scholarly journals Challenges and Approaches to Crop Improvement Through C3-to-C4 Engineering

2021 ◽  
Vol 12 ◽  
Author(s):  
Hongchang Cui
Keyword(s):  
Photosynthetic Efficiency ◽  
C4 Photosynthesis ◽  
Crop Improvement ◽  
Bundle Sheath ◽  
C4 Plants ◽  
System Level ◽  
C3 Plants ◽  
World Population ◽  
Kranz Anatomy ◽  
The Past

With a rapidly growing world population and dwindling natural resources, we are now facing the enormous challenge of increasing crop yields while simultaneously improving the efficiency of resource utilization. Introduction of C4 photosynthesis into C3 crops is widely accepted as a key strategy to meet this challenge because C4 plants are more efficient than C3 plants in photosynthesis and resource usage, particularly in hot climates, where the potential for productivity is high. Lending support to the feasibility of this C3-to-C4 engineering, evidence indicates that C4 photosynthesis has evolved from C3 photosynthesis in multiple lineages. Nevertheless, C3-to-C4 engineering is not an easy task, as several features essential to C4 photosynthesis must be introduced into C3 plants. One such feature is the spatial separation of the two phases of photosynthesis (CO2 fixation and carbohydrate synthesis) into the mesophyll and bundle sheath cells, respectively. Another feature is the Kranz anatomy, characterized by a close association between the mesophyll and bundle sheath (BS) cells (1:1 ratio). These anatomical features, along with a C4-specific carbon fixation enzyme (PEPC), form a CO2-concentration mechanism that ensures a high photosynthetic efficiency. Much effort has been taken in the past to introduce the C4 mechanism into C3 plants, but none of these attempts has met with success, which is in my opinion due to a lack of system-level understanding and manipulation of the C3 and C4 pathways. As a prerequisite for the C3-to-C4 engineering, I propose that not only the mechanisms that control the Kranz anatomy and cell-type-specific expression in C3 and C4 plants must be elucidated, but also a good understanding of the gene regulatory network underlying C3 and C4 photosynthesis must be achieved. In this review, I first describe the past and current efforts to increase photosynthetic efficiency in C3 plants and their limitations; I then discuss a systems approach to tackling down this challenge, some practical issues, and recent technical innovations that would help us to solve these problems.

scholarly journals LEAF ANATOMY CHARACTERIZATION OF FOUR Apochloa SPECIES: A C3 GENUS RELATED TO EVOLUTION OF C4 PATHWAY IN GRASSES

2020 ◽  
Vol 26 (1) ◽  
pp. 12-18
Author(s):  
Ane Marcela das Chagas Mendonça ◽  
Pedro Lage Viana ◽  
João Paulo Rodrigues Alves Delfino Barbosa
Keyword(s):  
Leaf Anatomy ◽  
C4 Photosynthesis ◽  
Water Status ◽  
Bundle Sheath ◽  
C3 Plants ◽  
Plant Water ◽  
Kranz Anatomy ◽  
C4 Pathway ◽  
Photosynthesis Pathway

Leaf anatomy characteristics provide important evidences about the transition between C3 and C4 pathways. The C4 photosynthesis pathway allowed to reduce the C3 photorespiratory rate, concentrating CO2 around the Rubisco site and using structures and machinery already presented in C3 plants. In monocots, it is observed a high number of C4 lineages, most of them phylogenetically related to C3 groups. The genus Apochloa (C3), subtribe Arthropogoninae, is related to two C4 genera Coleataenia and Cyphonanthus. The aim of this study was to evaluate four Apochloa species in order to establish anatomical characteristics related to the evolution of C4 pathway in this group. By means of transverse sections fully expanded leaves of A. euprepes, A. lorea, A. molinioides, and A. poliophylla were collected and the characteristics of the mesophyll (M) and bundle sheath (BS) cells were determined. These species showed a rustic Kranz anatomy with enlarged and radial arranged BS cells, which have few organelles organized in a centrifugal position. Although the modifications of BS cells are probably related to the maintenance of plant water status, we also discuss the evolution for the establishment of C4 photosynthesis in the related C4 genera.

