Revisiting RuBisCO

Since the discovery of its role in the CO₂ fixation reaction in photosynthesis, RuBisCO has been one of the most extensively researched enzymes in the fields of biochemistry, molecular biology, and molecular genetics as well as conventional plant physiology, agricultural chemistry, and crop science. In addition, the RuBisCO and RuBisCO-like genes of more than 2000 organisms have been sequenced during the past 20 years. During the course of those studies, the origin of the RuBisCO gene began to be discussed. Recent studies have reported that the RuBisCO gene emerged in methanogenic bacteria long before photosynthetic organisms appeared. The origin of similar early genes might have allowed this gene to overcome changes in global environments during ancient and recent eras and to participate in the fixation of 200 GT of CO₂ annually. In this review, I focus on several points that have not been discussed at length in the literature thus far.     

Effects of RuBisCO and CO2 concentration on cyanobacterial growth and carbon isotope fractionation

Carbon isotope biosignatures preserved in the Precambrian geologic record are primarily interpreted to reflect ancient cyanobacterial carbon fixation catalyzed by Form I RuBisCO enzymes. The average range of isotopic biosignatures generally follows that produced by extant cyanobacteria. However, this observation is difficult to reconcile with several environmental (e.g., temperature, pH, and CO₂ concentrations), molecular, and physiological factors that likely would have differed during the Precambrian and can produce fractionation variability in contemporary organisms that meets or exceeds that observed in the geologic record. To test a specific range of genetic and environmental factors that may impact ancient carbon isotope biosignatures, we engineered a mutant strain of the model cyanobacterium Synechococcus elongatus PCC 7942 that overexpresses RuBisCO across varying atmospheric CO₂ concentrations. We hypothesized that changes in RuBisCO expression would impact the net rates of intracellular CO₂ fixation versus CO₂ supply, and thus whole-cell carbon isotope discrimination. In particular, we investigated the impacts of RuBisCO overexpression under changing CO₂ concentrations on both carbon isotope biosignatures and cyanobacterial physiology, including cell growth and oxygen evolution rates. We found that an increased pool of active RuBisCO does not significantly affect the ¹³C/¹²C isotopic discrimination (ε_p) at all tested CO₂ concentrations, yielding ε_p of ≈ 23‰ for both wild-type and mutant strains at elevated CO₂. We therefore suggest that expected variation in cyanobacterial RuBisCO expression patterns should not confound carbon isotope biosignature interpretation. A deeper understanding of environmental, evolutionary, and intracellular factors that impact cyanobacterial physiology and isotope discrimination is crucial for reconciling microbially driven carbon biosignatures with those preserved in the geologic record.     

FACE-ing the global change: Opportunities for improvement in photosynthetic radiation use efficiency and crop yield

The earth is rapidly changing through processes such as rising [CO₂], [O₃], and increased food demand. By 2050 the projected atmospheric [CO₂] and ground level [O₃] will be 50% and 20% higher than today. To meet future agricultural demand, amplified by an increasing population and economic progress in developing countries, crop yields will have to increase by at least 50% by the middle of the century. FACE (Free Air Concentration Enrichment) experiments have been conducted for more than 20 years in various parts of world to estimate, under the most realistic agricultural conditions possible, the impact of the CO₂ levels projected for the middle of this century on crops. The stimulations of crop seed yields by the projected CO₂ levels across FACE studies are about 18% on average and up to 30% for the hybrid rice varieties and vary among crops, cultivars, nitrogen levels and soil moisture. The observed increase in crop yields under the projected CO₂ levels fall short of what would be required to meet the projected future food demand, even with the most responsive varieties. Crop biomass production and seed yield is the product of photosynthetic solar energy conversion. Improvement in photosynthetic radiation use efficiency stands as the most promising opportunity allowing for major increases in crop yield in a future that portends major changes in climate and crop growing environments. Our advanced understanding of the photosynthetic process along with rapidly advancing capabilities in functional genomics, genetic transformation and synthetic biology promises new opportunities for crop improvement by greater photosynthesis and crop yield. Traits and genes that show promise for improving photosynthesis are briefly reviewed, including enhancing leaf photosynthesis capacity and reducing photorespiration loss, manipulating plant hormones’ responses for better ideotypes, extending duration of photosynthesis, and increasing carbon partitioning to the sink to alleviate feedback inhibition of photosynthesis.     

