The Penalty of a Long, Hot Summer. Photosynthetic Acclimation to High CO2 and Continuous Light in “Living Fossil” Conifers

Deciduous forests covered the ice-free polar regions 280 to 40 million years ago under warm “greenhouse” climates and high atmospheric pCO2. Their deciduous habit is frequently interpreted as an adaptation for minimizing carbon losses during winter, but experiments with “living fossils” in a simulated warm polar environment refute this explanation. Measured carbon losses through leaf abscission of deciduous trees are significantly greater than losses through winter respiration in evergreens, yet annual rates of primary productivity are similar in all species. Here, we investigate mechanisms underlying this apparent paradox by measuring the seasonal patterns of leaf photosynthesis (A) under pCO2 enrichment in the same trees. During spring, A increased significantly in coastal redwood (Sequoia sempervirens), dawn redwood (Metasequoia glyptostroboides), and swamp cypress (Taxodium distichum) at an elevated pCO2 of 80 Pa compared with controls at 40 Pa. However, strong acclimation in Rubisco carboxylation capacity (Vc,max) completely offset the CO2 response of A in all species by the end of 6 weeks of continuous illumination in the simulated polar summer. Further measurements demonstrated the temporary nature of acclimation, with increases in Vc,max during autumn restoring the CO2 sensitivity of A. Contrary to expectations, the acclimation of Vc,max was not always accompanied by accumulation of leaf carbohydrates, but was associated with a decline in leaf nitrogen in summer, suggesting an alteration of the balance in plant sources and sinks for carbon and nitrogen. Preliminary calculations using A indicated that winter carbon losses through deciduous leaf abscission and respiration were recovered by 10 to 25 d of canopy carbon fixation during summer, thereby explaining the productivity paradox

The representation of root processes in models addressing the responses of vegetation to global change

The representation of root activity in models is here confined to considerations of applications assessing the impacts of changes in climate or atmospheric [CO2]. Approaches to modelling roots can be classified into four major types: models in which roots are not considered, models in which there is an interplay between only selected above-ground and below-ground processes, models in which growth allocation to all parts of the plants depends on the availability and matching of the capture of external resources, and models with explicit treatments of root growth, architecture and resource capture. All models seem effective in describing the major root activities of water and nutrient uptake, because these processes are highly correlated, particularly at large scales and with slow or equilibrium dynamics. Allocation models can be effective in providing a deeper, perhaps contrary, understanding of the dynamic underpinning to observations made only above ground. The complex and explicit treatment of roots can be achieved only in small-scale highly studied systems because of the requirements for many initialized variables to run the model

Carbon loss by deciduous trees in a CO2-rich ancient polar environment

Fossils demonstrate that deciduous forests covered the polar regions for much of the past 250 million years 1 when the climate was warm and atmospheric CO2 high 2. But the evolutionary significance of their deciduous character has remained a matter of conjecture for almost a century 3. The leading hypothesis 1,4-7 argues that it was an adaptation to photoperiod, allowing the avoidance of carbon losses by respiration from a canopy of leaves unable to photosynthesize in the darkness of warm polar winters 8-11. Here we test this proposal with experiments using 'living fossil' tree species grown in a simulated polar climate with and without CO2 enrichment. We show that the quantity of carbon lost annually by shedding a deciduous canopy is significantly greater than that lost by evergreen trees through wintertime respiration and leaf litter production, irrespective of growth CO2 concentration. Scaling up our experimental observations indicates that the greater expense of being deciduous persists in mature forests, even up to latitudes of 83 [degrees]N, where the duration of the polar winter exceeds five months. We therefore reject the carbon-loss hypothesis as an explanation for the deciduous nature of polar forests

Does Long-Term Elevation of CO2 Concentration Increase Photosynthesis in Forest Floor Vegetation? (Indiana Strawberry in a Maryland Forest).

