Physiological and iTRAQ based proteomics analyses reveal the mechanism of elevated CO2 concentration alleviating drought stress in cucumber (Cucumis sativus L.) seedlings.
Identifieur interne : 000132 ( Main/Exploration ); précédent : 000131; suivant : 000133Physiological and iTRAQ based proteomics analyses reveal the mechanism of elevated CO2 concentration alleviating drought stress in cucumber (Cucumis sativus L.) seedlings.
Auteurs : Qingqing Cui [République populaire de Chine] ; Yiman Li [République populaire de Chine] ; Xinrui He [République populaire de Chine] ; Shuhao Li [République populaire de Chine] ; Xin Zhong [République populaire de Chine] ; Binbin Liu [République populaire de Chine] ; Dalong Zhang [République populaire de Chine] ; Qingming Li [République populaire de Chine]Source :
- Plant physiology and biochemistry : PPB [ 1873-2690 ] ; 2019.
Descripteurs français
- KwdFr :
- MESH :
- métabolisme : Chloroplastes, Cucumis sativus, Dioxyde de carbone, Plant.
- méthodes : Protéomique.
- physiologie : Cucumis sativus, Plant.
- Sécheresses.
English descriptors
- KwdEn :
- MESH :
- chemical , metabolism : Carbon Dioxide.
- metabolism : Chloroplasts, Cucumis sativus, Seedlings.
- methods : Proteomics.
- physiology : Cucumis sativus, Seedlings.
- Droughts.
Abstract
Carbon dioxide is one of the most important anthropogenic greenhouse gases. We previously confirmed that elevated [CO2] alleviated the negative consequences of drought stress to cucumber seedlings, but the physiological mechanism remains unknown. We investigated the morphological and physiological characteristics as well as iTRAQ-based proteomics analyses in this study under different combinations [CO2] (400 and (800 ± 20) μmol·mol-1) and water conditions (no, moderate and severe drought stress simulated by polyethylene glycol 6000). The results showed: (1) elevated [CO2] significantly increased plant height, stem diameter, leaf area and relative water content (RWC) under drought stress; (2) drought stress significantly increased J and K peaks of the chlorophyll a fluorescence transient, indicating the damage of photosynthetic electron transport chain, while elevated [CO2] decreased them especially under moderate drought condition; (3) iTRAQ-based proteomics analyses indicated that elevated [CO2] increased the abundance of psbJ and the PSI reaction center subunit VI-2 in seedlings exposed to moderate drought stress; (4) the abundance of uroporphyrinogen decarboxylase 2 and tetrapyrrole-binding protein decreased in response to elevated [CO2] under severe drought condition; (5) elevated [CO2] regulated the expression of chloroplast proteins such as those related to stress and defense response, redox homeostasis, metabolic pathways. In conclusion, elevated [CO2] enhanced the efficiency of photosynthetic electron transport, limited the absorption of excess light energy, enhanced the ability of antioxidant and osmotic adjustment, and alleviated the accumulation of toxic substances under drought stress. These findings provide new clues for understanding the molecular basis of elevated [CO2] alleviated plant drought stress.
DOI: 10.1016/j.plaphy.2019.08.025
PubMed: 31493674
Affiliations:
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Electronic address: gslqm@sdau.edu.cn.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PubMed</idno><date when="2019">2019</date><idno type="RBID">pubmed:31493674</idno><idno type="pmid">31493674</idno><idno type="doi">10.1016/j.plaphy.2019.08.025</idno><idno type="wicri:Area/Main/Corpus">000109</idno><idno type="wicri:explorRef" wicri:stream="Main" wicri:step="Corpus" wicri:corpus="PubMed">000109</idno><idno type="wicri:Area/Main/Curation">000109</idno><idno type="wicri:explorRef" wicri:stream="Main" wicri:step="Curation">000109</idno><idno type="wicri:Area/Main/Exploration">000109</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en">Physiological and iTRAQ based proteomics analyses reveal the mechanism of elevated CO<sub>2</sub> concentration alleviating drought stress in cucumber (Cucumis sativus L.) seedlings.