[Full List of Publication :]


  • Chen, Q., Leshkowitz, D., Li, H., van Impel, A., Schulte-Merker, S., Amit, I., Rizzoti, K. and G. Levkowitz. (2023). Neural plate progenitors give rise to both anterior and posterior pituitary cells. Developmental Cell, pubmedPDF
  • Akinrinade, I., Kareklas, K., Reis, T.K., Gliksberg, M., Petri, G., Levkowitz, G., R.F. Oliveira. (2023) Evolutionarily conserved role of oxytocin in social fear contagion in zebrafish. Science 379, 1232–1237. pubmed, PDF  
     * Highlighted in "Science Perspective" by Ross S. Deangelis and Hans A. Hofmann  – The spread of fear in an empathetic fish. Science 379, 1186-1187. PDF
  • Parab, S., Card, O. A. Chen, Q., Buck, L. D., Quick, R. E., Horrigan, W. F., Levkowitz, G., Vanhollebeke, B., and R.L. Matsuoka. (2023) Local angiogenic interplay of Vegfc/d and Vegfa controls brain region-specific emergence of fenestrated capillaries. eLife  12:e86066. doi: 10.7554/eLife.86066. pubmed, PDF
  • Swaminathan, A., Gliksberg, M., Anbalagan, S., Wigoda, N. and G. Levkowitz. (2023) Stress resilience is established during development and is regulated by complement factors. Cell Reports 42, 111973 pubmedPDF
  • Peles, G., Swaminathan, A. and G. Levkowitz. (2022). Glucocorticoid-sensitive period of corticotroph development – implications for early life stress.  Journal of Neuroendocrinology e13229. pubmed  PDF
  • Nunes, A.R#, Gliksberg, M.#, Varela, S.A.M, Teles, M., Wircer, E., Blechman, J., Petri, G., Levkowitz, G.* and R.F. Oliveira* (2021) Developmental effects of oxytocin neurons on social affiliation and processing of social information. Journal of Neuroscience 10.1523/JNEUROSCI.2939-20.2021. pubmed  PDF
    #equal contribution(co-first); *co-corresponding authors 
     * Highlighted in This Week in the Journal (October 20, 2021 – Oxytocin Affects Development of Dopamine Neurons. PDF
  • Rajamannar , P. Arokidhas, I., Levkowitz, G.* and Biran, J.* (2021) The neurohypophysis and urophysis - ancient Piscine neurovascular interfaces. In Neuroanatomy of Neuroendocrine Systems, V. Grinevich and A. Dobolyi, eds. (Springer-Nature) Masterclass in Neuroendocrinology 12, PDF
    *co-corresponding authors
  • Gordon L. and G. Levkowitz. (2021). Through the open window: How does the brain talk to the body? Fronteirs for Young Minds. doi: 10.3389/frym.2021.534184. PDF
  • Segev-Hadar, A., Krispin, S., Olthof, A.M., Hyatt, K.C., Haller, L., Barki, A., Nitzan, T., Levkowitz, G., Kanadia, R.N., Cnaani, A. and J. Biran. (submitted) Oxytocin Signaling Regulates the Homeostatic Response to Cold Stress in Poikilothermic Vertebrates.  bioRxiv 2021.12.20.472748; doi:


  • Biran, J., Gliksberg, M., Shirat, I., Swaminathan, A., Levitas-Djerbi, T., Appelbaum, L. and G. Levkowitz. (2020). Splice-specific deficiency of the PTSD-associated gene PAC1 leads to a paradoxical age-dependent stress behavior. Scientific Reports 10:9559. pubmedPDF 

    * Faculty Opinions (formerly F1000prime) Recommendation by Pisegna J: In Faculty Opinions, 26 Jun 2020; 10.3410/f.738137336.793575648.

