Scharfman HE, MacLusky NJ. (2017) Sex differences in hippocampal area CA3 pyramidal cells. Journal of Neuroscience Research. 95(1-2):563-575

Harte-Hargrove LC, Varga-Wesson A, Duffy AM, Milner TA, Scharfman HE. (2015) Opioid receptor-dependent sex differences in synaptic plasticity in the hippocampal mossy fiber pathway of the adult rat. J. Neurosci. 35(4):1723-38.

Atwi S, McMahon D, Scharfman H, MacLusky NJ. (2014) Androgen Modulation of Hippocampal Structure and Function. Neuroscientist. 2014

Special issue of Neuroscience (2012): Steroid hormones in the CNS: The role of BDNF. Guest Editors: Scharfman HE, Kramar EA, Luine VN, Srivastava D.

Skucas VA, Duffy AM, Harte-Hargrove LC, Magagna-Poveda A, Radman T, Chakraborty G, Schroeder CE, MacLusky NJ, Scharfman HE (2013) Testosterone depletion in adult male rats increases mossy fiber transmission, LTP, and sprouting in area CA3 of hippocampus. J. Neurosci. 33: 2338-2355.

This study showed that gonadectomized male rats exhibited increased BDNF expression in the mossy fiber axons of hippocampus.  Mossy fiber transmission was increased in a manner consistent with a new BDNF-dependence.  Mossy fiber LTP was also increased and exhibited dependence on trkB receptors, which bind BDNF.  The mossy fiber axons exhibited sprouting into stratum oriens, and current source density analysis suggested that new synapses were made by sprouted axons. The remarkable increase in plasticity in gonadectomized male rats suggests that testosterone may normally suppress excitability and plasticity, which could be advantageous to prevent seizures and excitotoxicity, but at the ‘price’ of reduced plasticity.

Barouk S, Hintz TM, Li P, Maclusky NJ, Scharfman HE (2011) 17β-estradiol increases astrocytic vasclar endothelial growthfactor (VEGF) in female rat hippocampus. Endocrinology 152:1745-1751.

This paper shows that 17β-estradiol increases VEGF in the female rat, which is important because VEGF increases vascular permeability and angiogenesis.  Interestingly, the increase in VEGF was associated with astrocytes as well as blood vessels, suggesting a mechanism regulating the glia-vascular interface.


Supplemental Information:


Scharfman HE, Hintz TM, Kim M, MacLusky NJ (2009) Seizures and reproductive function: insights from an animal model. Ann Neurol 54:687-697.


Scharfman HE, Malthankar-Phatak GH, Friedman D, Pearce PP, McCloskey DP, Harden CJ, MacLusky NJ. (2009) A rat model of epilepsy in women: a tool to study the physiological interaction between the endocrine system and seizures. Endocrinology. 150 :4437-4452.


Scharfman HE, MacLusky NJ (2008) Estrogen-growth factor interactions and their contribution to neurological disorders. Headache 48: S2 77-89.

This article reviews the evidence that BDNF contributers to the actions of estrogen. The emphasis is on neurological disorders such as epilepsy and migraine.


MacBeth AH, Scharfman HE, MacLusky NJ, Luine VN (2008) Effects of multiparity on recognition memory, monoaminergic neurotransmitters, and brain-derived neurotrophic factor (BDNF). Horm Beh., 76: 36-44.


Scharfman HE, Hintz TM, Gomez J, Stormes KA, Barouk S, Malthanakar--Phatak GH, McCloskey DP, Luine VN, MacLusky NJ (2007) Changes in hippocampal fuction in ovariectomized rats after sequential low doses of estradiol to simulate the preovulatory estrogen surge. Eur J Neurosci.  26: 2595-2612.

This study showed that the effects of the estrous cycle in the female rat (J. Neurosci. 2003) could be explained by the rise in estradiol levels during the estrous cycle., and that BDNF played an important role.  It also provides the methods for a protocol that can be used in female rats to simulate the serum levels of estradiol during the preovulatory surge. 


Scharfman HE, MacLusky NJ (2006) The influence of gonadal hormones on excitability, seizures and epilepsy in the female. Epilepsia 47:1423-1440.



Scharfman HE, MacLusky NJ (2006) Estrogen and BDNF: complexity of steroid hormone-growth factor interactions. Front. Neuroendocrin. 27:415-435.

This review addresses the actions of estrogen that are potentially mediated by BDNF and NPY. It was published in a special issue that highlighted a symposium at one of the annual Society for Behavioral Neuroendcrinology meetings about reproductive steroids and growth factors.  The preface is provided below as well as a link to the article.



Staley K, Scharfman HE (2005) A woman’s preogative. Nature Neurosci 8:697-699.


Scharfman HE, Goodman JH, Rigoulot M, Berger R, Walling SW, Mercurio TM, Stormes KA, MacLusky NJ (2005) Seizure susceptibility of intact and ovariectomized female rats treated with the convulsant pilocarpine. Exp Neurol 196: 73-86.


Scharfman HE, MacLusky NJ (2004) Similarities between BDNF and estrogen: coincidence or clue? Trends Neurosci. 28: 79-85.


Scharfman HE et al (2003) Hippocampal excitability changes across the estrous cycle in the rat: a potential role for BDNF. J Neurosci. 23:11641-11652.

This paper showed that female rats have a cyclical increase in mossy fiber excitability and mossy fiber BDNF expression that follows the preovulatory estrogen surge.  The physiological changes in mossy fiber transmission were blocked by an antagonist of trk receptors, implicating BDNF mechanistically.