</title><author><name sortKey="Cui, Qingqing" sort="Cui, Qingqing" uniqKey="Cui Q" first="Qingqing" last="Cui">Qingqing Cui</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Bei'jing, 100081, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Bei'jing, 100081</wicri:regionArea><wicri:noRegion>100081</wicri:noRegion></affiliation></author><author><name sortKey="Li, Yiman" sort="Li, Yiman" uniqKey="Li Y" first="Yiman" last="Li">Yiman Li</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="He, Xinrui" sort="He, Xinrui" uniqKey="He X" first="Xinrui" last="He">Xinrui He</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="Li, Shuhao" sort="Li, Shuhao" uniqKey="Li S" first="Shuhao" last="Li">Shuhao Li</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="Zhong, Xin" sort="Zhong, Xin" uniqKey="Zhong X" first="Xin" last="Zhong">Xin Zhong</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="Liu, Binbin" sort="Liu, Binbin" uniqKey="Liu B" first="Binbin" last="Liu">Binbin Liu</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Biology, Tai'an, 271018, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Biology, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="Zhang, Dalong" sort="Zhang, Dalong" uniqKey="Zhang D" first="Dalong" last="Zhang">Dalong Zhang</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018, China. Electronic address: zdl0531@126.com.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author><author><name sortKey="Li, Qingming" sort="Li, Qingming" uniqKey="Li Q" first="Qingming" last="Li">Qingming Li</name><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018, China. Electronic address: gslqm@sdau.edu.cn.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018</wicri:regionArea><wicri:noRegion>271018</wicri:noRegion></affiliation></author></analytic><series><title level="j">Plant physiology and biochemistry : PPB</title><idno type="eISSN">1873-2690</idno><imprint><date when="2019" type="published">2019</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Carbon Dioxide (metabolism)</term><term>Chloroplasts (metabolism)</term><term>Cucumis sativus (metabolism)</term><term>Cucumis sativus (physiology)</term><term>Droughts (MeSH)</term><term>Proteomics (methods)</term><term>Seedlings (metabolism)</term><term>Seedlings (physiology)</term></keywords><keywords scheme="KwdFr" xml:lang="fr"><term>Chloroplastes (métabolisme)</term><term>Cucumis sativus (métabolisme)</term><term>Cucumis sativus (physiologie)</term><term>Dioxyde de carbone (métabolisme)</term><term>Plant (métabolisme)</term><term>Plant (physiologie)</term><term>Protéomique (méthodes)</term><term>Sécheresses (MeSH)</term></keywords><keywords scheme="MESH" type="chemical" qualifier="metabolism" xml:lang="en"><term>Carbon Dioxide</term></keywords><keywords scheme="MESH" qualifier="metabolism" xml:lang="en"><term>Chloroplasts</term><term>Cucumis sativus</term><term>Seedlings</term></keywords><keywords scheme="MESH" qualifier="methods" xml:lang="en"><term>Proteomics</term></keywords><keywords scheme="MESH" qualifier="métabolisme" xml:lang="fr"><term>Chloroplastes</term><term>Cucumis sativus</term><term>Dioxyde de carbone</term><term>Plant</term></keywords><keywords scheme="MESH" qualifier="méthodes" xml:lang="fr"><term>Protéomique</term></keywords><keywords scheme="MESH" qualifier="physiologie" xml:lang="fr"><term>Cucumis sativus</term><term>Plant</term></keywords><keywords scheme="MESH" qualifier="physiology" xml:lang="en"><term>Cucumis sativus</term><term>Seedlings</term></keywords><keywords scheme="MESH" xml:lang="en"><term>Droughts</term></keywords><keywords scheme="MESH" xml:lang="fr"><term>Sécheresses</term></keywords></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en">Carbon dioxide is one of the most important anthropogenic greenhouse gases. We previously confirmed that elevated [CO<sub>2</sub>] alleviated the negative consequences of drought stress to cucumber seedlings, but the physiological mechanism remains unknown. We investigated the morphological and physiological characteristics as well as iTRAQ-based proteomics analyses in this study under different combinations [CO<sub>2</sub>] (400 and (800 ± 20) μmol·mol<sup>-1</sup>) and water conditions (no, moderate and severe drought stress simulated by polyethylene glycol 6000). The results showed: (1) elevated [CO<sub>2</sub>] significantly increased plant height, stem diameter, leaf area and relative water content (RWC) under drought stress; (2) drought stress significantly increased J and K peaks of the chlorophyll a fluorescence transient, indicating the damage of photosynthetic electron transport chain, while elevated [CO<sub>2</sub>] decreased them especially under moderate drought condition; (3) iTRAQ-based proteomics analyses indicated that elevated [CO<sub>2</sub>] increased the abundance of psbJ and the PSI reaction center subunit VI-2 in seedlings exposed to moderate drought stress; (4) the abundance of uroporphyrinogen decarboxylase 2 and tetrapyrrole-binding protein decreased in response to elevated [CO<sub>2</sub>] under severe drought condition; (5) elevated [CO<sub>2</sub>] regulated the expression of chloroplast proteins such as those related to stress and defense response, redox homeostasis, metabolic pathways. In conclusion, elevated [CO<sub>2</sub>] enhanced the efficiency of photosynthetic electron transport, limited the absorption of excess light energy, enhanced the ability of antioxidant and osmotic adjustment, and alleviated the accumulation of toxic substances under drought stress. These findings provide new clues for understanding the molecular basis of elevated [CO<sub>2</sub>] alleviated plant drought stress.