  • Ribeiro, D., Nunes, A.R., Teles, M.C., Anbalagan, S., Blechman, J., Levkowitz, G. and R.F. Oliveira. (2020). Genetic variation in the social environment affects behavioral phenotypes of oxytocin receptor mutants in zebrafish. eLife 2020;9:e56973 DOI: 10.7554/eLife.56973. pubmed, PDF 

    *Highlighted in eLife’s "Insight Article"  by Elena Dreosti. Social Behaviours: Nurturing nature. eLife 2020;9:e61323 DOI: 10.7554/eLife.61323

  • Chen, Q., Leshkowitz, D. Blechman, J. and G. Levkowitz. (2020). Single-cell molecular and cellular architecture of the mouse neurohypophysis. eNeuro  10.1523/ENEURO.0345-19.2019 pubmedPDF 

    *Highlighted in eNeuro's "Featured Research"  

  • Journal of Neuroendocrinology coverWagner S., Grinevich V. and G. Levkowitz (2020). Advances in Neurohypophysial Hormones Research. Journal of Neuroendocrinology e12853 pubmedPDF 
* Special Issue of papers from the 13th World Congress on Neurohypophysial Hormones (WCNH2019). Edited by:  Gil Levkowitz Shlomo Wagner Valery Grinevich.


  • Nunes, A.R., Carreira, L., Anbalagan, S., Blechman, J. Levkowitz, G. and R.F. Oliveira. (2020). Perceptual mechanisms of social affiliation in zebrafish. Scientific Reports  10:3642 pubmed, PDF
  • Ribeiro, D., Nunes, A.R., Gliksberg, M., Anbalagan, S., Levkowitz, G. and R.F. Oliveira. (2020). Oxytocin receptor signaling impairs novelty recognition but not social preference in zebrafish. Journal of Neuroendocrinology  e12834 doi:10.1111/jne.12834. pubmedPDF
  • Gordon L., Blechman J., Shimoni E., Gur D., Anand-Apte B. and G. Levkowitz. (2019). Fenestrae-associated protein Plvap regulates the rate of blood-borne proteins passage into the hypophysis. Development  146: dev177790. doi:10.1242/dev.177790. pubmedPDF
  • Robinson, K.J., Bosch, O.J., Levkowitz, G., Busch, K.E, Jarman, A. and M.Ludwig. (2019). Social creatures: model animal systems for studying the neuroendocrine mechanisms of social behaviour. Journal of Neuroendocrinology 12:e12807 doi: 10.1111/JNE.12807. pubmed, PDF
  • Anbalagan S., Blechman J., Gliksberg M., Gordon L., Rotkopf R., Dadosh T., Shimoni E. and G. Levkowitz. (2019). Robo2 regulates synaptic oxytocin content by affecting actin dynamics. eLife 2019;8:e45650 DOI: 10.7554/eLife.45650. pubmedPDF
  • Denichenko P., Mogilevsky M., Cléry A., Welte T., Biran J., Shimshon O., Barnabas G.D., Danan-Gotthold M., Kumar S., Yavin E., Levanon E.Y., Allain F.H., Geiger T., Levkowitz G.and R. Karni (2019). Specific inhibition of splicing factor activity by decoy RNA oligonucleotides. Nature Communications  10:1590. pubmed,  PDF
  • Anbalagan S, Gordon L, Blechman J, Matsuoka R, Rajamannar P, Wircer E, Biran J, Reuveny A, Leshkowitz D, Stainier D.Y.R, Levkowitz G. (2018). Pituicyte cues regulate the development of permeable neuro-vascular interfaces. Developmental Cell  47, 711–726.
    pubmed, PDF 

    "Featured Article" in Dev. Cell's December 17th issue 
    * Highlighted in a Dev. Cell Preview by Thomas, J.L. and Eichmann, A. (2018). To BBB or Not to BBB. Developmental Cell 47, 689-691.

