Franceschi Cortina The C4 photosynthesis conference in Canberra in 2016 brought together world experts in the field ranging in discipline across biochemistry, physiology, molecular genetics and ecophysiology, and also included those involved in applied efforts to engineer C4 into C3 crops. It is now over half a century since the biochemical characterization of the C4 photosynthetic pathway, and this special issue highlights the sheer breadth of current knowledge. The process of photosynthesis in desert plants has evolved mechanisms to conserve water. AT Brutnell Recently, however, high resolution MALDI imaging mass spectrometry was used to examine the lipid composition of thylakoids of mesophyll and bundle sheath cells of maize (Dueñas et al., 2016). After the seminal experiments by Hatch and Slack, unravelling the biochemistry of the pathway in a number of species followed rapidly and provided the foundation of our current knowledge on the diverse biochemistry of C4 photosynthesis (Hatch, 1987; Hatch, 1992; Furbank, 2016). It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. Pignon M Oxford University Press is a department of the University of Oxford. By modelling the processes of diffusion, capture and release of CO2 and oxygen inside a typical Bienertia mesophyll cell, Jurić et al. RD Osborn Lauterbach Freitag (2017) and Schüßler et al. Mengin It has long been recognized that C4 plant species have a higher temperature optimum for photosynthesis than C3 plants due to the operation of a CO 2-concentrating system that inhibits Rubisco oxygenase activity (Berry and Björkman, 1980; Edwards and Walker, 1983).In C3 plants, inhibition of net photosynthesis (Pn) at moderately high temperatures has usually been ascribed to … Theories developed for the interpretation of stable isotope discrimination during C4 photosynthesis (Farquhar, 1983; Gillon and Yakir, 2000; Barbour et al., 2016) allow us to probe the interconnectivity of C3 and C4 cycle activity and CO2 diffusion properties into mesophyll cells. 19: C3 plants can perform photosynthesis only when the stomata are open. (2017) report on phylogenetic searches for C3–C4 species. 1.1. S This technique was used for the first time by Arrivault et al. Plants that use C4 carbon fixation concentrate carbon dioxide spatially, using “bundle sheath cells” which are inundated with CO 2. RE RF About 1500 species are a variety of dicot species, some of which form woody tissues and grow as shrubs and small trees. Denton 2. Aubry R . As gene and transcript sequencing costs plummet with third-generation technologies, what will be the new technological driver of C4 research? Gowik RF Guenther Gowik Australian Research Council Centre of Excellence for Translational Photosynthesis, Division of Plant Sciences, Research School of Biology, The Australian National University. S C4 Pathway (Hatch and Slack Pathway) Every photosynthetic plant follows Calvin cycle, but in some plants, there is a primary stage to the Calvin Cycle known as C4 pathway. S YJ The techniques of the 14C pulse chase which were used by Hatch and Slack to unravel the mysteries of C4 photosynthesis have been replaced by mass spectrometric measurements of 13CO2 labelling kinetics, which provide a wealth of information compared to past experiments. Barbour It will provide a welcome tool for establishing C4 metabolism in C3 species. U S GE The key difference between C3 and C4 plants is that the C3 plants form a three-carbon compound as the first stable product of the dark reaction while the C4 plants form a four-carbon compound as the first stable product of the dark reaction.. Photosynthesis is a light-driven process that converts carbon dioxide and water into energy-rich sugars in plants, algae and cyanobacteria. (2017) draw our attention to limitations connected to N metabolism and vein density that may have constrained the evolutionary transition of two Moricandia species (Brassicaceae) from C3–C4 into the C4 pathway. Schulze It is with sadness that we note that Roger Slack passed away on 24 October 2016. ; This reaction is catalyzed by the enzyme ribulose bisphosphate carboxylase oxygenase (RUBISCO). This 3 minute tutorial discusses the differences between the three types of photosynthesis: C3, C4 and CAM. The description of improved mathematical models, combined gas exchange and stable isotope tools, metabolic 13CO2 labelling kinetics and more efficient transformation systems for C4 plants (such as Setaria viridis) hold great promise for improving C4 photosynthesis in a crop environment. A typical plant on the earth that uses photosynthesis is a C3 plant. Schlüter et al. Ocampo Alonso-Cantabrana Published by Oxford University Press on behalf of the Society for Experimental Biology. . Kadereit CJ M Plants in tropical desert regions commonly follow the C4 pathway. If you're behind a web filter, please make sure that the domains *.kastatic.org and *.kasandbox.org are unblocked. Burgess P Voznesenskaya S Schwander Kocacinar . An Achilles heel for the dense modern stands of our most productive C, Variations in nitrogen use efficiency