As the partial pressure of CO2 (pCO2) in the atmosphere rises, photorespiratory loss of carbon in C3 photosynthesis will diminish and the net efficiency of light-limited photosynthetic carbon uptake should rise. We tested this expectation for Indiana strawberry (Duchesnea indica) growing on a Maryland forest floor. Open-top chambers were used to elevate the pCO2 of a forest floor habitat to 67 Pa and were paired with control chambers providing an ambient pCO2 of 38 Pa. After 3.5 years, D. indica leaves grown and measured in the elevated pCO2 showed a significantly greater maximum quantum efficiency of net photosynthesis (by 22%) and a lower light compensation point (by 42%) than leaves grown and measured in the control chambers. The quantum efficiency to minimize photorespiration, measured in 1% O2, was the same for controls and plants grown at elevated pCO2. This showed that the maximum efficiency of light-energy transduction into assimilated carbon was not altered by acclimation and that the increase in light-limited photosynthesis at elevated pCO2 was simply a function of the decrease in photorespiration. Acclimation did decrease the ribulose-1,5-bisphosphate carboxylase/oxygenase and light-harvesting chlorophyll protein content of the leaf by more than 30%. These changes were associated with a decreased capacity for light-saturated, but not light-limited, photosynthesis. Even so, leaves of D. indica grown and measured at elevated pCO2 showed greater light-saturated photosynthetic rates than leaves grown and measured at the current atmospheric pCO2. In situ measurements under natural forest floor lighting showed large increases in leaf photosynthesis at elevated pCO2, relative to controls, in both summer and fall. The increase in efficiency of light-limited photosynthesis with elevated pCO2 allowed positive net photosynthetic carbon uptake on days and at locations on the forest floor that light fluxes were insufficient for positive net photosynthesis in the current atmospheric pCO2

Modelling the recent historical impacts of atmospheric CO2 and climate change on Mediterranean vegetation

During the past century, annual mean temperature has increased by 0.75 degrees C and precipitation has shown marked variation throughout the Mediterranean basin. These historical climate changes may have had significant, but presently undefined, impacts on the productivity and structure of sclerophyllous shrubland, an important vegetation type in the region. We used a vegetation model for this functional type to examine climate change impacts, and their interaction with the concurrent historical rise in atmospheric CO2. Using only climate and soil texture as data inputs, model. predictions showed good agreement with observations of seasonal and regional variation in leaf and canopy physiology, net primary productivity (NPP), leaf area index (LAI) and soil water. Model simulations for shrubland sites indicated that potential NPP has risen by 25% and LAI by 7% during the past century, although the absolute increase in LAI was small. Sensitivity analysis suggested that the increase in atmospheric CO2 since 1900 was the primary cause of these changes, and that simulated climate change alone had negative impacts on both NPP and LAI. Effects of rising CO2 were mediated by significant increases in the efficiency of water-use in NPP throughout the region, as a consequence of the direct effect of CO2 on leaf gas exchange. This increase in efficiency compensated for limitation of NPP by drought, except in areas where drought was most severe. However, while water was used more efficiently, total canopy water loss rose slightly or remained unaffected in model simulations, because increases in LAI with CO2 counteracted the effects of reduced stomatal conductance on transpiration. Model simulations for the Mediterranean region indicate that the recent rise in atmospheric CO2 may already have had significant impacts on productivity, structure and water relations of sclerophyllous shrub vegetation, which tended to offset the detrimental effects of climate change in the region

Chicken Farming in Grassland Increases Environmental Sustainability and Economic Efficiency

Background: Grassland degradation caused by overgrazing poses a threat to both animal husbandry and environmental sustainability in most semi-arid areas especially north China. Although the Chinese Government has made huge efforts to restore degraded grasslands, a considerable attempt has unfortunately failed due to an inadequate consideration of economic benefits to local communities. Methodology/Principal Findings: A controlled field experiment was conducted to test our hypothesis that utilizing natural grasslands as both habitat and feed resources for chickens and replacing the traditional husbandry system with chicken farming would increase environmental sustainability and raise income. Aboveground plant biomass elevated from 25 g m22 for grazing sheep to 84 g m22 for chicken farming. In contrast to the fenced (unstocked) grassland, chicken farming did not significantly decrease aboveground plant biomass, but did increase the root biomass by 60% (p,0.01). Compared with traditional sheep grazing, chicken farming significantly improved soil surface water content (0–10 cm), from 5% to 15%. Chicken farming did not affect the soil bulk density, while the traditional sheep grazing increased the soil bulk density in the 0–10 cm soil layer by 35% of the control (p,0.05). Most importantly, the economic income of local herdsmen has been raised about six times compared with the traditional practice of raising sheep. Ecologically, such an innovative solution allowed large degraded grasslands to naturally regenerate. Grasslands also provided a high quality organic poultry product which could be marketed in big cities. Conclusion/Significance: Chicken farming is an innovative alternative strategy for increasing environmental sustainability and economic income, rather than a challenge to the traditional nomadic pastoral system. Our approach might be technically applicable to other large degraded grasslands of the world, especially in China

Carbon source-sink limitations differ between two Species with Contrasting Growth Strategies.