Evolutionary trends in RuBisCO kinetics and their co‐evolution with CO2 concentrating mechanisms

RuBisCO‐catalyzed CO₂ fixation is the main source of organic carbon in the biosphere. This enzyme is present in all domains of life in different forms (III, II, and I) and its origin goes back to 3500 Mya, when the atmosphere was anoxygenic. However, the RuBisCO active site also catalyzes oxygenation of ribulose 1,5‐bisphosphate, therefore, the development of oxygenic photosynthesis and the subsequent oxygen‐rich atmosphere promoted the appearance of CO₂ concentrating mechanisms (CCMs) and/or the evolution of a more CO₂‐specific RuBisCO enzyme. The wide variability in RuBisCO kinetic traits of extant organisms reveals a history of adaptation to the prevailing CO₂/O₂ concentrations and the thermal environment throughout evolution. Notable differences in the kinetic parameters are found among the different forms of RuBisCO, but the differences are also associated with the presence and type of CCMs within each form, indicative of co‐evolution of RuBisCO and CCMs. Trade‐offs between RuBisCO kinetic traits vary among the RuBisCO forms and also among phylogenetic groups within the same form. These results suggest that different biochemical and structural constraints have operated on each type of RuBisCO during evolution, probably reflecting different environmental selective pressures. In a similar way, variations in carbon isotopic fractionation of the enzyme point to significant differences in its relationship to the CO₂ specificity among different RuBisCO forms. A deeper knowledge of the natural variability of RuBisCO catalytic traits and the chemical mechanism of RuBisCO carboxylation and oxygenation reactions raises the possibility of finding unrevealed landscapes in RuBisCO evolution.     

RAF2 is a RuBisCO assembly factor in Arabidopsis thaliana

Ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RuBisCO) catalyzes the reaction between gaseous carbon dioxide (CO₂) and ribulose‐1,5‐bisphosphate. Although it is one of the most studied enzymes, the assembly mechanisms of the large hexadecameric RuBisCO is still emerging. In bacteria and in the C4 plant Zea mays, a protein with distant homology to p terin‐4α‐c arbinolamine d ehydratase (PCD) has recently been shown to be involved in RuBisCO assembly. However, studies of the homologous PCD‐like protein (RAF2, RuBisCO assembly factor 2) in the C3 plant Arabidopsis thaliana (A. thaliana) have so far focused on its role in hormone and stress signaling. We investigated whether A. thalianaRAF2 is also involved in RuBisCO assembly. We localized RAF2 to the soluble chloroplast stroma and demonstrated that raf2 A. thaliana mutant plants display a severe pale green phenotype with reduced levels of stromal RuBisCO. We concluded that the RAF2 protein is probably involved in RuBisCO assembly in the C3 plant A. thaliana.     

Natural genetic variation in photosynthesis: an untapped resource to increase crop yield potential?

Raising crop yield potential is a major goal to ensure food security for the growing global population. Photosynthesis is the primary determinant of crop productivity and any gain in photosynthetic CO₂ assimilation per unit of leaf area (A) has the potential to increase yield. Significant intraspecific variation in A is known to exist in various autotrophic organs that represent an unexploited target for crop improvement. However, the large number of factors that influence photosynthetic rates often makes it difficult to measure or estimate A under dynamic field conditions (i.e. fluctuating light intensities or temperatures). This complexity often results in photosynthetic capacity, rather than realized photosynthetic rates being used to assess natural variation in photosynthesis. Here we review the work on natural variation in A, the different factors determining A and their interaction in yield formation. A series of drawbacks and perspectives are presented for the most common analyses generally used to estimate A. The different yield components and their determination based on different photosynthetic organs are discussed with a major focus on potential exploitation of various traits for crop improvement. To conclude, an example of different possibilities to increase yield in wheat through enhancing A is illustrated.     