reflect the biochemical subtype while variations in water use efficiency reflect the evolutionary lineage of C4 grasses at inter-glacial CO, Regulatory gateways for cell-specific gene expression in C, Photosynthetic efficiency and carbon concentration in terrestrial plants: the C, A portrait of the C4 photosynthetic family on the 50, Photorespiration and the evolution of C4 photosynthesis, Exploiting the engine of C4 photosynthesis, Molecular phylogeny and forms of photosynthesis in tribe Salsoleae (Chenopodiaceae), A synthetic pathway for the fixation of carbon dioxide in vitro, Multigene manipulation of photosynthetic carbon assimilation increases CO, Global distribution of C-3 and C-4 vegetation: carbon cycle implications, Water-use efficiency and nitrogen-use efficiency of C, Biochemical models of leaf photosynthesis, Strategies for improving C4 photosynthesis, Kranz anatomy is not essential for terrestrial C4 plant photosynthesis, An untranslated cis-element regulates the accumulation of multiple C4 enzymes in. General features of the C 4 cycle. KA Fernie I. The evolution of C4 photosynthesis entails the recruitment of enzyme activities that are not involved in photosynthetic carbon fixation in C3 plants to photosynthesis. C4 plants use this 4-carbon compound to effectively “concentrate” CO2 around rubisco, so that rubisco is less likely re react with O2. P EA Gong MJ The C4 Plants, evolved from the C3 plants, have high nitrogen and water use efficiency. The ancestors of maize and miscanthus appear to have existed in very open habitats, where water and nutrient deficiencies would have limited leaf area. T S Lercher The first stable product formed in C3 cycle is a three carbon (3C) compound, hence the name. To use Khan Academy you need to upgrade to another web browser. Saladié This special issue follows two other recent volumes of Journal of Experimental Biology focused on C4 (‘Exploiting the engine of C4 photosynthesis’ – Volume 62, Issue 9, see Sage and Zhu, 2011; and ‘C4 and CAM photosynthesis in the new millennium’– Volume 65, Issue 13, see Sage, 2014). . M SJ AB . AR In the field of C4 engineering, synthetic biology has the potential to impact hugely on both basic and basic/strategic engineering approaches (Schwander et al., 2016). New genomic and transcriptomic information shows that multi-level regulation of gene expression is required for the pathway to function, yet we know it to be one of the most dynamic examples of convergent evolution. Maurino Li Hibberd C4 plants use the C4 carbon fixation pathway to increase their photosynthetic efficiency by reducing or suppressing photorespiration, which mainly occurs under low atmospheric CO 2 concentration, high light, high temperature, drought, and salinity. (2016) resolve the C3–C4 intermediate and C4 lineages in the Salsoleae family (Chenopodiaceae). von Caemmerer D Kadereit JE Despite this complexity, C4 photosynthesis is recognized as one of the most dynamic examples of convergent evolution, arising multiple times over the last 60 million years in warm semi-arid regions, with early occurrences coinciding with low atmospheric CO2 in the late Oligocene (Sage et al., 2011; Sage, 2016). (2017), who have developed a cross-species genome scanning approach to identify genes under positive selection in C4 evolution which is independent from knowledge of the biochemical pathways involved (see also the Insight article in this issue by Christin, 2017). This led to the definition of the C 4 dicarboxylic acid pathway, later abbreviated to C 4 photosynthesis, and the plants employing this process were termed C 4 plants. Burroughs Koteyeva This requires a different regulation of gene … Lee Tomato Protein Phosphatase 2C (SlPP2C3) influences fruit ripening onset and fruit glossiness, Fruit presence induces polar auxin transport in citrus and olive stem and represses IAA release from the bud, Leaf apoplastic alkalisation promotes transcription of the ABA synthesising enzyme Vp14 and stomatal closure in, Molecular and functional analysis of a brown planthopper resistance protein with two nucleotide binding site domains, The inverse relationship between solar-induced fluorescence yield and photosynthetic capacity: benefits for field phenotyping, About the Society for Experimental Biology, What can we learn from genomes and gene regulation in C, New technologies and mathematical models elucidate the physiology and biochemistry of C, http://creativecommons.org/licenses/by/4.0/, Receive exclusive offers and updates from Oxford Academic, Copyright © 2020 Society for Experimental Biology. Plants utilizing … Biology is brought to you with support from the Amgen Foundation. © The Author 2017. Photosynthesis - Photosynthesis - Carbon fixation in C4 plants: Certain plants—including the important crops sugarcane and corn (maize), as well as other diverse species that are thought to have expanded their geographic ranges into tropical areas—have developed a special mechanism of carbon fixation that largely prevents photorespiration. AJ Powell CO2 diffusion during C4 photosynthesis also remains an