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Summary: After entering the leaf, CO2 faces an intricate pathway to the site of photosynthetic fixation embedded within the chloroplasts. The efficiency of CO2 flux is hindered by a number of structural and biochemical barriers which, together, define the ease of flow of the gas within the leaf, termed mesophyll conductance. Previous authors have identified the key elements of this pathway, raising the prospect of engineering the system to improve CO2 flux and, thus, to increase leaf photosynthetic efficiency. In this review, we provide a perspective on the potential for improving the individual elements that contribute to this complex parameter. We lay particular emphasis on generation of the cellular architecture of the leaf which sets the initial boundaries of a number of mesophyll conductance parameters, incorporating an overview of the molecular transport processes which have been proposed as major facilitators of CO2 flux across structural boundaries along the pathway. The review highlights the research areas where future effort might be invested to increase our fundamental understanding of mesophyll conductance and leaf function and, consequently, to enable translation of these findings to improve the efficiency of crop photosynthesis. Significance Statement: Increasing CO2 flux has been identified as an element of leaf structure/function which could significantly improve photosynthetic efficiency and, thus, crop yield. This review highlights the challenges and opportunities of achieving this aim, bringing into focus our ignorance of some fundamental aspects of leaf developmental physiology, and identifying areas where future research investment could yield improved knowledge, understanding, and impact. [ABSTRACT FROM AUTHOR]

Summary: The quantitative and spatial coordination of stomatal pores in the epidermis and airspaces in the underlying mesophyll tissue is vital for efficient gas exchange in the leaf. The mechanisms that determine the distribution of stomata in the epidermis have been studied extensively, but how this relates to the regulation of mesophyll airspace configuration is poorly understood. Recent studies have investigated how development is coordinated between these tissue layers. The evidence suggests that multiple mechanisms are likely to work concurrently to coordinate stomatal and mesophyll development for optimal leaf gas exchange, and that both genetic and physiological factors contribute to this regulation. Such advances in our understanding of leaf development have important implications for potential improvement of crop water use efficiency. [ABSTRACT FROM AUTHOR]

Summary 1I. Introduction 1II. The shoot apical meristem: the creation of a field of cells from which leaves can be formed 2III. Patterning: the selection of leaf initiation site 2IV. Morphogenesis: the initial stages of leaf formation 4V. Differentiation and determination: transcriptional networks controlling determinancy 6VI. The elaboration of leaf form 6VII. Early steps in leaf histogenesis 7VIII. Later steps in leaf differentiation: epidermal cell fate 8IX. Compound and simple leaves: variation on a theme? 9X. Changes in leaf form: developmental and environmental influences 9XI. A model of leaf formation 9 Acknowledgements 10 References 10The formation of a leaf is a basic aspect of plant development. This review provides an overview of our present understanding of the process from initiation to the final form of the leaf. Molecular genetic and cell biology approaches have yielded significant advances in this area, adding not only to our knowledge of leaf development but also to fundamental principles in plant biology. These principles will be highlighted, as well as areas where our understanding is still incomplete, in particular the problem of coordinating the multifaceted steps involved in the generation of the leaf structure.New Phytologist(2004)doi: 10.1111/j.1469-8137.2004.01292.x© New Phytologist(2004) [ABSTRACT FROM AUTHOR]

The article provides a quantitative analysis of the process of capitulum growth and the gradual but controlled process by which elements within the capitulum become destined to form individual flowers. The data provide a foundation for the modelling of flower formation and for linking the observed parameters of floral morphogenesis with the wealth of molecular data on organogenesis in plants. An analysis of auxin flux and distribution during leaf initiation is also presented. The situation in the capitulum may be linked with the spatial and temporal control of factors specific to flower formation.