Effects of overexpressing photosynthetic carbon flux control enzymes in the cyanobacterium Synechocystis PCC 6803

Synechocystis PCC 6803 is a model unicellular cyanobacterium used in e.g. photosynthesis and CO₂ assimilation research. In the present study we examined the effects of overexpressing Ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO), sedoheptulose 1,7-biphosphatase (SBPase), fructose-bisphosphate aldolase (FBA) and transketolase (TK), confirmed carbon flux control enzymes of the Calvin-Bassham-Benson (CBB) cycle in higher plants, in Synechocystis PCC 6803. Overexpressing RuBisCO, SBPase and FBA resulted in increased in vivo oxygen evolution (maximal 115%), growth rate and biomass accumulation (maximal 52%) under 100 μmol photons m⁻² s⁻¹ light condition. Cells overexpressing TK showed a chlorotic phenotype but increased biomass by approximately 42% under 100 μmol photons m⁻² s⁻¹ light condition. Under 15 μmol photons m⁻² s⁻¹ light condition, cells overexpressing TK showed enhanced in vivo oxygen evolution. This study demonstrates increased growth and biomass accumulation when overexpressing selected enzymes of the CBB cycle. RuBisCO, SBPase, FBA and TK are identified as four potential targets to improve growth and subsequently also yield of valuable products from Synechocystis PCC 6803.     

Photosynthesis research under climate change

Increasing global population and climate change uncertainties have compelled increased photosynthetic efficiency and yields to ensure food security over the coming decades. Potentially, genetic manipulation and minimization of carbon or energy losses can be ideal to boost photosynthetic efficiency or crop productivity. Despite significant efforts, limited success has been achieved. There is a need for thorough improvement in key photosynthetic limiting factors, such as stomatal conductance, mesophyll conductance, biochemical capacity combined with Rubisco, the Calvin–Benson cycle, thylakoid membrane electron transport, nonphotochemical quenching, and carbon metabolism or fixation pathways. In addition, the mechanistic basis for the enhancement in photosynthetic adaptation to environmental variables such as light intensity, temperature and elevated CO₂ requires further investigation. This review sheds light on strategies to improve plant photosynthesis by targeting these intrinsic photosynthetic limitations and external environmental factors.

     

Physiological and ecological controls on carbon sequestering in terrestrial ecosystems

Carbon cycling processes in ecosystems are generally believed to be well understood. Carbon, hydrogen, oxygen and other essential elements are chemically converted from inorganic to organic compounds primarily in the process of photosynthesis. Secondary metabolic processes cycle carbon in and among organisms and carbon is ultimately released back to the environment as CO₂ by respiratory processes. Unfortunately, our understanding of this cycle was determined under the assumption that the primary inorganic form of C (CO₂ in the atmosphere) was relatively constant. With the emerging concensus that atmospheric carbon concentration is increasing, we must now reassess our understanding of the carbon cycle. How will plants, animals and decomposers respond to a doubling of carbon supply? Will biological productivity be accelerated? If plant productivity increases will a predictable percentage of the increase be accumulated as increased standing crop? Or, is it possible that doubling the availability of CO₂ will increase metabolic activity at all trophic levels resulting in no net increase in system standing crop? The purpose of this paper is to review evidence for physiological and growth responses of plants to carbon dioxide enhancement. Essentially no research has been completed on the ecological aspects of these questions. From this review, I conclude that accurate predictions of future ecosystem responses to increasing atmospheric carbon dioxide concentration are not possible without additional understanding of physiological and ecological mechanisms.     

Potential metabolic mechanisms for inhibited chloroplast nitrogen assimilation under high CO2

