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Cellular Signaling

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Team leader: Sabine Carpin, PhD


Associate Professors:

Sabine Carpin PhD (HDR), Christiane Depierreux PhD, François Héricourt PhD (HDR), Frédéric Lamblin PhD (HDR)


Engineers and technician:

Margot Bruat, Françoise Chefdor PhD, Mélanie Larcher


PhD student and post-doc:

Hanae Makhokh, Sonia Malik PhD



Earth's climate has changed throughout history and a global climate change is observed due to human activities. The last IPCC report confirms that extreme events and disasters as drought will become more intensive. In our regions, more intensive and frequent droughts will lead to reduced water availability to food crop, forest and populiculture.

Poplar culture which is economically important in the « Région Centre - Val de Loire », is typically restricted to well irrigated zones and will be consequently strongly affected by more drastic drought events. In the future, one major issue for poplar culture in this context will be to understand signaling pathways which contribute to perceive the stress leading to poplar adaptation to drought.

Such signaling pathways have been identified and one of them corresponds to a multi-step phosphorelay system (MSP). This kind of signaling pathway is composed by three partners : a receptor belonging to the Histidine-aspartate Kinase familly (HK), a shuttle protein belonging to the Histidine-containing Phosphotransfer protein family (HPt), and a third partner belonging to the Response Regulator protein family (RR) which comprises transcription factors (RR-B).

In poplar, we identified two HK proteins (HK1a and b) which could play a role in drought perception (Chefdor et al., 2006 ; Héricourt et al., 2016), 10 HPt proteins (HPt1 to 10) (Héricourt et al., 2013), 9 RR-B (Bertheau et al., 2012 ; 2013 ; 2015) and 10 RR-A (Chefdor et al., 2018).

A preferential partnership was identified between some of these proteins as shown in the Figure below.

Fonctionnement MSP

Scheme of MSP potentialy involved in perception of drought in poplar

MSP osmosensing peuplier

MSP partners and their relationship



Cytokinins have been shown as negative regulators during drought (Tran et al., 2007) and it is now accepted that these phytohormones can play a role in plant acclimation/adaptation/tolerance during drought via MSP pathway (Li et al., 2016). Indeed, the key to understand abiotic stress tolerance resides in understanding the plant’s capacity to accelerate/maintain or repress growth (Dolferus 2014).

Consequently, plants constantly oscillate between normal or stress conditions thus between growth (acceleration or maintain) and stress tolerance (growth repression) via a shared MSP pathway between these two signals, cytokinin and drought.

However, understanding how both signals, drought and cytokinins, perceived by distinct receptors, can result in specific responses through an identical pool of HPt and RR-B proteins, remains to be determined.

Question  HK

Our project focus on the study of MSP signaling pathways, trying to understand how those signaling pathway could permit plants to conserve specific responses to both signals and continue to grow during drought.

Our aims :

- To study HK1a and b roles and their operating mechanism

- To confirm the specificity between HPt2, 7 and 9 with HK1 compared to cytokinin receptors (HK2, 3 and 4).

- To study the roles of RR-B partners of HPt2, 7 and 9.

- To study RR-A roles in drought signaling.



In vitro culture, hydroponic culture, protoplasts transformation, poplar agrotransformation, yeast two-hybrid, classical molecular biology techniques (RNA and DNA extraction, cloning techniques, sequencing, PCR, RT-PCR, western blot, etc …)


Peuplier in vitro

Poplar transformation



2H test RR18/HPt


Academic collaborations

- Centre de Biophysique Moléculaire (CBM), CNRS, Orléans

- Institut de Chimie Organique et Analytique (ICOA), CNRS-Université d'Orléans

- Laboratoire de Biomolécules et Biotechnologie Végétale (BBV), Université de Tours

- Groupe d'Etudes des Interactions Hôtes-Pathogènes (GEIHP), CHU-Université d'Angers

- Laboratoire de GlycoBiologie et Matrice Extracellulaire Végétale (Glyco-MEV), Université de Rouen

- Dipartimento di Bioscienze e Territorio, Università degli Studi del Molise, Italie