Summary: Land plants evolved multiple adaptations to restrict transpiration. However, the underlying molecular mechanisms are not sufficiently understood.We used an ozone‐sensitivity forward genetics approach to identify Arabidopsis thaliana mutants impaired in gas exchange regulation.High water loss from detached leaves and impaired decrease of leaf conductance in response to multiple stomata‐closing stimuli were identified in a mutant of MURUS1 (MUR1), an enzyme required for GDP‐l‐fucose biosynthesis. High water loss observed in mur1 was independent from stomatal movements and instead could be linked to metabolic defects. Plants defective in import of GDP‐l‐Fuc into the Golgi apparatus phenocopied the high water loss of mur1 mutants, linking this phenotype to Golgi‐localized fucosylation events. However, impaired fucosylation of xyloglucan, N‐linked glycans, and arabinogalactan proteins did not explain the aberrant water loss of mur1 mutants.Partial reversion of mur1 water loss phenotype by borate supplementation and high water loss observed in boron uptake mutants link mur1 gas exchange phenotypes to pleiotropic consequences of l‐fucose and boron deficiency, which in turn affect mechanical and morphological properties of stomatal complexes and whole‐plant physiology. Our work emphasizes the impact of fucose metabolism and boron uptake on plant–water relations. [ABSTRACT FROM AUTHOR]

Summary: C4 plants typically operate a CO2 concentration mechanism from mesophyll (M) cells into bundle sheath (BS) cells. NADH dehydrogenase‐like (NDH) complex is enriched in the BS cells of many NADP‐malic enzyme (ME) type C4 plants and is more abundant in C4 than in C3 plants, but to what extent it is involved in the CO2 concentration mechanism remains to be experimentally investigated.We created maize and rice mutants deficient in NDH function and then used a combination of transcriptomic, proteomic, and metabolomic approaches for comparative analysis.Considerable decreases in growth, photosynthetic activities, and levels of key photosynthetic proteins were observed in maize but not rice mutants. However, transcript abundance for many cyclic electron transport (CET) and Calvin–Benson cycle components, as well as BS‐specific C4 enzymes, was increased in maize mutants. Metabolite analysis of the maize ndh mutants revealed an increased NADPH : NADP ratio, as well as malate, ribulose 1,5‐bisphosphate (RuBP), fructose 1,6‐bisphosphate (FBP), and photorespiration intermediates.We suggest that by optimizing NADPH and malate levels and adjusting NADP‐ME activity, NDH functions to balance metabolic and redox states in the BS cells of maize (in addition to ATP supply), coordinating photosynthetic transcript abundance and protein content, thus directly regulating the carbon flow in the two‐celled C4 system of maize. [ABSTRACT FROM AUTHOR]

Summary: Leaf structure plays an important role in photosynthesis. However, the causal relationship and the quantitative importance of any single structural parameter to the overall photosynthetic performance of a leaf remains open to debate. In this paper, we report on a mechanistic model, eLeaf, which successfully captures rice leaf photosynthetic performance under varying environmental conditions of light and CO2.We developed a 3D reaction‐diffusion model for leaf photosynthesis parameterised using a range of imaging data and biochemical measurements from plants grown under ambient and elevated CO2 and then interrogated the model to quantify the importance of these elements.The model successfully captured leaf‐level photosynthetic performance in rice. Photosynthetic metabolism underpinned the majority of the increased carbon assimilation rate observed under elevated CO2 levels, with a range of structural elements making positive and negative contributions. Mesophyll porosity could be varied without any major outcome on photosynthetic performance, providing a theoretical underpinning for experimental data.eLeaf allows quantitative analysis of the influence of morphological and biochemical properties on leaf photosynthesis. The analysis highlights a degree of leaf structural plasticity with respect to photosynthesis of significance in the context of attempts to improve crop photosynthesis. [ABSTRACT FROM AUTHOR]

Plasmodiophora brassicae (clubroot) infection leads to reprogramming of host development resulting in the formation of characteristic galls. In this work we explored the cellular events that underly gall formation in Arabidopsis thaliana with the help of molecular markers of cell division ( CYCB1:GUS) and meristematic activity ( ANT:GUS). Our results show that gall development involved the amplification of existing meristematic activities within the vascular cambium (VC) and phloem parenchyma (PP) cells in the region of the hypocotyl. Additionally we found that the increase in VC activity and prolonged maintenance of cambial-derived cells in a meristematic state was crucial for gall formation; disruption of the VC activity significantly decreased the gall size. Gall formation also perturbed vascular development with a significant reduction in xylem and increase in PP in infected plants. This situation was reflected in a decrease in transcripts of key factors promoting xylogenesis (VND6, VND7 and MYB46) and an increase in those promoting phloem formation and function (APL, SUC2). Finally we show, using the cell cycle inhibitor ICK1/KRP1 and a cle41 mutant with altered regulation of cambial stem cell maintenance and differentiation, that a decrease in gall formation did not prevent pathogen development. This finding demonstrates that although gall formation is a typical symptom of the disease and influences numbers of spores produced, it is not required for completion of the pathogen life cycle. Together, these results provide an insight into the relationship of the cellular events that accompany Plasmodiophora infection and their role in disease progression. [ABSTRACT FROM AUTHOR]