The Kranz syndrome in the Eragrostideae (Chloridoideae, Poaceae) as indicated by carbon isotopic ratios*

Bothalia ◽  
1984 ◽  
Vol 15 (3/4) ◽  
pp. 587-590
Author(s):  
Hector O. Panarello ◽  
Evangelina Sanchez
Keyword(s):  
Mass Spectrometry ◽  
C4 Photosynthesis ◽  
Leaf Tissue ◽  
C4 Plants ◽  
C3 Plants ◽  
Isotopic Ratios ◽  
Carbon Isotope Ratios ◽  
Carbon Isotopic ◽  
Δ 13C ◽  
Kranz Leaf Anatomy

13C/12C ratios are generally regarded as being very reliable indicators of C3 or C4 photosynthesis. These relative carbon isotope ratios are expressed as a negative δ 3C and fall into two distinct groups: Kranz (or C4) plants with δ between -9°/00 no and -18°/00 and non-Kranz (C3) plants with δ between -22°/00 and -280/00 no. In this paper, 29 taxa, representing 12 genera, of the tribe Eragrostideae were examined by mass spectrometry for their δ 13C in dried leaf tissue. All these taxa proved to be C4, plants with δ13C values ranging between -13,6°/oo and -10.9°/oo. These findings confirmed published leaf anatomical observations which showed that all the studied taxa had characteristic Kranz leaf anatomy.

scholarly journals Improving Light Use Efficiency in C4 Plants by Increasing Electron Transport Rate

Proceedings ◽  
2020 ◽  
Vol 36 (1) ◽  
pp. 203
Author(s):  
Maria Ermakova ◽  
Robert T. Furbank ◽  
Susanne von Caemmerer
Keyword(s):  
Electron Transport ◽  
Conversion Efficiency ◽  
C4 Photosynthesis ◽  
Foxtail Millet ◽  
Bundle Sheath ◽  
C4 Plants ◽  
Crop Species ◽  
Bundle Sheath Cells ◽  
Light Conversion Efficiency ◽  
Sheath Cells

C4 plants play a key role in world agriculture and strategies to manipulate and enhance C4 photosynthesis have the potential for major agricultural impacts. The C4 photosynthetic pathway is a biochemical CO2 concentrating mechanism that requires the coordinated functioning of mesophyll and bundle sheath cells of leaves. Chloroplast electron transport in C4 plants is shared between the two cell types; it provides resources for CO2 fixation therefore underpinning the efficiency of photosynthesis. Using the model monocot C4 species Setaria viridis (green foxtail millet) we demonstrated that the Cytochrome (Cyt) b6f complex regulates the electron transport capacity and thus the rate of CO2 assimilation at high light and saturating CO2. Overexpression of the Cyt b6f in both mesophyll and bundle sheath cells results in a higher electron throughput and allows better light conversion efficiency in both photosystems. Importantly, increased Cyt b6f abundance in leaves provides higher rates of C4 photosynthesis without marked changes in Rubisco or chlorophyll content. Our results demonstrate that increasing the rate of electron transport is a viable strategy for improving the light conversion efficiency in C4 crop species like maize and sorghum.

scholarly journals C4 photosynthesis with Kranz anatomy evolved in the Oryza coarctata Roxb

2020 ◽  
Author(s):  
Soni Chowrasia ◽  
Tapan Kumar Mondal
Keyword(s):  
C4 Photosynthesis ◽  
Expression Patterns ◽  
Bundle Sheath ◽  
Ultrastructural Observation ◽  
Mesophyll Cells ◽  
Potential Donor ◽  
Kranz Anatomy ◽  
Vein Density ◽  
Bundle Sheath Cells ◽  
Sheath Cells