- Institute of Molecular and Cellular Biosciences, University of Tokyo, Japon


Non academic partnerships

- Alban Muller

- GlycoDiag


- Comité de Développement Horticole de la Région CVL (CDHR-CVL)

- Institut Français de la Vigne et du Vin (IFV)

- Vinopôle Centre Val de Loire

- FiBois (anciennement Arbocentre)




Mongès A, Yaakoub H, Bidon B, Glévarec G, Héricourt F, Carpin S, Chauderon L, Drašarová L, Spichal L, Binder BM, Papon N, Rochange S (2023) Are histidine kinases of arbuscular mycorrhizal fungi involved in the response to ethylene and cytokinins? Mol. Plant-Microbe Interact, in press

MAKHOKH H, LAFITE P, LARCHER M, LAMBLIN F, CHEFDOR F, DEPIERREUX C, TANIGAWA M, MAEDA T, CARPIN S, HERICOURT F (2023) Searching for osmosensing determinants in poplar Histidine-aspartate Kinases. International Journal of Molecular Sciences, 24: 6318

BUSONT O, DURAMBOUR G, BERNARD S, PLASSON C, JOUDIOU C, BAUDE L, CHEFDOR F, DEPIERREUX C, HERICOURT F, LARCHER M, MALIK S, BOULOGNE I, DRIOUICH A, CARPIN S, LAMBLIN F (2023) Black poplar (Populus nigra L.) root extracellular trap, structural and molecular remodeling in response to osmotic stress. Cells, 12: 858


DJEGHDIR I, CHEFDOR F, BERTHEAU L, KOUDOUNAS K, CARQUEIJEIRO I, LEMOS CRUZ P, COURDAVAULT V, DEPIERREUX C, LARCHER M, LAMBLIN F, HERICOURT F, GLEVAREC G, OUDIN A, CARPIN S Evaluation of type-B RRs dimerization in poplar: a mechanism to preserve signaling specificity? Plant Science, 313: 111068


BIDON B, KABBARA S, COURDAVAULT V, GLEVAREC G, OUDIN A, HERICOURT F, CARPIN S, SPICHAL L, BINDER B, COCK JM, PAPON N (2020). Cytokinin and ethylene cell signalling pathways from prokaryotes to eukaryotes. Cells, 9, 2526


HERICOURT F, LARCHER M, CHEFDOR F, KOUDOUNAS K, CARQUEIJEIRO I, LEMOS CRUZ P, COURDAVAULT V, TANIGAWA M, MAEDA T, DEPIERREUX C, LAMBLIN F, GLEVAREC G, CARPIN S (2019) New insight on HPts as hubs in poplar cytokinin and osmosensing multistep phosphorelays: cytokinin pathway uses specific HPts. Plants, 8: 591

DAUDU D, KISIALA A, WERNER RIBEIRO C, MELIN C, PERROT L, CLASTRE M, COURDAVAULT V, PAPON N, OUDIN A, COURTOIS M, DUGE DE BERNONVILLE T, GAUCHER M, DEGRAVE A, LANOUE A, LANOTTE P, SCHOULER C, BRISSET M-N, EMERY RJN, PICHON O, CARPIN S, GIGLIOLI-GUIVARC'H N, CRECHE J, BESSEAU S, GLEVAREC G Setting-up a fast and reliable cytokinin biosensor based on a plant histidine kinase receptor expressed in Saccharomyces cerevisiae. Journal of Biotechnology, 289: 103-11

CHEFDOR F, HERICOURT F, KOUDOUNAS K, CARQUEIJEIRO I, COURDAVAULT V, MASCAGNI F, BERTHEAU L, LARCHER M, DEPIERREUX C, LAMBLIN F, RACCHI ML, CARPIN S Highlighting type A RRs as potential regulators of the dkHK1 multistep phosphorelay pathway in Populus. Plant Science 277: 68-78.