important issue in those species with Kranz anatomy. S P Alexander These functional models of C4 which allow the link between leaf biochemistry and gas exchange have become essential tools (von Caemmerer and Furbank, 1999; von Caemmerer, 2000). . Schulze Schmidt Rolland A transcriptomics/genomics approach and a review of gene expression across multiple lineages of C4 plants (Aubry et al., 2014; Williams et al., 2016; Reeves et al., 2017) have led to the conclusion that regulation of gene expression at multiple levels (including transcriptional control by promoter regions, as demonstrated by Gowik et al., 2017) is required for C4 photosynthesis to function. (2017) go a step further and combine physiological, anatomical and transcriptomic approaches to elucidate the molecular transition from the C3 to the C4 state in the leaves of Salsola soda (Chenopodiaceae). ; The resulting 6-carbon compound breaks down into two molecules of 3-phosphoglyceric acid (PGA). Zhu The C4 pathway isn’t just about biochemistry, rather it is a complex combination of biochemical and morphological specialization. Melzer Gene discovery through genomics approaches reveals gene candidates and evidence for the importance of certain genes in evolution or for plant performance, but these must be experimentally validated. It is the gas-tight nature of the bundle sheath that allows the decarboxylation of C4 acids in this compartment to elevate CO2 partial pressure around Rubisco. C4 photosynthesis is a CO 2-concentrating mechanism present in about 7000 species of higher plants. . The prized goal has been the mining of C3–C4 species to identify anatomical, biochemical and molecular features that underlie C4 evolution. Furbank (2016) points to a wealth of genomic and transcriptomic information now available for C4 leaves, and leaves of closely related C3 plants, which is catalysing a new generation of research into the C4 mechanism and the genetic architecture underpinning it. GE Marshall-Colon JM . R Bräutigam Gowik Cousins Sheehy Williams U This led to the definition of the C4 dicarboxylic acid pathway, later abbreviated to C4 photosynthesis, and the plants employing this process were termed C4 plants. Westhoff (2017) show that a spatial separation as low as 10 μm between the primary and the secondary carboxylases can provide enough diffusive resistance to sustain an efficient C4 pathway, demonstrating that single-cell C4 photosynthesis is a viable option. All plant species rely on this process to produce their source of energy. Edwards Dueñas GE BMC If you're behind a web filter, please make sure that the domains *.kastatic.org and *.kasandbox.org are unblocked. . What are C3 Plants. Simkin DT In this issue, both Voznesenskaya et al. TJ EJ Kellogg In comparison to C3 crops such as rice, C4 crops have higher yields and increased water and nitrogen use efficiency (Hibberd et al., 2008; Langdale, 2011). E AP C4 Photosynthesis. RS Hibberd Weber Hibberd Reyna-Llorens RE In 50 years we have seen the expansion of the field from the examination of a rudimentary biochemical pathway in just a few species to the construction of complex evolutionary models and assembly of massive genomic and transcriptomic data sets from a large range of both crop and wild C4 species, as well as multiple efforts to engineer C4 traits into C3 crops and model species. VG It appears that posttranscriptional control may also be important (Fankhauser and Aubry, 2017) and that many of the mechanisms for regulation of C4 gene expression are indeed present in C3 plants and recruited to a C4 function (Reeves et al., 2017). G C4 plants grow ‘cheaper’ leaves which allows them to produce 50 per cent more roots than C3 species. D Voznesenskaya et al. Long M CAM Photosynthesis, Calvin Cycle, C3 Photosynthesis, C4 Photosynthesis, Photorespiration. Plants have different systems for harvesting energy depending on their environment. Weber KL . However, plants have evolved another form of photosynthesis to help reduce these losses in hot, dry environments. In his Darwin review, Sage (2016) outlines the evolution of the 61 independent C4 lineages which have resulted in more than 8000 species in grasses, sedges and eudicots and looks at the biogeography of these species. Sharwood Aubry Photorespiration: C3, C4, and CAM plants C3, C4, and CAM plants How the C4 and CAM pathways help minimize photorespiration. RT XG How the C4 and CAM pathways help minimize photorespiration. S EV H Denton et al. Hibberd Lercher ME Covshoff Furbank, in one of two comprehensive Darwin reviews in this issue, retraces these historical events in detail ( Furbank, 2016 ). T (2017) use leaf maturation in C3 and C4Flaveria species to identify transcription factors. This led to the development of distinguishing gas exchange features of C4 CO2 assimilation rates. Mettler-Altmann This inhibits its oxygenase activity allowing it to operate close to its maximal rate. P A Kurz P Burgener Reeves I J Smith-Unna From a biochemical and modelling perspective, the confounding nature of the two-compartment C4 system for ‘grind and find’ extraction of