AbstractThe C4 cycle is a complex biochemical pathway that has been evolved in plants to deal with the adverse environmental conditions. Mostly C4 plants grow in arid, water-logged area or poor nutrient habitats. Wild species, Oryza coarctata (genome type KKLL; chromosome number (4x) =48, genome size 665 Mb) belongs to the genus of Oryza which thrives well under high saline as well as submerged conditions. Here, we report for the first time that O. coarctata is a C4 plant by observing the increased biomass growth, morphological features such as vein density, anatomical features including ultrastuctural characteristics as well as expression patterns of C4 related genes. Leaves of O. coarctata have higher vein density and possess Kranz anatomy. The ultrastructural observation showed chloroplast dimorphism i.e. presence of agranal chloroplasts in bundle sheath cells whereas, mesophyll cells contain granal chloroplasts. The cell walls of bundle sheath cells contain tangential suberin lamella. The transcript level of C4 specific genes such as phosphoenolpyruvate carboxylase, pyruvate orthophosphate dikinase, NADP-dependent malic enzyme and malate dehydrogenase was higher in leaves of O. coarctata compare to high yielding rice cultivar (IR-29). These anatomical, ultra structural as well as molecular changes in O. coarctata for C4 photosynthesis adaptation might be might be due to its survival in wide diverse condition from aquatic to saline submerged condition. Being in the genus of Oryza, this plant could be potential donor for production of C4 rice in future through conventional breeding, as successful cross with rice has already been reported.

scholarly journals Photosynthesis solutions to enhance productivity

2017 ◽  
Vol 372 (1730) ◽  
pp. 20160374 ◽  
Author(s):  
Christine H. Foyer ◽  
Alexander V. Ruban ◽  
Peter J. Nixon
Keyword(s):  
Grain Yield ◽  
Photosynthetic Efficiency ◽  
Crop Improvement ◽  
Step Change ◽  
Crop Plants ◽  
Light Energy ◽  
Yield Potential ◽  
Carbon Gain ◽  
The Past ◽  
Energy Assimilation

The concept that photosynthesis is a highly inefficient process in terms of conversion of light energy into biomass is embedded in the literature. It is only in the past decade that the processes limiting photosynthetic efficiency have been understood to an extent that allows a step change in our ability to manipulate light energy assimilation into carbon gain. We can therefore envisage that future increases in the grain yield potential of our major crops may depend largely on increasing the efficiency of photosynthesis. The papers in this issue provide new insights into the nature of current limitations on photosynthesis and identify new targets that can be used for crop improvement, together with information on the impacts of a changing environment on the productivity of photosynthesis on land and in our oceans. This article is part of the themed issue ‘Enhancing photosynthesis in crop plants: targets for improvement’.

scholarly journals Changes in the plasmodesma structure and permeability at the bundle sheath and mesophyll interface during the maize C4 leaf development

2020 ◽  
Author(s):  
Peng Gao ◽  
Baijuan Du ◽  
Pinghua Li ◽  
Byung-Ho Kang
Keyword(s):  
Cell Wall ◽  
Organic Acids ◽  
Molecular Diffusion ◽  
C4 Photosynthesis ◽  
Solute Concentration ◽  
Bundle Sheath ◽  
Solute Diffusion ◽  
Cell Wall Modification ◽  
Maize Leaf ◽  
Kranz Anatomy

AbstractPlasmodesmata are intercellular channels that facilitate molecular diffusion between neighboring plant cells. The development and functions of plasmodesmata are controlled by multiple intra- and intercellular signaling pathways. Plasmodesmata are critical for dual-cell C4 photosynthesis in maize because plasmodesmata at the mesophyll and bundle sheath interface mediate exchange of CO2-carrying organic acids. We examined developmental profiles of plasmodesmata and chloroplasts in the maize leaf from young cells in the base to mature cell in the tip using microscopy approaches. Young mesophyll and bundle sheath cells in the leaf base had proplastids, and their plasmodesmata were simple, devoid of cytoplasmic sleeves. In maturing cells where Kranz anatomy and dimorphic chloroplasts were evident, we observed extensive remodeling of plasmodesmata that included acquisition of an electron-dense ring on the mesophyll side and cytoplasmic sleeves on the bundle sheath side. Interestingly, the changes in plasmodesmata involved a drop in symplastic dye mobility and suberin accumulation in the cell wall, implying a more stringent mesophyll-bundle sheath transport. We compared kinetics of the plasmodesmata and the cell wall modification in wildtype leaves with leaves from ppdk and dct2 mutants with defective C4 pathways. Plasmodesmata development, symplastic transport inhibition, and cell wall suberization were accelerated in the mutant lines, probably due to the aberrant C4 cycle. Transcriptomic analyses of the mutants confirmed the expedited changes in the cell wall. Our results suggest that a regulatory machinery at the mesophyll-bundle sheath boundary suppresses erroneous flux of C4 metabolites in the maize leaf.Significance StatementPlasmodesmata in the maize Kranz anatomy mediate the exchange of organic acids between mesophyll and bundle sheath. Since solute diffusion through plasmodesmata is governed by solute concentration gradients, a balanced distribution of C4 metabolites is critical for concentration of CO2 in the bundle sheath. Plasmodesmata bridging the mesophyll and bundle sheath cytoplasm have a cylindrical cavity, which can facilitate molecular movements, and a valve-like attachment. Construction of the sophisticated plasmodesmata was linked to C4 photosynthesis, and plasmodesmata assembly finished more rapidly in maize mutants with defective C4 pathways than in wild-type plants. These results suggest that the specialized plasmodesmata contribute to controlled transport of C4 metabolites.