Improving photosynthesis is considered a major and feasible option to dramatically increase crop yield potential. Increased atmospheric CO₂ concentration often stimulates both photosynthesis and crop yield, but decreases protein content in the main C₃ cereal crops. This decreased protein content in crops constrains the benefits of elevated CO₂ on crop yield and affects their nutritional value for humans. To support studies of photosynthetic nitrogen assimilation and its complex interaction with photosynthetic carbon metabolism for crop improvement, we developed a dynamic systems model of plant primary metabolism, which includes the Calvin–Benson cycle, the photorespiration pathway, starch synthesis, glycolysis–gluconeogenesis, the tricarboxylic acid cycle, and chloroplastic nitrogen assimilation. This model successfully captures responses of net photosynthetic CO₂ uptake rate (A), respiration rate, and nitrogen assimilation rate to different irradiance and CO₂ levels. We then used this model to predict inhibition of nitrogen assimilation under elevated CO₂. The potential mechanisms underlying inhibited nitrogen assimilation under elevated CO₂ were further explored with this model. Simulations suggest that enhancing the supply of α-ketoglutarate is a potential strategy to maintain high rates of nitrogen assimilation under elevated CO₂. This model can be used as a heuristic tool to support research on interactions between photosynthesis, respiration, and nitrogen assimilation. It also provides a basic framework to support the design and engineering of C₃ plant primary metabolism for enhanced photosynthetic efficiency and nitrogen assimilation in the coming high-CO₂ world.

Simulations with a dynamic systems model of C₃ primary metabolism show that the decreased supply of reducing equivalent and 2-oxoglutaric acid cause decreased nitrogen assimilation under elevated CO₂.     

Biochemical approaches to C4 photosynthesis evolution studies: the case of malic enzymes decarboxylases

C₄ photosynthesis enables the capture of atmospheric CO₂ and its concentration at the site of RuBisCO, thus counteracting the negative effects of low atmospheric levels of CO₂ and high atmospheric levels of O₂ (21 %) on photosynthesis. The evolution of this complex syndrome was a multistep process. It did not occur by simply recruiting pre-exiting components of the pathway from C₃ ancestors which were already optimized for C₄ function. Rather it involved modifications in the kinetics and regulatory properties of pre-existing isoforms of non-photosynthetic enzymes in C₃ plants. Thus, biochemical studies aimed at elucidating the functional adaptations of these enzymes are central to the development of an integrative view of the C₄ mechanism. In the present review, the most important biochemical approaches that we currently use to understand the evolution of the C₄ isoforms of malic enzyme are summarized. It is expected that this information will help in the rational design of the best decarboxylation processes to provide CO₂ for RuBisCO in engineering C₃ species to perform C₄ photosynthesis.     

The interactive effects of elevated CO2 and O3 concentration on photosynthesis in spring wheat

This study investigated the interacting effects of carbon dioxide and ozone on photosynthetic physiology in the flag leaves of spring wheat (Triticum aestivum L. cv. Wembley), at three stages of development. Plants were exposed throughout their development to reciprocal combinations of two carbon dioxide and two ozone treatments: [CO₂] at 350 or 700 mol mol^–1, [O₃] at < 5 or 60 nmol mol^–1. Gas exchange analysis, coupled spectrophotometric assay for RuBisCO activity, and SDS-PAGE, were used to examine the relative importance of pollutant effects on i) stomatal conductance, ii) quantum yield, and iii) RuBisCO activity, activation, and concentration. Independently, both elevated [CO₂] and elevated [O₃] caused a loss of RuBisCO protein and V_cmax. In combination, elevated [CO₂] partially protected against the deleterious effects of ozone. It did this partly by reducing stomatal conductance, and thereby reducing the effective ozone dose. Elevated [O₃] caused stomatal closure largely via its effect on photoassimilation.     

Enhancing photosynthetic CO2 use efficiency in rice: approaches and challenges

It is estimated that 925 million people mainly in developing countries suffer from malnutrition due to food shortage. This situation will deteriorate as world population will reach 9 billion by 2050. It is obvious that current rate of increase in crop yields is not sufficient to solve the problem of food security worldwide, especially in Asia, where at least 50 % increase in rice yield is needed to satisfy the increasing population. Depending on advanced development of modern biotechnology, several strategies have been provided for ‘supercharging’ photosynthesis to increase rice yield. In this review, we updated four major approaches: namely improving the performance of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco), establishment of photorespiratory bypass, installing single-cell and two-celled C₄ photosynthesis. The first approach aimed at direct manipulation of Rubisco for more efficient catalytic character by directed molecular evolution. The second approach focused on reducing the loss of photorespiratory CO₂ by direct manipulation of photorespiratory pathway. The last two concentrated on introduction of C₄ pathway into rice, based on the observation that the efficiency of C₄ photosynthesis is 50 % higher than that of C₃ photosynthesis.     