  • Blechman J., S. Anbalagan, G.G. Matthews and G. Levkowitz. (2018). Genome editing reveals idiosyncrasy of CNGA2 ion channel-directed antibody immunoreactivity towards oxytocin. Front. Cell Dev. Biol. 6:117. doi: 10.3389/fcell.2018.00117. pubmedPDF
  • Biran, J., Wircer, E., Blechman, J. and G. Levkowitz.  (2018). Development and function of the zebrafish neuroendocrine system. In: Model animals in neuroendocrinology: From worm to mouse to man, M. Ludwig and G. Levkowitz, eds. (Wiley-Blackwell), pp101-131. PDF
  • D. Gur, J.‐D. Nicolas, V. Brumfeld, O. Bar‐Elli, D. Oron, G. Levkowitz. (2018). The Dual Functional Reflecting Iris of the Zebrafish. Advanced Science 5(8):1800338. pubmedPDF  
Highlighted in Advanced Science News: "Fishing for the Structure of the Iris"
  • Edited book: Ludwig M, Levkowitz G. eds. (2018) Model Animals in Neuroendocrinology: From Worm to Mouse to Man. Hoboken, NJ: Wiley Blackwell. ISBN: 978-1-119-39094-7. PDF 
*Part of the “Masterclass in Neuroendocrinology Series”, written for students, trainees, established researchers, and teachers as a joint venture between the International Neuroendocrine Federation (INF) and Wiley-Blackwell. 
  • Gliksberg M. and G. Levkowitz. (2017). Smells Familiar: Pheromone-induced neurotransmitter switching mediates social discrimination. Neuron 95:1229-1231. pubmed, PDF
  • Wircer E., J. Blechmann, N. Borodovsky, M. Tsoory, A.R. Nunes, R.F. Oliveira and G. Levkowitz. (2017). Homeodomain protein Otp affects developmental neuropeptide switching in oxytocin neurons associated with a long-term effect on social behavior. eLife 2017;6: e22170. DOI: 10.7554/eLife.22170. pubmed, PDF
  • Blechman J., Levkowitz, G. and Y. Gothilf. (2017). The not-so-long history of zebrafish research in Israel. Int. J. Dev. Biol. 61:149-157. pubmedPDF 
  • Biran J, and G. Levkowitz.  (2016). Zebrafish reel in phenotypic suppressors of autism. Neuron 89:673-675.
  • Wircer E., S. Ben-Dor and G. Levkowitz. (2016).  In: Non-mammalian models for neurohypophysial peptides. In Molecular Neuroendocrinology: From Genome to Physiology, D. Murphy and H. Gainer, eds. (Wiley-Blackwell), pp. 301–328. PDF


  • Biran J, Tahor M, Wircer E and G. Levkowitz.  (2015). Role of developmental factors in hypothalamic function. Frontiers in Neuroanatomy 9:47. doi:10.3389/fnana.2015.00047.
  • Blechman J. and G. Levkowitz. (2013). Alternative splicing of the pituitary adenylate cyclase-activating polypeptide receptor PAC1: mechanisms of fine tuning of brain activity. Frontiers in Endocrinology. May 21;4:55. doi: 10.3389/fendo.2013.00055.
    pubmed, PDF
  • Gutnick A. and G. Levkowitz. (2012). The neurohypophysis – fishing for new insights. J. Neuroendo. 24(6):973-974.
    Also published as Topical Briefings of the British Society for Neuroendocrinology.
  • Amir-Zilberstein L., J. Blechman, Y. Sztainberg, W.H.J. Norton, A. Reuveny, J.L., N. Borodovsky, M. Tahor, Bonkowsky, L. Bally-Cuif, A. Chen and G. Levkowitz. (2012). Homeodomain protein Otp and activity-dependent splicing modulate neuronal adaptation to stress. Neuron 73:279-291.
    pubmed, PDF
    * Selected for Neuron’s cover illustration (Jan. 26 issue) and highlighted in Neuron’s Spotlight On Disease  – Regulating Stress through Splicing.
  • Blechman J., A. Gutnick, L. Amir-Zilberstein and G. Levkowitz. (2011). The metabolic regulator PGC-1α directly controls the expression of the hypothalamic neuropeptide oxytocin. J. Neurosci. 31(42):14835-14840.
    pubmed, PDF
     * Highlighted in This Week in the Journal (October 19, 2011)  – PGC-1α may suppress eating while increasing metabolism.
  • Gutnick, A., J. Blechman, J. Kaslin, L. Herwig, H-G. Belting, M. Affolter, J.L. Bonkowsky and G. Levkowitz. (2011). The hypothalamic neuropeptide oxytocin is required for formation of the neuro-vascular interface of the pituitary. Developmental Cell 21:642–654.
    pubmed, PDF 
    * Highlighted in a Dev. Cell Preview by Cariboni, A., and Ruhrberg, C. (2011). The hormone of love attracts a partner for life. Dev. Cell 21, 602-604. PDF
  • Machluf, Y., Gutnick A. and Levkowitz G. (2011).  Development of the zebrafish hypothalamus.  Ann. N.Y. Acad. Sci. 1220(1):93-105
  • Machluf, Y. and Levkowitz G. (2011). Visualization of mRNA expression in the zebrafish embryo. Methods Mol. Biol. 714:83-102.