Are crassulacean acid metabolism and C4 photosynthesis incompatible?

2002 ◽  
Vol 29 (6) ◽  
pp. 775 ◽  
Author(s):  
Rowan F. Sage
Keyword(s):  
Crassulacean Acid Metabolism ◽  
Acid Metabolism ◽  
C4 Photosynthesis ◽  
Bundle Sheath ◽  
C4 Plants ◽  
Cam Plants ◽  
Bundle Sheath Cells ◽  
C4 Pathway ◽  
Cam Species ◽  
Cam Photosynthesis

This paper originates from a presentation at the IIIrd International Congress on Crassulacean Acid Metabolism, Cape Tribulation, Queensland, Australia, August 2001. Despite sharing a similar metabolism, crassulacean acid metabolism (CAM) and C4 photosynthesis are not known to occur in identical species, with the exception of Portulaca spp. In Portulaca, C4 and weak CAM photosynthesis occur in distinct regions of the leaf, rather than in the same cells. This is in marked contrast to the situation in most CAM species where C3 and CAM photosynthesis are active in the same cell over the course of a day and growing season. The lack of CAM and C4 photosynthesis in identical cells of a plant indicates these photosynthetic pathways are incompatible. Incompatibilities between CAM and C4 photosynthesis could have a number of biochemical, anatomical and evolutionary explanations. Biochemical incompatibilities could result from the requirement for spatial separation of C3 and C4 phases in C4 plants versus temporal separation in CAM plants. In C4 plants, regulatory systems coordinate mesophyll and bundle sheath metabolism, with light intensity being the major environmental signal. In CAM plants, a circadian oscillator coordinates day and night phases of CAM. The requirement for rapid intercellular transport in C4 plants may be incompatible with the intracellular transport and storage needs of CAM. For example, the large vacuole required for malate storage in CAM could impede metabolite diffusion between mesophyll and bundle sheath cells in C4 plants. Anatomical barriers could also exist because both CAM and the C4 pathway require distinct leaf anatomies for efficient function. Efficient function of the C4 pathway generally requires an outer layer of cells specialized for phosphoenolpyruvate (PEP) carboxylation and regeneration and an inner layer for CO2 accumulation and refixation, while CAM species require enlarged vacuoles and tight cell packing. In evolutionary terms, barriers preventing CAM and C4 photosynthesis in the same species may be the initial steps in the respective evolutionary pathways from C3 ancestors. The first steps in C4 photosynthesis are related to scavenging photorespiratory CO2 via localization of glycine decarboxylase in the bundle sheath cells. The initial steps in CAM evolution are associated with the scavenging of respiratory CO2 at night by PEP carboxylation. In each, simplified versions of the specialized anatomy may need to be present for the evolutionary sequence to begin. For C4 evolution, enhanced bundle sheath size may be required in C3 ancestors; for CAM evolution, succulence may be required. Thus, before CAM or C4 photosynthesis began to evolve, the outcome of the evolutionary experiment may have been predetermined.