Expression of foreign type I ribulose-1,5-bisphosphate carboxylase/ oxygenase (EC 4.1.1.39) stimulates photosynthesis in cyanobacterium Synechococcus PCC7942 cells

A reporter gene assay revealed that promoters derived from Synechococcus PCC7942 (S.7942) psbAI and Synechocystis PCC6803 (S.6803) psbAII were suitable for the expression of foreign ribulose-bisphosphate carboxylase (RuBisCO; EC 4.1.1.39) in S.7942 cells. Transformational vectors with a promoter and a foreign RuBisCO gene, cvrbc originated from Allochromatium vinosum, were constructed on a binary vector, pUC303, and introduced to S.7942 cells. When the cvrbc was expressed with the S.7942 psbAI promoter, the total RuBisCO activity increased 2.5- to 4-fold than that of the wild type cell. The S.6803 psbAII promoter increased the activity of the transformant 1.5–2 times of that of wild type cell. There was a significant increase in the rate of photosynthesis depending on the increase of RuBisCO activity. The maximum rate of photosynthesis of the transformant cell was 1.63 times higher than that of the wild type under the illumination of 400 μmol m⁻² s⁻¹, at 20 mM bicarbonate and at 30 °C. Although the photosynthesis of the higher plant is limited by the ability of photosystems under high irradiance and the high CO₂ concentration, that of the S.7942 cell is limited by the RuBisCO activity, even at high CO₂ concentrations and under high irradiance.     

Crop responses to CO2 enrichment

Carbon dioxide is rising in the global atmosphere, and this increase can be expected to continue into the foreseeable future. This compound is an essential input to plant life. Crop function is affected across all scales from biochemical to agro-ecosystem. An array of methods (leaf cuvettes, field chambers, free-air release systems) are available for experimental studies of CO₂ effects. Carbon dioxide enrichment of the air in which crops grow usually stimulates their growth and yield. Plant structure and physiology are markedly altered. Interactions between CO₂ and environmental factors that influence plants are known to occur. Implications for crop growth and yield are enormous. Strategies designed to assure future global food security must include a consideration of crop responses to elevated atmospheric CO₂. Future research should include these targets: search for new insights, development of new techniques, construction of better simulation models, investigation of belowground processes, study of interactions, and the elimination of major discrepancies in the scientific knowledge base.     

