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Frontiers in Biology

Front Biol    2012, Vol. 7 Issue (2) : 96-112     https://doi.org/10.1007/s11515-012-1193-3
REVIEW
The stomata frontline of plant interaction with the environment-perspectives from hormone regulation
Mengmeng ZHU1, Shaojun DAI1,2, Sixue CHEN1()
1. Department of Biology, Genetics Institute, Interdisciplinary Center for Biotechnology Research (ICBR), University of Florida, Gainesville, FL 32610, USA; 2. Alkali Soil Natural Environmental Science Center, Northeast Forestry University, Key Laboratory of Saline-alkali Vegetation Ecology Restoration in Oil Field, Ministry of Education, Harbin 150040, China
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Abstract

Plants have evolved elaborate mechanisms to perceive and integrate signals from various environmental conditions. On leaf surface, stomata formed by pairs of guard cells mediate gas exchange, water transpiration as well as function in response to abiotic and biotic stresses. Stomatal closure could be induced by drought, salt, pathogen and other adverse conditions. This constitutes an instant defense response to prevent further damage to plants. Abscisic acid (ABA) is a major plant hormone involved in stress responses. Stress-activated ABA synthesis causes stomatal closure and prevents opening to reduce water loss and cell dehydration. Key regulatory receptor complex and other important components in the ABA signaling pathway have been identified. However, our knowledge of ABA signal transduction in guard cells is far from complete. Jasmonates are a group of phytohormones generally known to be important for plant defense against insects and necrotrophic pathogens. The increased levels of methyl jasmonate (MeJA) induced by herbivory and pathogen invasion show a similar effect on stomatal movement associated with ROS production as ABA. Investigation of guard cell signaling networks involving the two important phytohormones is significant and exciting. Information about protein and metabolite components and how they interact in guard cells is lacking. Here we review recent advances on hormone signaling networks in guard cells and how the networks integrate environmental signals to plant physiological output.

Keywords stomata      guard cells      hormone      signaling      molecular networks     
Corresponding Author(s): CHEN Sixue,Email:schen@ufl.edu   
Issue Date: 01 April 2012
 Cite this article:   
Shaojun DAI,Sixue CHEN,Mengmeng ZHU. The stomata frontline of plant interaction with the environment-perspectives from hormone regulation[J]. Front Biol, 2012, 7(2): 96-112.
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http://journal.hep.com.cn/fib/EN/10.1007/s11515-012-1193-3
http://journal.hep.com.cn/fib/EN/Y2012/V7/I2/96
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Fig.1  Overview of the ABA signaling networks in guard cells. [Ca], cytosolic free Ca concentration; ABA, abscisic acid; ABC, ATP binding cassette; ABI1, ABA insensitive 1; ABI2, ABA insensitive 2; ABI5, ABA insensitive 5; AREB 2, ABA responsive element binding protein 2; Asc, ascorbic acid; ATGPX3, glutathione peroxidase 3; AtRboh, respiratory burst oxidase protein; CDPK, calcium-dependent protein kinase; CP, carotenoid precursor; ETR1, ethylene response 1; G, glucosinolate; GCA2, growth controlled by abscisic acid 2; GCR2, G protein-coupled receptor; GPA1, α-subunit of the trimeric G protein; GRX, glutaredoxin; HAB1&2, homology to ABI1 1&2; IP, inositol trisphosphate; ITC, isothiocyanate; KAT1, potassium channel 1; M, myrosinase; GTG, G protein coupled receptor (GPCR) type protein; MAPK, mitogen-activated protein kinase; OST1, open stomata 1; PA, phosphatidic acid; PI3K, phosphatidylinositol-3-kinase; PI4K, phosphatidylinositol-4-kinase; PIP, phosphatidylinositol-4,5-bisphosphate; PLC, phospholipase C; PLD, phospholipase D; POX, peroxidase; PP2A, protein phosphatase 2A; PP2C, protein phosphatase 2C; PYL, pyrabactin resistance-like; PYR, pyrabactin resistance; RCAR, regulatory component of ABA receptor; ROS, reactive oxygen species; SLAC1, slow anion channel 1; SnRK2, sucrose non-fermenting 1-related protein kinase 2; TF, transcription factor; TRX, thioredoxin.
Fig.2  implified model of crosstalk between ABA and MeJA signaling in stomatal closure. [Ca], cytosolic calcium concentration; ABA, abscisic acid; ; , ; I channels, Ca ion channel; MeJA, methyl jasmonate; , ; pH, cytosolic pH; RCN1, root curling in n-naphthylphthalamic acid 1; ROS, reactive oxygen species; .
ProteinGene locusGeneReferences
ABA signalingMeJA signaling
PYR/PYL/RCAR (14) Bet v I domain proteinAt4g17870PYR1Ma et al., 2009
At5g46790PYL1Park et al., 2009
At2g26040PYL2Santiago et al., 2009
At2g38310PYL4Nishimura et al., 2010Lackman et al., 2011
At5g53160PYL8/RCAR3
At1g01360PYL9/RCAR1
PP2C Group A (9) Mg2+-dependent Ser/Thr protein phosphataseAt4g26080ABI1Leung et al., 1994Munemasa et al., 2007
At5g57050ABI2Saez et al., 2004Islam et al., 2009
At1g72770HAB1Saez et al., 2004
At1g17550HAB2Leonhardt et al., 2004
At3g11410AtPP2CAYoshida et al., 2005
At5g51760AHG1Nishimura et al., 2007
SnRK2 (10) Ser/Thr protein kinaseAt3g50500SnRK2.2Merlot et al., 2002
At5g66880SnRK2.3Fujii et al., 2007
At4g33950SnRK2.6/OST1Mustilli et al., 2002Suhita et al., 2004
CaM/CML (57) Calmodulin (-like)At3g51920CML9Delk et al., 2005
At5g37770CML24Magnan et al., 2008
CDPK (34) Ca2+-dependent Ser/Thr kinaseAt4g23650CPK3Choi et al., 2005
At4g09570CPK4Mori et al., 2006
At2g17290CPK6Ma and Wu, 2007
At1g35670CPK11Zhu et al., 2007
At4g04720CPK21Geiger et al., 2011
At4g07470CPK23
At3g57530CPK32
F-box protein (>568)At2g39940COI1Xie et al., 1998
Katsir et al., 2008
Jasmonate-ZIM domain protein (12)At1g19180JAZ1Sheard et al., 2010
Thines et al., 2007
CBL (10) Calcineurin-B likeAt4g17615CBL1/SCABP5Cheong et al., 2003
At5g47100CBL9Pandey et al., 2008
CIPK/SnRK3 (25) Ser/Thr protein kinaseAt5g01810CIPK15/PKS3Cheong et al., 2007
At1g30270CIPK23Pandey et al., 2008
Rboh (10) NADPH oxidaseAt5g47910AtRbohDKwak et al., 2003Suhita et al., 2004
At1g64060AtRbohFSuhita et al., 2004
PP2A (26) Protein phosphatase regulatory subunitAt1g25490RCN1Kwak et al., 2002Saito et al., 2008
Murata et al., 2001
Myrosinase (6)At5g26000TGG1Zhao et al., 2008Islam et al., 2009
At5g25980TGG2Islam et al., 2009
Transcription factors (>1500)At1g32640MYC2/JAI1/JIN1Abe et al., 2003Lorenzo et al., 2004
At2g47190MYB2
Tab.1  Protein components of guard cell ABA and MeJA signaling pathways in
Fig.3  Diagram of potential modifications of redox sensitive cysteines in guard cell proteins. The reversible cysteine modifications may play important signaling roles in stomatal movement as redox switches.
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