A classical view is that leaf shape is the result of local promotion of growth linked to cell proliferation. However, an alternative hypothesis is that leaf form is the result of local repression of growth in an otherwise growing system. Here we show that leaf form can indeed be manipulated in a directed fashion by local repression of growth. We show that targeting expression of an inhibitor of a cyclin-dependent kinase ( KRP1) to the sinus area of developing leaves of Arabidopsis leads to local growth repression and the formation of organs with extreme lobing, including generation of leaflet-like organs. Directing KRP1 expression to other regions of the leaf using an miRNA target sequence tagging approach also leads to predictable novel leaf forms, and repression of growth in the leaf margin blocks the outgrowth of lobes, leading to a smoother perimeter. In addition, we show that decreased growth around the perimeter and across the leaf abaxial surface leads to a change in 3D form, as predicted by mechanical models of leaf growth. Our analysis provides experimental evidence that local repression of growth influences leaf shape, suggesting that it could be part of the mechanism of morphogenesis in plants in the context of an otherwise growing system. [ABSTRACT FROM AUTHOR]

Summary PDR5-like proteins represent one group of the ABC superfamily of transporters. Members of this group are present in plants and, due to the function of PDR5-related proteins in fungi in the excretion of xenobiotics (including antifungal agents), it has been proposed that they might play a similar role in plants in the response to and detoxification of herbicides and fungicides. However, until now no functional data has been presented showing an altered plant response to any herbicide or fungicide as a result of manipulating the expression of a PDR5-like gene in plants. In this paper, we show that the plant SpTUR2 PDR5-like ABC transporter is localised to the plasma membrane and that expression of this protein in Arabidopsis leads to the acquisition of resistance to the diterpenoid antifungal agent sclareol. These data both define a possible endogenous substrate for this transporter and highlight the potential of manipulating plant chemical resistance via modulating the expression of specific PDR5-like transporters. [ABSTRACT FROM AUTHOR]

Summary To identify genes expressed at the earliest stages of leaf development, we have performed a differential display analysis using portions of meristems destined to form leaves. Our analysis led to the identification of five genes showing an asymmetric pattern of gene expression within the meristem associated with leaf formation. Surprisingly, three of these genes encoded enzymes involved in carbohydrate metabolism (ADPglucose pyrophosphorylase, sucrose synthase and an SNF1-like kinase). Furthermore, specific transcript patterns were responsive to specific sugar and hormonal treatments. The other two genes identified encoded a Phantastica-like myb transcription factor (associated with the acquisition of leaf dorsiventrality) and CYP85 (a cytochrome P450, which plays a pivotal role in brassinolide metabolism). These data, firstly, identify a novel set of marker genes for the analysis of the earliest stages of leaf formation. Secondly, the function of the proteins encoded by these genes and their expression patterns within the meristem indicate that carbohydrate metabolism is spatially regulated within a tissue involved in key developmental processes. Finally, our data provide the first indication of an asymmetry in gene expression related to hormone biosynthesis in the meristem. [ABSTRACT FROM AUTHOR]

AIM: Management of competition with predators is an important consideration for fisheries, particularly within marine protected areas (MPAs) where conservation is a primary objective. We aimed to test whether static no‐take zones within a large, sustainable‐use MPA prevented overlap between gentoo penguins and a krill fishery during two winters with contrasting prey fields. LOCATION: South Georgia, Southwest Atlantic Ocean. METHOD: We used satellite tracking (N = 16, June–September 2018) to describe gentoo penguin movements and distribution and quantified their overlap with the MPA's no‐take zone (NTZ) and the krill fishing grounds. DNA metabarcoding of scats (N = 220, April–September 2018) was used to quantify diet. RESULTS: When krill were at moderate densities and evenly distributed in 2001, gentoo penguins would have spent all of their time within the 12 NM NTZ, but when availability was low in 2018, they spent 46.3% of their time outside the NTZ and 9.6% within the krill fishing grounds. The extension of the NTZ to 30 km in response to this finding would have produced a 14.9% increase in protection for penguins and displaced 4% of fishery hauls. Gentoo penguin diet comprised 25.8% krill, which is lower than in the late 1980s but more than in 2009. MAIN CONCLUSIONS: Gentoo penguins extend their foraging range when krill is scarce, which increases the potential for spatial overlap with the krill fishery during periods of nutritional stress. Current regulations allow for expansion of both extent and catches by the krill fishery and, should this occur, gentoo penguins may face heightened risks from competition. A dynamic ocean management framework, that extends closed areas in response to near real‐time data on penguin movements and krill density estimates, may reduce the potential for competition in this sustainable‐use MPA while allowing a profitable krill fishery. [ABSTRACT FROM AUTHOR]

