Welcome to the Citrus Lab

Unraveling citrus physiology to enhance tree performance and resilience in the face of climate change


Citrus molecular physiology; 
Citrus genome editing;
 Abiotic stress responses;
Guard cell signaling;
  Tree architecture

Gilor Kelly

Principal Investigator

Tamar Basha

Research Associate / Lab Manager

Dayakar Boddupally

Post-Doctoral Fellow

Shay Igon-Hagbi

PhD Student

Co-supervision, Prof. Nir Sade (TAU)

Yoav Yoktan

MSc Student

Co-supervision, Prof. Avi Sadka (ARO)

Shmulik (Shmuel) Gorlik

MSc Student

Co-supervision, Prof. Leor Eshed-Williams (HUJI)

11/25/2025

In planta genome editing approach in citrus

Targeted breeding for specific trait improvements in already successful cultivars is likely to revolutionize fruit tree breeding and will pave the way for accelerating the development of high-quality citrus cultivars.

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11/25/2025

Guard cells and plant development

Guard cells are capable of coordinating hypocotyl elongation and plant development via sugar sensors that have the opposite effect of light on hypocotyl elongation, converging at PIF4.

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11/25/2025

Guard cells specific activity for improved water use efficiency

Our findings support the existence of an ongoing diurnal regulation of transpiration by the light responsive transcription factors HY5 and PIF4 in the stomata, which ultimately determine the whole-plant water use efficiency.

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Research articles

  • Kelly G, Plesser E, Bdolach E, Arroyave M, Belausov E, Faigenboim A, Rozen A, Zemach H, Zach Y Y, Goldenberg L, Arad T, Yaniv Y, Sade N, Sherman A, Eyal Y and Carmi N. (2025) In-planta genome editing approach in citrus using developmental regulators. The Plant Journal. 122 (2): e70155 Link 

  • Goldenberg L, Ghuge S, Faigenboim A, Weissberg M, Shaya F, Rozen A, Dahan Y, Plesser E, Kelly G, Yaniv Y, Arad T, Ophir R, Sherman A, Carmi N, Eyal Y. (2025) A Citrus 2OGD multi-gene cluster encompasses functional and tissue specificity that direct furanocoumarin and pyranocoumarin biosynthesis in citrus. New Phytologist. 245 (4): 1547-1562. Link 

  • Gal A, Dalal A, Anfang M, Sharma D, Binenbaum J, Muchaki, Kumar R, Egbaria A, Estefani Duarte K, Kelly G, Rodrigo de Souza W, Sade N. (2023) Plasma membrane aquaporins regulate root hydraulic conductivity in the model plant Setaria viridis. Plant physiology. 193 (4): 2640-2660. Link 

  • Kelly G, Yaaran A, Gal A, Egbaria A, Brandsma D, Belausov E, Wolf D, David-Schwartz R, Granot D, Eyal Y, Carmi N, Sade N. (2023) Guard cell activity of PIF4 and HY5 control transpiration. Plant Science. 328: 111583.  Link 

  • Kelly G, Brandsma D, Egbaria A, Stein O, Doron-Faigenboim A, Lugassi N, Belausov E, Zemach H, Shaya F, Carmi N, Sade N, and Granot D. (2021) Guard cells control hypocotyl elongation through HXK1, HY5 and PIF4. Communications Biology. 4: 765. Link

  • Lugassi N, Kelly G, Arad T, Farkash C, Yaniv Y, Yeselson Y, Schaffer A. A, Raveh E, Granot D and Carmi N. (2020) Expression of hexokinase in stomata of citrus fruit reduces fruit transpiration and affects seed development. Frontiers in Plant Science 11: 255. Link

  • Galsurker O, Kelly G, Doron-Faigenboim A, Aruchamy K, Salam B B, Teper-Bamnolker P, Lers A and Eshel D. (2020) Endogenous sugar level is associated with differential heat tolerance in onion bulb scales. Postharvest Biology and Technology 163: 1-7. Link

  • Lugassi N, Yadav B, Egbaria A, Wolf D, Kelly G, Neuhaus E, Raveh E, Carmi N and Granot D. (2019)
    Expression of Arabidopsis hexokinase in tobacco guard cells increases water-use efficiency and confers tolerance to drought and salt Stress. Plants-Basel 8 (12): 613. Link