Improving Photosynthesis

Photosynthesis is the basis of plant growth, and improving photosynthesis can contribute toward greater food security in the coming decades as world population increases. Multiple targets have been identified that could be manipulated to increase crop photosynthesis. The most important target is Rubisco because it catalyses both carboxylation and oxygenation reactions and the majority of responses of photosynthesis to light, CO₂, and temperature are reflected in its kinetic properties. Oxygenase activity can be reduced either by concentrating CO₂ around Rubisco or by modifying the kinetic properties of Rubisco. The C₄ photosynthetic pathway is a CO₂-concentrating mechanism that generally enables C₄ plants to achieve greater efficiency in their use of light, nitrogen, and water than C₃ plants. To capitalize on these advantages, attempts have been made to engineer the C₄ pathway into C₃ rice (Oryza sativa). A simpler approach is to transfer bicarbonate transporters from cyanobacteria into chloroplasts and prevent CO₂ leakage. Recent technological breakthroughs now allow higher plant Rubisco to be engineered and assembled successfully in planta. Novel amino acid sequences can be introduced that have been impossible to reach via normal evolution, potentially enlarging the range of kinetic properties and breaking free from the constraints associated with covariation that have been observed between certain kinetic parameters. Capturing the promise of improved photosynthesis in greater yield potential will require continued efforts to improve carbon allocation within the plant as well as to maintain grain quality and resistance to disease and lodging.Photosynthesis is the process plants use to capture energy from sunlight and convert it into biochemical energy, which is subsequently used to support nearly all life on Earth. Plant growth depends on photosynthesis, but it is simplistic to think that growth rate directly reflects photosynthetic rate. Continued growth requires the acquisition of water and nutrients in addition to light and CO₂ and, in many cases, involves competition with neighboring plants. Biomass must be invested by the plant to acquire these resources, and respiration is necessary to maintain all the living cells in a plant. Photosynthetic rate is typically measured by enclosing part of a leaf in a chamber, but to understand growth, one needs to consider the daily integral of photosynthetic uptake by the whole plant or community and how it is allocated. Almost inevitably, changing photosynthesis in some way requires more resources. Consequently, in order to improve photosynthesis, one needs to consider the tradeoffs elsewhere in the system. The title, “Improving Photosynthesis,” could be interpreted in many ways. For this review, I am restricting the scope to focus on crop species growing under favorable conditions.To support the forecast growth in human population, large increases in crop yields will be required (Reynolds et al., 2011; Ziska et al., 2012). Dramatic increases in yield were achieved by the Green Revolution through the introduction of dwarfing genes into the most important C₃ cereal crops rice (Oryza sativa) and wheat (Triticum aestivum). This allowed greater use of fertilizer, particularly nitrogen, without the risk of lodging, where the canopy collapses under the weight of the grain, causing significant yield losses (Stapper and Fischer, 1990). It also meant that biomass allocation within the plant could be altered to increase grain mass at the expense of stem mass now that the plants were shorter. Retrospective comparisons of cultivars released over time, but grown concurrently under favorable conditions with weed, pest, and disease control and physical support to prevent lodging, reveal that while modern cultivars yield more grain, they have similar total aboveground biomass (Austin et al., 1980, 1989).It is interesting to revisit the review by Gifford and Evans (1981): “over the course of evolution from the wild plant to modern cultivar, carbon partitioning was improved. Thus, as remaining scope for further improvement in carbon allocation must be small, it would be better to aim at increasing photosynthetic and growth rates. Alternatively, as partitioning is where flexibility has been manipulated in the past, it is better to aim for further increases in harvest index.” Just over 30 years have passed since this was published, and yield gains made by plant breeders have continued to come largely from increasing carbon allocation into grain (Fischer and Edmeades, 2010) and selecting for increased early vigor (Richards et al., 2010). By contrast, selection based on improving photosynthesis has yet to be achieved. Plants need leaves and roots to capture light, water, and nutrients for growth and stems to form the leaf canopy and support the flowers and grain, so further increases in harvest index may lead to a decrease in yield. Therefore, in order to increase yield potential further, it is necessary to increase total biomass. If light interception through the growing season is already fully exploited, then increasing biomass requires that photosynthesis be increased. It is the realization that further significant increases in yield potential will not be possible by continuing the current strategy that has turned attention toward improving photosynthesis. Recent technological developments now provide us with the means to engineer changes to photosynthesis that would not have been possible previously.     

Microalgae and Cyanobacteria: How Exploiting These Microbial Resources Can Address the Underlying Challenges Related to Food Sources and Sustainable Agriculture: A Review

For thousands of years, crop production has almost entirely depended on conventional agriculture. However, the reality is changing. The ever-growing population, global climate change, soil degradation and biotic/abiotic stresses are a growing threat to food production and security. Thus, sustainable alternatives to increase crop production for a population projected to reach 9.8 billion by 2050 are a major priority. In addition to vertical and soilless farming, innovative products based on bioresources, including plant growth stimulants, have been a target for sustainable food production. Such solutions have led to the exploitation of microorganisms, including microalgae and cyanobacteria as potential bioresources for food and plant biostimulant products. Microalgae (eukaryotic) and cyanobacteria (prokaryotic) are photosynthetic microorganisms with the capacity to synthesize a vast array of bioactive metabolites from atmospheric CO₂ and inorganic nutrients. The present review outlines the nutritional value of microalgae and cyanobacteria as alternative food resources. The potential aspects of microalgae and cyanobacteria as stabilizers of the net change in soil organic carbon (C) levels for reduced farmland degradation are also highlighted. The applications of microalgae and cyanobacteria as remedies for improved soil structure and fertility, and as enhancers of crop productivity and abiotic stress tolerance in agricultural settings are outlined. This review also discusses the co-cultivation of crops with microalgae or cyanobacteria in hydroponic systems to favor optimum root CO₂/O₂ levels for optimized crop production.