The pattern of cell division, growth and separation during leaf development determines the pattern and volume of airspace in a leaf. The resulting balance of cellular material and airspace is expected to significantly influence the primary function of the leaf, photosynthesis, and yet the manner and degree to which cell division patterns affect airspace networks and photosynthesis remains largely unexplored. In this paper we investigate the relationship of cell size and patterning, airspace and photosynthesis by promoting and repressing the expression of cell cycle genes in the leaf mesophyll. Using micro CT imaging to quantify leaf cellular architecture and fluorescence/gas exchange analysis to measure leaf function, we show that increased cell density in the mesophyll of Arabidopsis can be used to increase leaf photosynthetic capacity. Our analysis suggests that this occurs both by increasing tissue density (decreasing the relative volume of airspace) and by altering the pattern of airspace distribution within the leaf. Our results indicate that cell division patterns influence the photosynthetic performance of a leaf, and that it is possible to engineer improved photosynthesis via this approach. [ABSTRACT FROM AUTHOR]

Vegetation is sparsely distributed over Antarctica's ice-free ground, and distinct plant communities are present in each of the continent's 15 recently identified Antarctic Conservation Biogeographic Regions (ACBRs). With rapidly increasing human activity in Antarctica, terrestrial plant communities are at risk of damage or destruction by trampling, overland transport, and infrastructure construction and from the impacts of anthropogenically introduced species, as well as uncontrollable pressures such as fur seal (Arctocephalus gazella) activity and climate change. Under the Protocol on Environmental Protection to the Antarctic Treaty, the conservation of plant communities can be enacted and facilitated through the designation of Antarctic Specially Protected Areas (ASPAs). We examined the distribution within the 15 ACBRs of the 33 ASPAs whose explicit purpose includes protecting macroscopic terrestrial flora. We completed the first survey using normalized difference vegetation index (NDVI) satellite remote sensing to provide baseline data on the extent of vegetation cover in all ASPAs designated for plant protection in Antarctica. Large omissions in the protection of Antarctic botanical diversity were found. There was no protection of plant communities in 6 ACBRs, and in another 6, <0.4% of the ACBR area was included in an ASPA that protected vegetation. Protected vegetation cover within the 33 ASPAs totaled 16.1 km2 for the entire Antarctic continent; over half was within a single protected area. Over 96% of the protected vegetation was contained in 2 ACBRs, which together contributed only 7.8% of the continent's ice-free ground. We conclude that Antarctic botanical diversity is clearly inadequately protected and call for systematic designation of ASPAs protecting plant communities by the Antarctic Treaty Consultative Parties, the members of the governing body of the continent. [ABSTRACT FROM AUTHOR]

Leaf shape in Arabidopsis is modulated by patterning events in the margin that utilize a PIN-based auxin exporter/CUC2 transcription factor system to define regions of promotion and retardation of growth, leading to morphogenesis. In addition to auxin exporters, leaves also express auxin importers, notably members of the AUX1/ LAX family. In contrast to their established roles in embryogenesis, lateral root and leaf initiation, the function of these transporters in leaf development is poorly understood. We report that three of these genes ( AUX1, LAX1 and LAX2) show specific and dynamic patterns of expression during early leaf development in Arabidopsis, and that loss of expression of all three genes is required for observation of a phenotype in which morphogenesis (serration) is decreased. We used these expression patterns and mutant phenotypes to develop a margin-patterning model that incorporates an AUX1/ LAX1/ LAX2 auxin import module that influences the extent of leaf serration. Testing of this model by margin-localized expression of axr3-1 ( AXR17) provides further insight into the role of auxin in leaf morphogenesis. [ABSTRACT FROM AUTHOR]

