Associate Professor

Basic Sciences
Division of Pharmacology
School of Medicine
Loma Linda University
Loma Linda, CA 92350

Phone: (909) 651-5334
Fax: (909) 558-0119

Research Interests:

•Cardio/Cerebrovascular regulation of blood flow during aging and development of neurodegenerative disease

•Genetic, structural, and functional basis of the interplay between calcium, oxidative, and electrical signaling in vascular cells and neurons

Selected Publications:

  1. Hakim MA and Behringer EJ. Development of Alzheimer's disease progressively alters sex-dependent KCa and sex-independent KIR channel function in cerebrovascular endothelium. J Alzheimer’s Dis., 76(4):1423-1442, 2020. DOI: 10.3233/JAD-200085.
  2. Hakim MA, Nye-Chum P, Buchholz JN, Behringer EJ. Aging alters cerebrovascular endothelial GPCR and K+ channel function: Divergent role of biological sex. J Gerontol A Biol Sci Med Sci. pii:glz275, 2019. DOI: 10.1093/gerona/glz275.
  3. Stoll S, Xi J, Ma B, Leimena C, Behringer EJ, Qin G, Qiu H. The valosin-containing protein protects the heart against pathological Ca2+ overload by modulating Ca2+ uptake proteins. Toxicol Sci. pii: kfz164, 2019. DOI: 10.1093/toxsci/kfz164.
  4. Behringer EJ and Hakim MA. Functional Interaction among KCa and TRP Channels for Cardiovascular Physiology: Modern Perspectives on Aging and Chronic Disease. Int. J. Mol. Sci. 20: 1380, 2019. DOI:10.3390/ijms20061380.
  5. Hakim MA and Behringer EJ. Simultaneous Measurements of Intracellular Calcium and Membrane Potential in Freshly Isolated and Intact Mouse Cerebral Endothelium. J Vis Exp. Jan 20: 143, 2019. DOI: 10.3791/58832.
  6. Hakim MA, Buchholz JN, Behringer EJ. Electrical dynamics of isolated cerebral and skeletal muscle endothelial tubes: differential roles of G-Protein coupled receptors and K+ channels. Pharmacol Res Perspect. 6(2): e00391, 2018.
  7. Kapela A, Behringer EJ, Segal SS and Tsoukias NM. Biophysical properties of microvascular endothelium: Requirements for initiating and conducting electrical signals. Microcirculation. 25(2): 1-15, 2018.
  8. Behringer EJ*, Scallan JP*, Jafarnejad M, Castorena-Gonzalez JA, Zawieja SD, Moore JE Jr, Davis MJ, Segal SS. Calcium and electrical dynamics in lymphatic endothelium. J Physiol. 595(24): 7347-7368, 2017. *Equal contribution.
  9. Behringer EJ and Segal SS. Impact of aging on calcium signaling and membrane potential in endothelium of resistance arteries: A role for mitochondria. J Gerontol A Biol Sci Med Sci. 72(12): 1627-1637, 2017.
  10. Behringer EJ. Calcium and electrical signaling in arterial endothelial tubes: New insights into cellular physiology and cardiovascular function. Microcirculation. 24(3): 1-12, 2017.
  11. Behringer EJ and Segal SS. Membrane potential governs calcium influx into microvascular endothelium: Integral role for muscarinic receptor activation. J Physiol. 593(20): 4531-48, 2015.
  12. Socha MJ, Boerman EM, Behringer EJ, Shaw RL, Domeier TL, and Segal SS. Advanced age protects microvascular endothelium from aberrant Ca2+ influx and cell death induced by hydrogen peroxide. J Physiol. 593(9): 2155-69, 2015.
  13. Behringer EJ, Shaw RL, Westcott EB, Socha MJ, and Segal SS. Aging impairs electrical conduction along endothelium of resistance arteries through enhanced Ca2+-activated K+ channels activation. Arterioscler Thromb Vasc Biol. 33(8): 1892-1901, 2013.
  14. Behringer EJ and Segal SS. Spreading the signal for vasodilatation: Implications for skeletal muscle blood flow control and the effects of aging. J Physiol. 590 (Pt 24): 6277-84, 2012.
  15. Socha MJ, Domeier TL, Behringer EJ, Segal SS. Coordination of intercellular Ca2+ signaling in endothelial cell tubes of resistance arteries. Microcirculation. 19(8): 757-70, 2012.
  16. Behringer EJ and Segal SS. Tuning electrical conduction along endothelial cell tubes through Ca2+-activated K+ channels. Circ Res. 110(10): 1311-1321, 2012.
  17. Behringer EJ, Socha MJ, Polo-Parada L, Segal SS. Electrical conduction along endothelial cell tubes from mouse feed arteries: confounding actions of glycyrrhetinic acid derivatives. Br J Pharmacol. 166(2): 774-787, 2012.
  18. Socha MJ*, Behringer EJ*, Segal SS. Calcium and electrical signaling along endothelium of the resistance vasculature. Basic Clin Pharmacol Toxicol. 110(1): 80-86, 2012. *Equal contribution
  19. Behringer EJ, Leite LD, Buchholz NE, Keeney M, Pearce WJ, Vanterpool CK, Wilson SM and Buchholz JN. Maturation and long-term hypoxia alters calcium-induced calcium release in sheep cerebrovascular sympathetic neurons.  J Appl Physiol. 107(4): 1223-1234, 2009.
  20. Behringer EJ, Vanterpool CK, Pearce WJ, Wilson SM and Buchholz JN.  Advancing age alters the contribution of calcium release from smooth endoplasmic reticulum stores in superior cervical ganglion cells. J Gerontol A Biol Sci Med Sci. 64A: 34-44, 2009.
  21. Buchholz JN, Behringer EJ, Pottorf BJ, Pearce WJ and Vanterpool CK.  Age–dependent changes in Ca2+ homeostasis in peripheral neurons:  implications for changes in function.  Aging Cell. 6(3):  285-296, 2007.


Book Chapters

  1. Buchholz JN and Behringer EJ (Editors). Calcium and Signal Transduction. Physiology Series, Volume 1. Publisher, Intech Open; ISBN: 978-1-78984-249-4, 2019.
  2. Behringer EJ and Segal SS. Vascular Ion Channels in Physiology and Disease, Chapter title: Ion Channels in Control of Blood Flow: Electrical Conduction Along Endothelium of Resistance Arteries. Publisher, Springer. ISBN: 978-3319296333, 2016.
  3. Buchholz JN, Pottorf WJ, Vanterpool CK, Behringer EJ and Duckles SP. Senescence, Chapter title: Calcium Regulation in Neuronal Function with Advancing Age: Limits of Homeostasis. Publisher, In-Tech Open Access. ISBN: 978-953-308-28-6, 2012.