metabolites, transcripts and proteins has been a challenge. Lercher (2017) in maize to establish pool sizes and gradients of metabolites using cell type fractionation. Recent technical advances have greatly facilitated the measurements of isotope discrimination concurrently with gas exchange (Gong et al., 2017; Osborn et al., 2017). Carbon dioxide is absorbed from the atmosphere and oxygen is released. Stomata – these are pores (holes) in the leaves that are responsible for the exchange of gases between the plant leaves and the atmosphere. MD T We now have a good understanding of how C4 photosynthesis responds to environmental variables such as light, temperature and CO2 (Long, 1999). I Therefore, C4 plants dominate grassland floras and biomass production in the warmer climates of the tropical and A Upper and lower epidermis – the upper epidermis is the outer layer of the cells that controls the amount of water that is lost through transpiration. A A (2017), who use these models (following Heckmann et al., 2013) to outline how to combine genetic and evolutionary engineering to establish C4 metabolism in C3 plants. MJ N Fixing carbon is the way plants remove the carbon from atmospheric carbon dioxide and turn it into organic molecules like carbohydrates. S Biochemical and molecular studies have elucidated the various types of photosynthetic intermediacy, which range from the simpler C2 mode involving the glycine or photorespiratory shuttle with rudimentary bundle sheath to a C4-like pathway with well-developed Kranz anatomy and functional C4 pathway (Sage et al., 2012). O It is interesting that molecular genetics, genomics and transcriptomics are now commonly being used in a biochemical and evolutionary research perspective as affordable approaches to answer questions in C4 photosynthesis research, rather than operating in isolation as stand-alone fields (Box 2). PA Hilary Warren (then Johnson) was the first PhD student of Hal Hatch. González-Pérez H With appropriate rapid kill and cryopreservation, this technique may hold promise for measuring metabolites during photosynthesis in mesophyll and bundle sheath compartments more accurately. TP Physiological studies have revealed a clear lowering of the CO2 compensation point (CO2 partial pressure where there is no net CO2 exchange) for all types of C3–C4 intermediates, but advantages related to improved water and nitrogen use efficiency are only expressed in intermediate plants possessing a degree of C4 acid fixation (Vogan et al., 2011; Pinto et al., 2016). U NS Roger Slack, Hilary Warren and Hal Hatch at the opening of the conference ‘C4 Photosynthesis: past, present and future’ in April 2016. Westhoff C4 plants use C4 photosynthesis to avoid photorespiration. MJ Klein U C3, C4 and CAM are the three different processes that plants use to fix carbon during the process of photosynthesis. Christin The drawback to C4 photosynthesis is the extra energy in the form of ATP that is used to pump the 4-carbon acids to the bundle sheath cell and the pumping of the 3-carbon compound back to the mesophyll cell for conversion to PEP. JM JM In plants employing full CAM photosynthesis, the stomata in the leaves are closed during daylight hours to lessen evapotranspiration and open at night in order to take in carbon dioxide. The meeting was held at the ARC Centre of Excellence for Translational Photosynthesis in Canberra, Australia to commemorate the discovery of the C4 pathway and its significance in today’s plant biology and agricultural research (see Hibberd and Furbank, 2016). McAusland Edwards Evolution from C 3 to C 4 photosynthesis. P-A C3 and C4 Plants Photosynthesis is a complex pathway which is used by plants to fix carbon, present in the atmosphere, into sugar. (2017) documented dynamic variation in bundle sheath leakiness of a perennial C4 grass with short-term variation in atmospheric CO2 concentration. Yandeau-Nelson . The multidisciplinary approaches used and needed to unravel the secrets of C, Metabolite pools and carbon flow during C, Deep evolutionary comparison of gene expression identifies parallel recruitment of trans-factors in two independent origins of C4 photosynthesis, The proceedings of the fourth international congress on photosynthesis, Traces of strong selective pressures in the genomes of C, Freeze-quenched maize mesophyll and bundle sheath separation uncovers bias in previous tissue-specific RNA-Seq data, High-spatial resolution mass spectrometry imaging reveals the genetically programmed, developmental modification of the distribution of thylakoid membrane lipids among individual cells of the maize leaf, Post-transcriptional regulation of photosynthetic genes is a key driver of C, On the nature of carbon isotope discrimination in C, Naturally low carbonic anhydrase activity in C, Bundle-sheath leakiness and intrinsic water use efficiency of a perennial C. Evolution of C4 photosynthesis in the genus Flaveria: how many and which genes does it take to make C4? C C4 plants make a four-carbon sugar during the Calvin cycle. Weber D If you're seeing this message, it means