The early evolution of plants required the acquisition of a number of key adaptations to overcome physiological difficulties associated with survival on land. One of these was a tough sporopollenin wall that enclosed reproductive propagules and provided protection from desiccation and UV- B radiation. All land plants possess such walled spores (or their derived homologue, pollen)., We took a reverse genetics approach, consisting of knock-out and complementation experiments to test the functional conservation of the sporopollenin-associated gene MALE STERILTY 2 (which is essential for pollen wall development in Arabidopsis thaliana) in the bryophyte Physcomitrella patens., Knock-outs of a putative moss homologue of the A. thaliana MS2 gene, which is highly expressed in the moss sporophyte, led to spores with highly defective walls comparable to that observed in the A. thaliana ms2 mutant, and extremely compromised germination. Conversely, the moss MS2 gene could not rescue the A. thaliana ms2 phenotype., The results presented here suggest that a core component of the biochemical and developmental pathway required for angiosperm pollen wall development was recruited early in land plant evolution but the continued increase in pollen wall complexity observed in angiosperms has been accompanied by divergence in MS2 gene function. [ABSTRACT FROM AUTHOR]

The causal relationship between cell division and growth in plants is complex. Although altered expression of cell-cycle genes frequently leads to altered organ growth, there are many examples where manipulation of the division machinery leads to a limited outcome at the level of organ form, despite changes in constituent cell size. One possibility, which has been under-explored, is that altered division patterns resulting from manipulation of cell-cycle gene expression alter the physiology of the organ, and that this has an effect on growth. We performed a series of experiments on retinoblastoma-related protein ( RBR), a well characterized regulator of the cell cycle, to investigate the outcome of altered cell division on leaf physiology. Our approach involved combination of high-resolution micro CT imaging and physiological analysis with a transient gene induction system, providing a powerful approach for the study of developmental physiology. Our investigation identifies a new role for RBR in mesophyll differentiation that affects tissue porosity and the distribution of air space within the leaf. The data demonstrate the importance of RBR in early leaf development and the extent to which physiology adapts to modified cellular architecture resulting from altered cell-cycle gene expression. [ABSTRACT FROM AUTHOR]

•Significant progress has been made in the identification of the genetic factors controlling leaf shape. However, no integrated solution for the quantification and categorization of leaf form has been developed. In particular, the analysis of local changes in margin growth, which define many of the differences in shape, remains problematical. •Here, we report on a software package (leafprocessor) which provides a semi-automatic and landmark-free method for the analysis of a range of leaf-shape parameters, combining both single metrics and principal component analysis. In particular, we explore the use of bending energy as a tool for the analysis of global and local leaf perimeter deformation. •As a test case for the implementation of theleafprocessor program, we show that this integrated analysis leads to deeper insights into the morphogenic changes underpinning a series of previously identified Arabidopsis leaf-shape mutants. Our analysis reveals that many of these mutants which, at first sight, show similar leaf morphology, can be distinguished via our shape analysis. •Theleafprocessor program provides a novel integrated tool for the analysis of leaf shape. [ABSTRACT FROM AUTHOR]

The role of the margin in leaf development has been debated over a number of years. To investigate the molecular basis of events in the margin, we performed an enhancer trap screen to identify genes specifically expressed in this tissue. Analysis of one of these lines revealed abnormal differentiation in the margin, accompanied by an abnormal leaf size and shape. Further analysis revealed that this phenotype was due to insertion of the trap into DWF4, which encodes a key enzyme in brassinolide biosynthesis. Transcripts for this gene accumulated in a specific and dynamic pattern in the epidermis of young leaf primordia. Targeted expression of DWF4 to a subset of these cells (the leaf margin) in a dwf4 mutant background led to both restoration of differentiation of a specific group of leaf cells (margin cells) and restoration of wild-type leaf shape (but not leaf size). Ablation of these cells led to abrogation of leaf development and the formation of small round leaves. These data support the hypothesis that events in the margin play an essential role in leaf morphogenesis, and implicate brassinolide in the margin as a key mediator in the control of leaf shape, separable from a general function of this growth factor in the control of organ size. [ABSTRACT FROM AUTHOR]