     

How do vascular plants perform photosynthesis in extreme environments? An integrative ecophysiological and biochemical story

In this work, we review the physiological and molecular mechanisms that allow vascular plants to perform photosynthesis in extreme environments, such as deserts, polar and alpine ecosystems. Specifically, we discuss the morpho/anatomical, photochemical and metabolic adaptive processes that enable a positive carbon balance in photosynthetic tissues under extreme temperatures and/or severe water‐limiting conditions in C₃ species. Nevertheless, only a few studies have described the in situ functioning of photoprotection in plants from extreme environments, given the intrinsic difficulties of fieldwork in remote places. However, they cover a substantial geographical and functional range, which allowed us to describe some general trends. In general, photoprotection relies on the same mechanisms as those operating in the remaining plant species, ranging from enhanced morphological photoprotection to increased scavenging of oxidative products such as reactive oxygen species. Much less information is available about the main physiological and biochemical drivers of photosynthesis: stomatal conductance (g_s), mesophyll conductance (g_m) and carbon fixation, mostly driven by RuBisCO carboxylation. Extreme environments shape adaptations in structures, such as cell wall and membrane composition, the concentration and activation state of Calvin–Benson cycle enzymes, and RuBisCO evolution, optimizing kinetic traits to ensure functionality. Altogether, these species display a combination of rearrangements, from the whole‐plant level to the molecular scale, to sustain a positive carbon balance in some of the most hostile environments on Earth.     

Canopy warming caused photosynthetic acclimation and reduced seed yield in maize grown at ambient and elevated [CO2]

Rising atmospheric CO₂ concentration ([CO₂]) and attendant increases in growing season temperature are expected to be the most important global change factors impacting production agriculture. Although maize is the most highly produced crop worldwide, few studies have evaluated the interactive effects of elevated [CO₂] and temperature on its photosynthetic physiology, agronomic traits or biomass, and seed yield under open field conditions. This study investigates the effects of rising [CO₂] and warmer temperature, independently and in combination, on maize grown in the field throughout a full growing season. Free‐air CO₂ enrichment (FACE) technology was used to target atmospheric [CO₂] to 200 μmol mol^?1 above ambient [CO₂] and infrared heaters to target a plant canopy increase of 3.5 °C, with actual season mean heating of ~2.7 °C, mimicking conditions predicted by the second half of this century. Photosynthetic gas‐exchange parameters, leaf nitrogen and carbon content, leaf water potential components, and developmental measurements were collected throughout the season, and biomass and yield were measured at the end of the growing season. As predicted for a C₄ plant, elevated [CO₂] did not stimulate photosynthesis, biomass, or yield. Canopy warming caused a large shift in aboveground allocation by stimulating season‐long vegetative biomass and decreasing reproductive biomass accumulation at both CO₂ concentrations, resulting in decreased harvest index. Warming caused a reduction in photosynthesis due to down‐regulation of photosynthetic biochemical parameters and the decrease in the electron transport rate. The reduction in seed yield with warming was driven by reduced photosynthetic capacity and by a shift in aboveground carbon allocation away from reproduction. This field study portends that future warming will reduce yield in maize, and this will not be mitigated by higher atmospheric [CO₂] unless appropriate adaptation traits can be introduced into future cultivars.     

The decline of plant mineral nutrition under rising CO2: physiological and molecular aspects of a bad deal

The elevation of atmospheric CO₂ concentration has a strong impact on the physiology of C3 plants, far beyond photosynthesis and C metabolism. In particular, it reduces the concentrations of most mineral nutrients in plant tissues, posing major threats on crop quality, nutrient cycles, and carbon sinks in terrestrial agro-ecosystems. The causes of the detrimental effect of high CO₂ levels on plant mineral status are not understood. We provide an update on the main hypotheses and review the increasing evidence that, for nitrogen, this detrimental effect is associated with direct inhibition of key mechanisms of nitrogen uptake and assimilation. We also mention promising strategies for identifying genotypes that will maintain robust nutrient status in a future high-CO₂ world.