant species (e.g. Panicum virgatum L. (switchgrass)) to climate changes can drive broader ecosystem processes such as primary productivity. Using a rainfall mesocosm facility, three ecotypes of P. virgatum (collected from Kansas, Oklahoma and Texas, USA) were subjected to three precipitation regimes (average, -25%, +25%) to determine the physiological and growth responses to altered precipitation in a common garden setting. Results showed mean maximum photosynthetic rates, stomatal conductance, transpiration, midday water potential and dark-adapted chlorophyll fluorescence were lowest in the Kansas ecotypes. Increased precipitation treatments raised the mean midday water potentials and lowered water-use efficiency. Aboveground biomass responded positively to changes in precipitation, but flowering initiation was later and rates were lower for Texas ecotypes. In general, ecotype origin was a better predictor of differences in physiological responses and flowering, whereas the precipitation treatments had greater control over biomass production. Depending on the growth variable measured, these results show responses for P. virgatum are under varying ecotypic or environmental control with few interactions, suggesting that future predictions to climate change need not inherently consider localised adaptations in this economically important and widely distributed species.

Elevated CO2 influences the expression of floral-initiation genes in Arabidopsis thaliana.

Atmospheric CO(2) concentration ([CO(2)]) is rising on a global scale and is known to affect flowering time. Elevated [CO(2)] may be as influential as temperature in determining future changes in plant developmental timing, but little is known about the molecular mechanisms that control altered flowering times at elevated [CO(2)]. Using Arabidopsis thaliana, the expression patterns were compared of floral-initiation genes between a genotype that was selected for high fitness at elevated [CO(2)] and a nonselected control genotype. The selected genotype exhibits pronounced delays in flowering time when grown at elevated [CO(2)], whereas the control genotype is unaffected by elevated [CO(2)]. Thus, this comparison provides an evolutionarily relevant system for gaining insight into the responses of plants to future increases in [CO(2)]. Evidence is provided that elevated [CO(2)] influences the expression of floral-initiation genes. In addition, it is shown that delayed flowering at elevated [CO(2)] is associated with sustained expression of the floral repressor gene, FLOWERING LOCUS C (FLC), in an elevated CO(2)-adapted genotype. Understanding the mechanisms that account for changes in plant developmental timing at elevated [CO(2)] is critical for predicting the responses of plants to a high-CO(2) world of the near future.

Flowering time and elevated atmospheric CO2.

Flowering is a critical milestone in the life cycle of plants, and changes in the timing of flowering may alter processes at the species, community and ecosystem levels. Therefore understanding flowering-time responses to global change drivers, such as elevated atmospheric carbon dioxide concentrations, [CO(2)], is necessary to predict the impacts of global change on natural and agricultural ecosystems. Here we summarize the results of 60 studies reporting flowering-time responses (defined as the time to first visible flower) of both crop and wild species at elevated [CO(2)]. These studies suggest that elevated [CO(2)] will influence flowering time in the future. In addition, interactions between elevated [CO(2)] and other global change factors may further complicate our ability to predict changes in flowering time. One approach to overcoming this problem is to elucidate the primary mechanisms that control flowering-time responses to elevated [CO(2)]. Unfortunately, the mechanisms controlling these responses are not known. However, past work has indicated that carbon metabolism exerts partial control on flowering time, and therefore may be involved in elevated [CO(2)]-induced changes in flowering time. This review also indicates the need for more studies addressing the effects of global change drivers on developmental processes in plants.

Photosynthetic responses of forest understory tree species to long-term exposure to elevated carbon dioxide concentration at the Duke Forest FACE experiment.

We examined the photosynthetic responses of four species of saplings growing in the understory of the Duke Forest FACE experiment during the seventh year of exposure to elevated CO2 concentration ([CO2]). Saplings of these same species were measured in the first year of the Duke Forest FACE experiment and at that time showed only seasonal fluctuations in acclimation of photosynthesis to elevated [CO2]. Based on observations from the Duke Forest FACE experiment, we hypothesized that after seven years of exposure to elevated [CO2] significant photosynthetic down-regulation would be observed in these tree species. To test our hypothesis, photosynthetic CO2-response and light-response curves, along with chlorophyll fluorescence, chlorophyll concentration and foliar N were measured twice during the summer of 2003. Exposure to elevated [CO2] continued to increase photosynthesis in all species measured after seven years of treatment with the greatest photosynthetic increase observed near saturating irradiances. In all species, elevated [CO2] increased electron transport efficiency but did not significantly alter carboxylation efficiency. Quantum yield estimated by light curves, chlorophyll concentration, and foliar nitrogen concentrations were unaffected by elevated [CO2]. Contrary to our hypothesis, there is little evidence of progressive N limitation of leaf-level processes in these understory tree species after seven years of exposure to elevated [CO2] in the Duke Forest FACE experiment.

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide.

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].