</div></front></TEI><pubmed><MedlineCitation Status="MEDLINE" Owner="NLM"><PMID Version="1">31493674</PMID><DateCompleted><Year>2020</Year><Month>02</Month><Day>03</Day></DateCompleted><DateRevised><Year>2020</Year><Month>09</Month><Day>30</Day></DateRevised><Article PubModel="Print-Electronic"><Journal><ISSN IssnType="Electronic">1873-2690</ISSN><JournalIssue CitedMedium="Internet"><Volume>143</Volume><PubDate><Year>2019</Year><Month>Oct</Month></PubDate></JournalIssue><Title>Plant physiology and biochemistry : PPB</Title><ISOAbbreviation>Plant Physiol Biochem</ISOAbbreviation></Journal><ArticleTitle>Physiological and iTRAQ based proteomics analyses reveal the mechanism of elevated CO<sub>2</sub> concentration alleviating drought stress in cucumber (Cucumis sativus L.) seedlings.</ArticleTitle><Pagination><MedlinePgn>142-153</MedlinePgn></Pagination><ELocationID EIdType="pii" ValidYN="Y">S0981-9428(19)30331-6</ELocationID><ELocationID EIdType="doi" ValidYN="Y">10.1016/j.plaphy.2019.08.025</ELocationID><Abstract><AbstractText>Carbon dioxide is one of the most important anthropogenic greenhouse gases. We previously confirmed that elevated [CO<sub>2</sub>] alleviated the negative consequences of drought stress to cucumber seedlings, but the physiological mechanism remains unknown. We investigated the morphological and physiological characteristics as well as iTRAQ-based proteomics analyses in this study under different combinations [CO<sub>2</sub>] (400 and (800 ± 20) μmol·mol<sup>-1</sup>) and water conditions (no, moderate and severe drought stress simulated by polyethylene glycol 6000). The results showed: (1) elevated [CO<sub>2</sub>] significantly increased plant height, stem diameter, leaf area and relative water content (RWC) under drought stress; (2) drought stress significantly increased J and K peaks of the chlorophyll a fluorescence transient, indicating the damage of photosynthetic electron transport chain, while elevated [CO<sub>2</sub>] decreased them especially under moderate drought condition; (3) iTRAQ-based proteomics analyses indicated that elevated [CO<sub>2</sub>] increased the abundance of psbJ and the PSI reaction center subunit VI-2 in seedlings exposed to moderate drought stress; (4) the abundance of uroporphyrinogen decarboxylase 2 and tetrapyrrole-binding protein decreased in response to elevated [CO<sub>2</sub>] under severe drought condition; (5) elevated [CO<sub>2</sub>] regulated the expression of chloroplast proteins such as those related to stress and defense response, redox homeostasis, metabolic pathways. In conclusion, elevated [CO<sub>2</sub>] enhanced the efficiency of photosynthetic electron transport, limited the absorption of excess light energy, enhanced the ability of antioxidant and osmotic adjustment, and alleviated the accumulation of toxic substances under drought stress. These findings provide new clues for understanding the molecular basis of elevated [CO<sub>2</sub>] alleviated plant drought stress.</AbstractText><CopyrightInformation>Copyright © 2019 Elsevier Masson SAS. All rights reserved.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Cui</LastName><ForeName>Qingqing</ForeName><Initials>Q</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Bei'jing, 100081, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Li</LastName><ForeName>Yiman</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>He</LastName><ForeName>Xinrui</ForeName><Initials>X</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Li</LastName><ForeName>Shuhao</ForeName><Initials>S</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Zhong</LastName><ForeName>Xin</ForeName><Initials>X</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Liu</LastName><ForeName>Binbin</ForeName><Initials>B</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Biology, Tai'an, 271018, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Zhang</LastName><ForeName>Dalong</ForeName><Initials>D</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018, China. Electronic address: zdl0531@126.com.