DAUDU D, ALLION E, LIESECKE F, PAPON N, COURDAVAULT V, DUGE DE BERNONVILLE T, MELIN C, OUDIN A, CLASTRE M, LANOUE A, COURTOIS M, PICHON O, GIRON D, CARPIN S, GIGLIOLI-GUIVARC'H, CRECHE J, BESSEAU S, GLEVAREC G (2017) CHASE-Containing Histidine Kinase Receptors in Apple Tree: From a Common Receptor Structure to Divergent Cytokinin Binding Properties and Specific Functions. Frontiers in Plant Science 8: 1614.


Héricourt F., Chefdor F., Djeghdir I., Larcher M., Lafontaine F., Courdavault V., Auguin D., Coste F., Depierreux C., Tanigawa M., Maeda T., Glévarec G. and Carpin S. Functional Divergence of Poplar Histidine-Aspartate Kinase HK1 Paralogs in Response to Osmotic Stress. International Journal of Molecular Sciences 17: 2061-2079.


Bertheau L., Djeghdir I., Fourreau E., Chefdor F., Glévarec G., Oudin A., Depierreux C., Morabito D., Brignolas F., Courdavault V., Héricourt F., Auguin D., Carpin S. Insights into the involvement of B-type RRs in the osmosensing signaling pathway in Populus. Plant Physiology and Biochemistry 94: 244-252.


Bertheau L., Miranda M., Foureau E., Rojas Hoyos L.F., Chefdor F., Héricourt F., Depierreux C., Morabito D., Papon N., Clastre M., Scippa G.S., Brignolas F., Courdavault V., Carpin S. In planta validation of HK1 homodimerization and recruitment of preferential HPt downstream partners involved in poplar multistep phosphorelay systems. Plant Biosystems 147: 991-995.

Héricourt F.*, Chefdor F.*, Bertheau L., Tanigawa M., Maeda T., Guirimand G., Courdavault V., Larcher M., Depierreux C., Bénédetti H., Morabito D., Brignolas F., Carpin S. Characterization of histidine-aspartate kinase HK1 and identification of Histidine Phosphotransfer proteins as potential partners in a Populus multistep phosphorelay. Physiologia Plantarum 149: 188-199 (* co-premiers auteurs).


Bertheau L., Chefdor F., Guirimand G., Courdavault V., Depierreux C., Morabito D., Brignolas F., Héricourt F., Carpin S. Identification of five B-type response regulators 1 as members of a multistep phosphorelay system interacting with Histidine-containing Phosphotransfer partners of Populus osmosensor, BMC Plant Biology 12: 241-254.


Chefdor F., Bénédetti H., Depierreux C., Delmotte F., Morabito D., Carpin S. Osmotic stress sensing in Populus: Components identification of a phosphorelay system. FEBS Letters 580: 77-81.



  • January 2023

- Conférence du Dr Sabine Carpin

International Conference on Food and Nutritional Security

NABI-CIAB, Mohali, Inde

"Agroforestry to mitigate drought threats by an efficient stress signalling pathway: Poplar MultiStep Phosphorelay as model"


  • Octobre 2022

- Conference from PhD student Hanae Makhokh

34ème colloque Biotechnocentre

Ferme de Courcimont, Nouan le Fuzelier (45)

"Caractérisation d'osmosenseurs de peuplier : recherche de domaine fonctionnel"


  • February 2022

- Conference from Dr Sonia Malik

Conférences Cosmétosciences

Amphi Blaise, Polytech site Vinci, Université d'Orléans

"Biotechnological production of specialised plant metabolites"


  • January 2022

- Conference from Dr Sabine Carpin

Newton Bhabha Fund Researcher Link Workshop on Sustaining food production under environmental stress

NABI, Mohali, inde et Université de Birmingham et distanciel

"Multistep phosphorelay as a drought perception pathway in the poplar agroforestry species"

- Conference from Dr Sonia Malik

Chercheuse post-doctorante pour le projet CosmetoPop

"A brief overview of my research activities in biotechnological production of plant specialized metabolites: a case study of shikonin derivatives"


  • Octobre 2021

- Conference from Dr Kris Vissenberg

Chercheur du département de Biologie de l'Université d'Anvers

"Control of root hair growth and development by auxin and kinases in Arabidopsis thaliana"


