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ACEBO - Template-Literature-Review
ACEBO - Template-Literature-Review
Acebo
Stomata, the small pores on the surfaces of leaves and stalks, regulate the flow of gases in
and out of leaves and thus plants as a whole. They adapt to local and global changes on all
timescales from minutes to millennia. Recent data from diverse fields are establishing their central
importance to plant physiology, evolution and global ecology. Stomatal morphology, distribution and
behaviour respond to a spectrum of signals, from intracellular signalling to global climatic change.
Such concerted adaptation results from a web of control systems, reminiscent of a 'scale-free'
network, whose untangling requires integrated approaches beyond those currently used.
The plant hormone abscisic acid (ABA) plays a central role in the regulation of stomatal
movements under water-deficit conditions. The identification of ABA receptors and the ABA signaling
core consisting of PYR/PYL/RCAR ABA receptors, PP2C protein phosphatases and SnRK2 protein
kinases has led to studies that have greatly advanced our knowledge of the molecular mechanisms
mediating ABA-induced stomatal closure in the past decade. This review focuses on recent progress
in illuminating the regulatory mechanisms of ABA signal transduction, and the physiological
importance of basal ABA signaling in stomatal regulation by CO2 and, as hypothesized here, vapor-
pressure deficit. Furthermore, advances in understanding the interactions of ABA and other stomatal
signaling pathways are reviewed here. We also review recent studies investigating the use of ABA
signaling mechanisms for the manipulation of stomatal conductance and the enhancement of drought
tolerance and water-use efficiency of plants.
By controlling the opening and closure of the stomatal pores through which gas exchange
occurs, guard cells regulate two of the most important plant physiological processes: photosynthesis
and transpiration. Accordingly, guard cells have evolved exquisite sensory systems. Here we
summarize recent literature on guard cell sensing of light, drought (via the phytohormone abscisic
acid (ABA)), and CO2. New advances in our understanding of how guard cells satisfy the energetic
and osmotic requirements of stomatal opening and utilize phosphorylation to regulate the anion
channels and aquaporins involved in ABA-stimulated stomatal closure are highlighted. Omics and
modeling approaches are providing new information that will ultimately allow an integrated
understanding of guard cell physiology.
Guard cells define and regulate stomatal pores, the portals through which terrestrial plants
exchange gases with the atmosphere. Guard cells therefore regulate photosynthetic CO2 capture and
plant water status and, accordingly, have evolved a multitude of mechanisms for sensing and
responding to both physical and biotic stimuli. Here we briefly summarize some of the recent
discoveries regarding guard cell responses to light, ABA and CO2, and highlight one particular aspect
of guard cell function: anion channel regulation by phosphorylation.
The control of gaseous exchange between the leaf and external atmosphere is governed by
stomatal conductance (gs); therefore, stomata play a critical role in photosynthesis and transpiration
and overall plant productivity. Stomatal conductance is determined by both anatomical features and
behavioral characteristics. Here we review some of the osmoregulatory pathways in guard cell
metabolism, genes and signals that determine stomatal function and patterning, and the recent work
that explores coordination between gs and carbon assimilation (A) and the influence of spatial
distribution of functional stomata on underlying mesophyll anatomy. We also evaluate the current
literature on mesophyll-driven signals that may coordinate stomatal behavior with mesophyll carbon
assimilation and explore stomatal kinetics as a possible target to improve A and water use efficiency.
By understanding these processes, we can start to provide insight into manipulation of these
regulatory pathways to improve stomatal behavior and identify novel unexploited targets for altering
stomatal behavior and improving crop plant productivity.
Plants and microbes have long coevolved in a constant battle to overcome the mechanisms of
defense andattack from both sides. Plants have developed means to
prevent pathogen attack by hampering invasion ofplant tissues and actively warding off pathogen
colonization. On the other hand, pathogens have evolvedstrategies to mask their presence and/or
evade hostdefenses. Plant-microbe interaction starts with molecular recognition of each other,
leading to a cascade ofsignaling events with the final output of plant resistance or susceptibility to the
pathogen. In this molecularwar, epidermis of plants is the first barrier that pathogens need to
overtake. Natural openings on the leafsurface, such as stomata, provide an entry site to pathogens.
Plants have evolved a mechanism to closestomata upon sensing microbe-associated molecular
patterns (MAMPs). This mechanism is known as stomatal defense. A decade has passed since the
discovery
of stomatal defense, and the field has expanded considerably with significant understanding of the
basicmechanisms underlying the process. Here, we give aperspective of these findings and their
implications inthe understanding of plant-microbe interactions.