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Li</LastName><ForeName>Qingming</ForeName><Initials>Q</Initials><AffiliationInfo><Affiliation>College of Horticulture Science and Engineering, Shandong Agricultural University, Tai'an, 271018, China; State Key Laboratory of Crop Biology, Tai'an, 271018, China. Electronic address: gslqm@sdau.edu.cn.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2019</Year><Month>08</Month><Day>29</Day></ArticleDate></Article><MedlineJournalInfo><Country>France</Country><MedlineTA>Plant Physiol Biochem</MedlineTA><NlmUniqueID>9882449</NlmUniqueID><ISSNLinking>0981-9428</ISSNLinking></MedlineJournalInfo><ChemicalList><Chemical><RegistryNumber>142M471B3J</RegistryNumber><NameOfSubstance UI="D002245">Carbon Dioxide</NameOfSubstance></Chemical></ChemicalList><CitationSubset>IM</CitationSubset><MeshHeadingList><MeshHeading><DescriptorName UI="D002245" MajorTopicYN="N">Carbon Dioxide</DescriptorName><QualifierName UI="Q000378" MajorTopicYN="Y">metabolism</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D002736" MajorTopicYN="N">Chloroplasts</DescriptorName><QualifierName UI="Q000378" MajorTopicYN="N">metabolism</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D018553" MajorTopicYN="N">Cucumis sativus</DescriptorName><QualifierName UI="Q000378" MajorTopicYN="Y">metabolism</QualifierName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D055864" MajorTopicYN="Y">Droughts</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D040901" MajorTopicYN="N">Proteomics</DescriptorName><QualifierName UI="Q000379" MajorTopicYN="Y">methods</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D036226" MajorTopicYN="N">Seedlings</DescriptorName><QualifierName UI="Q000378" MajorTopicYN="N">metabolism</QualifierName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading></MeshHeadingList><KeywordList Owner="NOTNLM"><Keyword MajorTopicYN="N">Chloroplast</Keyword><Keyword MajorTopicYN="N">Cucumber seedling</Keyword><Keyword MajorTopicYN="N">Drought stress</Keyword><Keyword MajorTopicYN="N">Elevated [CO(2)]</Keyword><Keyword MajorTopicYN="N">Proteomics analyses</Keyword><Keyword MajorTopicYN="N">iTRAQ</Keyword></KeywordList></MedlineCitation><PubmedData><History><PubMedPubDate PubStatus="received"><Year>2019</Year><Month>07</Month><Day>11</Day></PubMedPubDate><PubMedPubDate PubStatus="revised"><Year>2019</Year><Month>08</Month><Day>26</Day></PubMedPubDate><PubMedPubDate PubStatus="accepted"><Year>2019</Year><Month>08</Month><Day>27</Day></PubMedPubDate><PubMedPubDate PubStatus="pubmed"><Year>2019</Year><Month>9</Month><Day>8</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate><PubMedPubDate PubStatus="medline"><Year>2020</Year><Month>2</Month><Day>6</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate><PubMedPubDate PubStatus="entrez"><Year>2019</Year><Month>9</Month><Day>8</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate></History><PublicationStatus>ppublish</PublicationStatus><ArticleIdList><ArticleId IdType="pubmed">31493674</ArticleId><ArticleId IdType="pii">S0981-9428(19)30331-6</ArticleId><ArticleId IdType="doi">10.1016/j.plaphy.2019.08.025</ArticleId></ArticleIdList></PubmedData></pubmed><affiliations><list><country><li>République populaire de Chine</li></country></list><tree><country name="République populaire de Chine"><noRegion><name sortKey="Cui, Qingqing" sort="Cui, Qingqing" uniqKey="Cui Q" first="Qingqing" last="Cui">Qingqing Cui</name></noRegion><name sortKey="He, Xinrui" sort="He, Xinrui" uniqKey="He X" first="Xinrui" last="He">Xinrui He</name><name sortKey="Li, Qingming" sort="Li, Qingming" uniqKey="Li Q" first="Qingming" last="Li">Qingming Li</name><name sortKey="Li, Shuhao" sort="Li, Shuhao" uniqKey="Li S" first="Shuhao" last="Li">Shuhao Li</name><name sortKey="Li, Yiman" sort="Li, Yiman" uniqKey="Li Y" first="Yiman" last="Li">Yiman Li</name><name sortKey="Liu, Binbin" sort="Liu, Binbin" uniqKey="Liu B" first="Binbin" last="Liu">Binbin Liu</name><name sortKey="Zhang, Dalong" sort="Zhang, Dalong" uniqKey="Zhang D" first="Dalong" last="Zhang">Dalong Zhang</name><name sortKey="Zhong, Xin" sort="Zhong, Xin" uniqKey="Zhong X" first="Xin" last="Zhong">Xin Zhong</name></country></tree></affiliations></record>Pour manipuler ce document sous Unix (Dilib)
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|texte= Physiological and iTRAQ based proteomics analyses reveal the mechanism of elevated CO2 concentration alleviating drought stress in cucumber (Cucumis sativus L.) seedlings.
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