we're having trouble loading external resources on our website. L M (2017) demonstrate that the family Portulacaceae has a C3–C4 Cryptopetala clade and a diverse C4 Pilosa clade, while Schüßler et al. Now, a focus on the molecular transition from C3–C4 intermediates, together with improved mathematical models, experimental tools and transformation systems, holds great promise for improving C4 photosynthesis in crops. Schüssler Huang DeFries Voznesenskaya C4 photosynthesis evolved multiple times independently from ancestral C3 photosynthesis in a broad range of flowering land plant families and in both monocots and dicots. (2017) remind us, though, that caution must be used in interpreting gene expression data, particularly cell- or tissue-specific data, which may include biases due to RNA preparation methods. Still JA Sage Gong et al. Pignon et al. J G BM S Search for other works by this author on: ARC Centre of Excellence for Translational Photosynthesis and Hawkesbury Institute for the Environment, Western Sydney University, Box 2. Knerova GJ (A) C 3 photosynthesis: the Calvin–Benson cycle and photorespiration work independently in both cell types. Beans, Rice, Wheat, and Potatoes are an example of plants that follow the C3 pathway. LE Many variations of C 4 photosynthesis exist, reflecting at least 45 independent origins in 19 families of higher plants. . . BJ Denton Alternatives such as producing large panels of mutants by non-targeted mutagenic approaches or by crossing genetic material to develop near-isogenic lines with and without genetic polymorphisms is outside the scope of most small research laboratories. Gowik . More recent studies, including those represented in this issue, have focused on documenting the phylogenetic diversity of C3–C4 taxa and elucidating the molecular elements underscoring the evolutionary, and in rare cases, the developmental, transitions from C3 to C4 (Gowik et al., 2011). V Among these are important crops such as maize, sorghum and sugarcane, but also weeds and invasive pla… 3. A comparison of C3, C4 and CAM plants. H These plants are called C4 plants, because the first product of carbon fixation is a 4-carbon compound (instead of a 3-carbon compound as in C3 or “normal” plants). C4 plants play a key role in world agriculture – crops such as maize and sorghum are major contributors to world food production in both developed and developing nations, and the C4 grasses sugarcane, miscanthus and switchgrass are the major plant sources of bioenergy. Smith-Unna C4 plants can do photosynthesis even in the closed condition of stomata. Most C4 species are characterized by Kranz anatomy, but there are a small number, such as Bienertia cycloptera, that perform C4 photosynthesis within individual mesophyll cells (Voznesenskaya et al., 2001; King et al., 2012). Lawson Billakurthi I JM JM Koteyeva It evolved as an adaptation to high light intensities, high temperatures, and dryness. Similarly, if CRISPR/Cas9 technology is combined with high-efficiency C4 grass transformation systems, production of allele mimics of potentially important genes occurring in nature and engineering of novel enzyme properties in C4 plants would advance rapidly. Where will C4 research go next? The agronomic use of C4 species, as well as their substantial influence on terrestrial CO2 fixation (Still et al., 2003), provides the scientific drive for understanding what has allowed the evolution of C4 photosynthesis to happen so many times. Obata Külahoglu Arrivault M AP . von Caemmerer Furbank K M Stanley There is a vast array of information and new technology now at our fingertips. GE Your comment will be reviewed and published at the journal's discretion. Three-fourths of the C4 species are grasses and sedges of warm-temperate, subtropical, and tropical zones. .Kasandbox.Org are unblocked as shrubs and small trees the Calvin–Benson cycle and photorespiration work independently in both cell types while! May therefore have been little evolutionary pressure for maintenance of photosynthetic efficiency in shade conditions there a. In bundle sheath cells in plant species rely on this article C4 made. 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The recruitment of enzyme activities that are not involved in photosynthetic carbon cycle is the first student! For full access to this pdf, sign in to an existing account, or purchase an annual subscription technology! Having trouble loading external resources on our website reaction is catalyzed by the ribulose... Concentrate carbon dioxide spatially, using “bundle sheath cells” which are inundated with CO 2 the. Cortina NS Erb TJ to produce 50 per cent more roots than species... By the enzyme ribulose bisphosphate carboxylase oxygenase ( rubisco ) light and dark of. Away on 24 October 2016 by Oxford University Press is a three (..., Research School of Biology, the light and dark reactions of the (! That are not involved in photosynthetic carbon fixation or the Hatch–Slack pathway is one of two comprehensive Darwin reviews this...

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