  • Kelly G, Egbaria A, Khamaisi B, Lugassi N, Attia Z, Moshelion M and Granot D. (2019) Guard-cell hexokinase increases water- use efficiency under normal and drought conditions. Frontiers in Plant Science. Link

  • Negin B, Yaaran A, Kelly G, Zait Y and Moshelion M. (2019) Mesophyll ABA restrains early growth and flowering and does not directly suppress photosynthesis. Plant Physiology 180 (2): 910-925. Link

  • Kottapalli J, David-Schwartz R, Khamaisi B, Brandsma D, Lugassi N, Egbaria A, Kelly G, Granot D. (2018) Sucrose-induced stomatal closure is conserved across evolution. PLoS ONE 13 (10): e0205359. Link

  • Fox H, Doron-Faigenboim A, Kelly G, Bourstein R, Attia Z, Zhou J, Moshe Y, Moshelion M, David-Schwartz R. (2017) Transcriptome analysis of Pinus halepensis under drought stress and during recovery. Tree Physiology 38 (3): 423-441. Link

  • Kelly G, Lugassi N, Belausov E, Wolf D, Khamaisi B, Brandsma D, Kottapalli J, Fidel L, Ben-Zvi B, Egbaria A, Acheampong A.K, Zheng C, Or E, Distelfeld A, David-Schwartz R, Carmi N, Granot D. (2017) The Solanum tuberosum KST1 partial promoter as a tool for guard cell expression in multiple plant species. Journal of Experimental Botany. 68 (11): 2885-2897. Link

  • Kelly G, Sade N, Doron-Faigenboim A, Lerner S, Shatil-Cohen A, Yeselson Y, Egbaria A, Kottapalli J, Schaffer A.A, Moshelion M, Granot D (2017) Sugar and hexokinase suppress expression of PIP aquaporins and reduce leaf hydraulics that preserves leaf water potential. The Plant Journal 91: 325-339 Link

  • Lugassi N, Kelly G, Fidel L, Yaniv Y, Attia Z, Levi A, Alchanatis V, Moshelion M, Raveh E, Carmi N and Granot D (2015) Expression of Arabidopsis hexokinase in citrus guard cells controls stomatal aperture and reduces transpiration. Frontiers in Plant Science 6: 1114. Link

  • Sade N, Shatil-Cohen A, Attia Z, Maurel C, Boursiac Y, Kelly G, Granot D, Yaaran A, Lerner S, Moshelion M (2014) The role of plasma membrane aquaporins in regulating the bundle sheath-mesophyll continuum and leaf hydraulics. Plant Physiology 166 (3) 1609-1620 Link

  • Kelly G, Sade N, Attia Z, Secchi F, Zwieniecki M, Holbrook NM, Levi A, Alchanatis V, Moshelion M, Granot D (2014) Relationship between hexokinase and the aquaporin PIP1 in the regulation of photosynthesis and plant growth. PLoS ONE 9 (2): e87888. Link

  • Kelly G, Moshelion M, David-Schwartz R, Halperin O, Wallach R, Attia Z, Belausov E, Granot D (2013) Hexokinase mediates stomatal closure. The Plant Journal 75: 977-988. Link

  • Kelly G, David-Schwartz R, Sade N, Moshelion M, Levi A, Alchanatis V, Granot D (2012) The pitfalls of transgenic selection and new roles of AtHXK1: a high level of AtHXK1 expression uncouples hexokinase1-dependent sugar signaling from exogenous sugar. Plant Physiology 159: 47-51. Link                                                          


Reviews

  • Granot D and Kelly G (2019) Evolution of guard-cell theories: the story of sugars. Trends in Plant Science 24 (6): 507-518. Link 

  • Lawson T, Simkin A, Kelly G, Granot D (2014) Mesophyll photosynthesis and guard cell metabolism impacts on stomatal behavior. New Phytologist 203: 1064-1081.Link 

  • Granot D, Kelly G, Stein O, David-Schwartz R (2014) Substantial roles of hexokinase and fructokinase in the effects of sugars on plant physiology and development. Journal of Experimental Botany 65 (3):809-819. Link 

  • Granot D, David-Schwartz R, Kelly G (2013) Hexose kinases and their role in sugar-sensing and plant development. Frontiers in Plant Science 4 (44): 1-17. Link



  • The Volcani center, Israel
  •  Mon-Fri - 08:00-19:00