scholarly journals Accelerated remodeling of the mesophyll-bundle sheath interface in the maize C4 cycle mutant leaves

2021 ◽  
Author(s):  
Peng Gao ◽  
Pengfei Wang ◽  
Baijuan Du ◽  
Pinghua Li ◽  
Byung-Ho Kang
Keyword(s):  
Cell Wall ◽  
Organic Acids ◽  
C4 Photosynthesis ◽  
Bundle Sheath ◽  
Wild Type ◽  
Symplastic Transport ◽  
Maize Leaf ◽  
Kranz Anatomy ◽  
Chloroplast Differentiation ◽  
Kinetics Of

Abstract C4 photosynthesis in the maize leaf involves the exchange of organic acids between mesophyll (M) and the bundle sheath (BS) cells. The transport is mediated by plasmodesmata embedded in the suberized cell wall. We examined the maize Kranz anatomy with a focus on the plasmodesma and cell wall suberization with microscopy methods. In the young leaf zone where M and BS cells had indistinguishable proplastids, plasmodesmata were simple and no suberin was detected. In leaf zones where dimorphic chloroplasts were evident, the plasmodesma acquired sphincter and cytoplasmic sleeves, and suberin was discerned. These modifications were accompanied by a drop in symplastic dye mobility at the M-BS boundary. We compared the kinetics of chloroplast differentiation and the modifications in M-BS connectivity in ppdk and dct2 mutants where C4 cycle is affected. The rate of chloroplast diversification did not alter, but plasmodesma remodeling, symplastic transport inhibition, and cell wall suberization were observed from younger leaf zone in the mutants than in wild type. Our results indicate that inactivation of the C4 genes accelerated the changes in the M-BS interface and the reduced permeability suggests that symplastic transport between M and BS could be gated probably for suppressing erroneous flux of C4 metabolites.

Functional compartmentation of C4 photosynthesis in the triple layered chlorenchyma of Aristida (Poaceae)

2005 ◽  
Vol 32 (1) ◽  
pp. 67 ◽  
Author(s):  
Elena V. Voznesenskaya ◽  
Simon D. X. Chuong ◽  
Nuria K. Koteyeva ◽  
Gerald E. Edwards ◽  
Vincent R. Franceschi
Keyword(s):  
Vascular Tissue ◽  
C4 Photosynthesis ◽  
Cell Types ◽  
Bundle Sheath ◽  
Mesophyll Cells ◽  
Western Blots ◽  
Kranz Anatomy ◽  
Bundle Sheath Cells ◽  
Low Levels ◽  
And Function

The genus Aristida (Poaceae), is composed of species that have Kranz anatomy and C4 photosynthesis. Kranz anatomy typically consists of two photosynthetic cell types: a layer of mesophyll cells where atmospheric CO2 is fixed into C4 acids, and an internal, chlorenchymatous vascular bundle sheath to which C4 acids are transferred and then decarboxylated to donate CO2 to the C3 cycle. The anatomy of Aristida species is unusual as it has three distinct layers of chlorenchyma cells surrounding the vascular tissue: an inner bundle sheath, an outer bundle sheath and the mesophyll cells. In this study of Aristida purpurea Nutt. var. longiseta, the functions of the three layers of chlorenchyma cells relative to the C4 photosynthetic mechanism were determined using ultrastructural analysis, western blots, immunolocalisation of photosynthetic enzymes and starch histochemistry. The results indicate that mesophyll cells contain high levels of phosphoenolpyruvate carboxylase (PEPC) and pyruvate Pi dikinase (PPDK), and function to capture CO2 in the C4 cycle. The inner bundle sheath, which is high in Rubisco and contains NADP-malic enzyme and glycine decarboxylase, functions to transfer CO2 to the C3 cycle through decarboxylation of C4 acids and by decarboxylation of glycine in the glycolate pathway. The outer chlorenchymatous sheath is where ADPG pyrophosphorylase is mainly located, and this cell layer functions as the primary site of starch storage. The outer sheath, which has low levels of Rubisco and PEPC, may also have a role in refixation of any CO2 that leaks from the inner bundle sheath cells.

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