  • Septembre 2021

- Conference from Dr François Héricourt

Meeting INRAE : gestion innovante des stress abiotiques

"Projet ProBioSec : recherche de biostimulant pour la tolérance des plantes à la sécheresse"


  • February 2021

- Conference from Dr François Héricourt

Webinar Végépolys Valley : innovation pour la transition agro-écologique en région CVL"

"Présentation du projet ProBioSec dans le cadre de la coalition Innov'AgroEco"


  • November 2020

- Conference from Dr François Héricourt

Webinar Végépolys Valley : valorisation des co-produits du bois pour la nutrition et la santé des plantes

"Projet ProBioSec : recherche de biostimulant pour la tolérance des plantes à la sécheresse à partir de feuilles de peuplier"


  • September 2020

- Conference from Pr Nicolas Papon

Professor in" Groupe d'Etudes des Interactions Hôtes-Pathogènes", CHU-Université d'Angers

"Evolution des systèmes de signalisation cellulaire à relais multiple de phosphorylation"


  • December 2019

- Conference from Dr Claire Rosnoblet

Lecturer in "Monoxyde d'azote et Réactions de Défense" team, UMR Agroécologie (Dijon)

"Caractérisation fonctionnelle de la protéine Cdc48 dans la réponse immunitaire induite par la cryptogéine chez le tabac"


  • May 2019

- Conference from Dr Sabine Carpin

Erasmus program, University of Firenze (Italy)

"Multistep phosphorelay system, a signaling pathway between growth and drought tolerance"


- Conference from Dr Roberto de Philippis, DAGRI, University of Firenze (Italy)

UFR Sciences et Techniques, Université d'Orléans

"Exploitation of microbial photosynthesis for environmentally sustainable biotechnological processes"


  • April 2019

- Conference from Dr Sabine Carpin

EMBO Symposium, Grand Hotel, New Delhi (India)

"Stress sensing in tree species"

- Conference from Dr Sabine Carpin

Workshop ICGEB, New Delhi (India)

"During drought, how to choose between growth and tolerance?"

- Conference from Dr François Héricourt

Workshop ICGEB, New Delhi (India)

"Perception of drought in poplar: search for osmosensor and its ligand"


  • Novembre 2018

- Seminar from Dr Françoise Chefdor

Equipe Signalisation Cellulaire, LBLGC, Université d'Orléans

"Highlighting type A RRs as potential regulators of the dkHK1 multistep phosphorelay pathway in Populus"


  • Septembre 2018

- Conference from Pr Ashwani Pareek, University Jawarharlal Nerhu, New Delhi (India)

Hôtel Dupanloup, Studium Orléans

"Towars improving abiotic stress tolerance in rice"


Students training by research

  • PhD thesis

- Hanae Makhokh (to be defended in 2023)

- Inès Djeghdir (defended on 15 decembre 2016)

- Lucie Bertheau (defended on 19 decembre 2013)

- Françoise Chefdor (defended on 19 decembre 2006)

  • Master degree

- Master Agrosciences, Environnement, Territoires, Paysage, Forêt, Orléans

        Spécialité Biologie Intégrative et Changements Globaux

- Master Science du Vivant, Orléans

        Spécialité Biotechnologie, Biologie Moléculaire et Cellulaire

- Master Biologie des Plantes et des Micro-organismes, Biotechnologies, Bioprocédés, Montpellier

        Spécialité Biologie des Plantes

- Master Biologie, Agronomie, Santé, Rennes

        Spécialité Sciences Cellulaire et Moléculaire du Vivant

  • Undergraduate degree

- Licence 3 parcours BOPE et BMC, Orléans

- BTS Biotechnologie, Lycée Jacques Monod, St Jean de Braye



- Conseil régional CVL : ARD CosmétoSciences 2020, CosmétoPop

- Conseil régional CVL : APR d’intérêt régional 2020, ProBioSec

- Conseil régional du Centre : projet d’initiative académique 2013, SiSCyLi

- Conseil régional du Centre : projet « recherche » Appel Blanc 2008, SiSeP

- Conseil Général du Loiret : projet de recherche innovant 2008