  • Russek-Blum N., Nabel-Rosen H. and Levkowitz G. (2010). Two-photon- based photo activation in live zebrafish embryos. J. Vis. Exp. 46: e1902.
    pubmedvideo article link
  • Palevitch, O., Abraham E. , Borodovsky N., Levkowitz G., Zohar Y. and Gothilf Y. (2010). Cxcl12a-Cxcr4b signaling is important for proper development of forebrain GnRH system in zebrafish. Gen. and Comp. Endocrinology 165(2):262-268.
  • Russek-Blum N, Nabel-Rosen H and Levkowitz G. (2009). High resolution fate map of the zebrafish diencephalon. Dev. Dyn. 238(7):1827-1835.
    pubmed, PDF
  • Borodovsky N, Ponomaryov T, Frenkel S and Levkowitz G. (2009). Neural protein Olig2 acts upstream of the transcriptional regulator Sim1 to specify diencephalic dopaminergic neurons. Dev. Dyn. 238(4):826–834.
    pubmed, PDF
  • Palevitch O, Abraham E, Borodovsky N, Levkowitz G, Zohar Y and Gothilf Y. (2009). Nasal embryonic LHRH factor plays a role in the developmental migration and projection of gonadotropin-releasing hormone 3 neurons in zebrafish. Dev. Dyn. 238(1):66-75.
    pubmed, PDF
  • Russek-Blum, N., Gutnick, A., Nabel-Rosen, H., Blechman, J., Staudt, N., Dorsky, R.I., Houart, C. and Levkowitz, G. (2008) Dopaminergic neuronal cluster size is determined during early forebrain patterning. Development 135(20):3401-3413.
    pubmed, PDF
  • Blechman, J., Borodovsky, N., Eisenberg, M., Nabel-Rosen, H., Grimm, J. and Levkowitz, G. (2007) Specification of hypothalamic neurons by dual regulation of the homeodomain protein Orthopedia. Development 134(24):4417-4426.
    pubmed, PDF
  • Wilcock, D. M., A. Rojiani, A. Rosenthal, G. Levkowitz, S. Subbarao, J. Alamed, D. Wilson, N. Wilson. M. J. Freeman, M. N. Gordon, D. Morgan. 2004, Passive amyloid immunotherapy clears amyloid and transiently activates microglia in a transgenic mouse model of amyloid deposition. 2004. Journal of Neuroscience 24: 6144-6151. pubmed, PDF
  • Levkowitz, G., Zeller, J., Sirotkin, H.I., French, D., Schilbach, S., Hashimoto, H., Hibi, M., Talbot, W.S. and Rosenthal, A. (2003) Zinc finger protein too few controls the development of monoaminergic neurons. Nat. Neurosci. 6(1):28-33.
    pubmed, PDF