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximise crop yield. We investigated how differences in growth rate arise from source-sink limitations, using a model system comparing a fast-growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow-growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub-ambient and elevated CO2 concentrations ([CO2 ]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO2 ] indicating that source strength was near maximal at current [CO2 ]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO2 ] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO2 ], and lower non-structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximising growth of elite cereals under future elevated [CO2 ]

Is stimulation of leaf photosynthesis by elevated carbon dioxide concentration maintained in the long term? A test with Lolium perenne grown for 10 years at two nitrogen fertilization levels under Free Air CO2 Enrichment (FACE)

Photosynthesis is commonly stimulated in grasslands with experimental increases in atmospheric CO2 concentration ([CO2]), a physiological response that could significantly alter the future carbon cycle if it persists in the long term.. Yet an acclimation of photosynthetic capacity suggested by theoretical models and short-term experiments could completely remove this effect of CO2. Perennial ryegrass (Lolium perenne L. cv. Bastion) was grown under an elevated [CO2] of 600 mumol mol(-1) for 10 years using Free Air CO2 Enrichment (FACE), with two contrasting nitrogen levels and abrupt changes in the source: sink ratio following periodic harvests. More than 3000 measurements characterized the response of leaf photosynthesis and stomatal conductance to elevated [CO2] across each growing season for the duration of the experiment. Over the 10 years as a whole, growth at elevated [CO2] resulted in a 43% higher rate of light-saturated leaf photosynthesis and a 36% increase in daily integral of leaf CO2 uptake. Photosynthetic stimulation was maintained despite a 30% decrease in stomatal conductance and significant decreases in both the apparent, maximum carboxylation velocity (V-c,V-max) and the maximum rate of electron transport (J(max)). Immediately prior to the periodic (every 4-8 weeks) cuts of the L. perenne stands, V-c,V-max and J(max), were significantly lower in elevated than in ambient [CO2] in the low-nitrogen treatment. This difference was smaller after the cut, suggesting a dependence upon the balance between the sources and sinks for carbon. In contrast with theoretical expectations and the results of shorter duration experiments, the present results provide no significant change in photosynthetic stimulation across a 10-year period, nor greater acclimation in V-c,V-max and J(max) in the later years in either nitrogen treatment

Crop origins explain variation in global agricultural relevance

Human food production is dominated globally by a small number of crops. Why certain crops have attained high agricultural relevance while others have remained minor might partially stem from their different origins. Here, we analyse a dataset of 866 crops to show that seed crops and species originating from seasonally dry environments tend to have the greatest agricultural relevance, while phylogenetic affinities play a minor role. These patterns are nuanced by root and leaf crops and herbaceous fruit crops having older origins in the aseasonal tropics. Interestingly, after accounting for these effects, we find that older crops are more likely to be globally important and are cultivated over larger geographical areas than crops of recent origin. Historical processes have therefore left a pervasive global legacy on the food we eat today

Stomatal dynamics in Alloteropsis semialata arise from the evolving interplay between photosynthetic physiology, stomatal size and biochemistry

C4 plants are expected to have faster stomatal movements than C3 species because they tend to have smaller guard cells. However, little is known about how the evolution of C4 photosynthesis influences stomatal dynamics in relation to guard cell size and environmental factors. We studied photosynthetically diverse populations of the grass Alloteropsis semialata, showing that the origin of C4 photosynthesis in this species was associated with a shortening of stomatal guard and subsidiary cells. However, for a given cell size, C4 and C3–C4 intermediate individuals had similar or slower light-induced stomatal opening speeds than C3 individuals. Conversely, when exposed to decreasing light, stomata in C4 plants closed as fast as those in non-C4 plants. Polyploid formation in some C4 plants led to larger stomatal cells and was associated with slower stomatal opening. Conversely, diversification of C4 diploid plants into wetter environments was associated with an acceleration of stomatal opening. Overall, there was significant relationship between light-saturated photosynthesis and stomatal opening speed in the C4 plants, implying that photosynthetic energy production was limiting for stomatal opening. Stomatal dynamics in this wild grass therefore arise from the evolving interplay between photosynthetic physiology and the size and biochemical function of stomatal complexes