Stomata are microscopic valves on plant surfaces that originated over 400 million years (Myr)
ago and facilitated the greening of Earth's continents by permitting efficient shoot-atmosphere gas
exchange and plant hydration 1. However, the core genetic machinery regulating stomatal
development in non-vascular land plants is poorly understood 2-4 and their function has remained a
matter of debate for a century 5. Here, we show that genes encoding the two basic helix-loop-helix
proteins PpSMF1 (SPEECH, MUTE and FAMA-like) and PpSCREAM1 (SCRM1) in the moss
Physcomitrella patens are orthologous to transcriptional regulators of stomatal development in the
flowering plant Arabidopsis thaliana and essential for stomata formation in moss. Targeted P. patens
knockout mutants lacking either PpSMF1 or PpSCRM1 develop gametophytes indistinguishable from
wild-type plants but mutant sporophytes lack stomata. Protein-protein interaction assays reveal
heterodimerization between PpSMF1 and PpSCRM1, which, together with moss-angiosperm gene
complementations 6, suggests deep functional conservation of the heterodimeric SMF1 and SCRM1
unit is required to activate transcription for moss stomatal development, as in A. thaliana 7. Moreover,
stomata-less sporophytes of ΔPpSMF1 and ΔPpSCRM1 mutants exhibited delayed dehiscence,
implying stomata might have promoted dehiscence in the first complex land-plant sporophytes. ©
2016 Macmillan Publishers Limited, part of Springer Nature. All rights reserved.
Stomata exert control on fluxes of CO2 and water (H2O) in the majority of vascular plants and
thus are pivotal for planetary fluxes of carbon and H2O. However, in mosses, the significance and
possible function of the sporophytic stomata are not well understood, hindering understanding of the
ancestral function and evolution of these key structures of land plants.Infrared gas analysis and
13CO2 labelling, with supporting data from gravimetry and optical and scanning electron microscopy,
were used to measure CO2 assimilation and water exchange on young, green, ± fully expanded
capsules of 11 moss species with a range of stomatal numbers, distributions, and aperture
sizes.Moss sporophytes are effectively homoiohydric. In line with their open fixed apertures, moss
stomata, contrary to those in tracheophytes, do not respond to light and CO2 concentration. Whereas
the sporophyte cuticle is highly impermeable to gases, stomata are the predominant sites of 13CO2
entry and H2O loss in moss sporophytes, and CO2 assimilation is closely linked to total stomatal
surface areas.Higher photosynthetic autonomy of moss sporophytes, consequent on the presence of
numerous stomata, may have been the key to our understanding of evolution of large, gametophyte-
independent sporophytes at the onset of plant terrestrialization.
It has been demonstrated that the leaves of a range of forest tree species have responded to
the rising concentration of atmospheric CO2 over the last 200 years by a decrease in both stomatal
density and stomatal index. This response has also been demonstrated experimentally by growing
plants under elevated CO2 concentrations. Investigation of Quaternary fossil leaves has shown a
corresponding stomatal response to changing CO2 concentrations through a glacial-interglacial cycle,
as revealed by ice core data. Tertiary leaves show a similar pattern of stomatal density change, using
palynological evidence of palaeo-temperature as a proxy measure of CO2 concentration. The present
work extends this approach into the Palaeozoic fossil plant record. The stomatal density and index of
Early Devonian, Carboniferous and Early Permian plants has been investigated, to test for any
relationship that they may show with the changes in atmospheric CO2 concentration, derived from
physical evidence, over that period. Observed changes in the stomatal data give support to the
suggestion from physical evidence, that atmospheric CO2 concentrations fell from an Early Devonian
high of 10-12 times its present value, to one comparable to that of the present day by the end of the
Carboniferous. These results suggest that stomatal density of fossil leaves has potential value for
assessing changes in atmospheric CO2 concentration through geological time.
Plants have evolved developmental plasticity which allows the up- or down-regulation of
photosynthetic and water loss capacities as new leaves emerge. This developmental plasticity
enables plants to maximise fitness and to survive under differing environments. Stomata play a
pivotal role in this adaptive process. These microscopic pores in the epidermis of leaves control gas
exchange between the plant and its surrounding environment. Stomatal development involves
regulated cell fate decisions that ensure optimal stomatal density and spacing, enabling efficient gas
exchange. The cellular patterning process is regulated by a complex signalling pathway involving
extracellular ligand–receptor interactions, which, in turn, modulate the activity of three master
transcription factors essential for the formation of stomata. Here, we review the current understanding
of the biochemical interactions between the epidermal patterning factor ligands and the ERECTA
family of leucine-rich repeat receptor kinases. We discuss how this leads to activation of a kinase
cascade, regulation of the bHLH transcription factor SPEECHLESS and its relatives, and ultimately
alters stomatal production