Table 1.

Summary of the effects of ovarian hormones on cognition

Ability	Sample Characteristics	Younger	Older	References
Attention
	Humans
	Premenopause	F > L		Pletzer et al., 2014
	Postmenopause (55–65 years)		HRT > no HRT	Castonguay et al., 2015
	Postmenopause		HRT > no HRT	Fedor-Feyberg, 1977;
	Postmenopause (∼60 years)		ERT > no ERT	Schmidt et al., 1996
	Postmenopause (∼65 years)		HRT > no HRT	Smith et al., 2001
	Postmenopause (∼53 years)		HRT = no HRT	Gleason et al., 2015
	Postmenopause (47–65 years)		ERT = no ERT	Polo-Kantola et al., 1998b
	Postmenopause (53–72 years)		HRT = no HRT	Alhola et al., 2006
	Postmenopause (∼53 years)		HRT = no HRT	Keenan et al., 2001
	Postmenopause (58–75 years)		HRT = no HRT	Wolf et al., 2005
	Animals
	OVX macaques (6–11 years)	OVX < intact E2 > vehicle		Voyko 2002
	OVX macaques (7–16 years)	E2 = Vehicle		Lacreuse et al., 2009
	OVX macaques (∼20 years)		E2 > No E2	Voytko et al., 2009
	OVX macaques (18–25 years)		E2 > No E2	Kohama et al., 2016
	17-mo OVX rats		E2 > No E2	Bohacek & Daniel, 2010
	Intact rats	Proestrous < Estrous/Diestrous		Quinlan et al., 2010
		E2 < Vehicle		Almey et al., 2013
Non-spatial working memory
	Humans
Maintenance
Digit Span forward:	Postmenopause		HRT = no HRT	Alhola et al., 2006; Castonguay et al., 2015; Duff & Hampson, 2000; Gleason et al., 2015; Keenan et al., 2001; Polo-Kantola et al., 1998b; Schmidt et al., 1996
Manipulation/Updating
		Early F = Late F		Joseph et al., 2012
	Postmenopause (∼56 years)		HRT > no HRT	Duff & Hampson, 2000
	Postmenopause (55–65 years)		HRT = no HRT HRT > no HRT	Castonguay et al., 2015
	Postmenopause (∼53 years)		HRT = no HRT	Gleason et al., 2015
	Postmenopause (∼53 years)		HRT > no HRT	Keenan et al., 2001
	Postmenopause (50–65 years)		3 days ERT > no ERT	Krug et al., 2006
	Postmenopause (∼69 years)		HRT = no HRT	Janowsky et al., 2000
	Postmenopause (∼70 years)		ERT > no ERT	Baker et al., 2012
	Animals
Object-DRST Face-DRST	OVX macaques (6–9 years)	EE2 = no EE2 EE2 < no EE2		Lacreuse & Herndon, 2003
Object-DRST	OVX macaques (7–16 years)	EB = no EB		Lacreuse et al., 2009
	OVX rats	Low EB > no EB Mod-high EB < no EB		Wide et al., 2004
	OVX rats	High E2 < no E2		Wang et al., 2008
	OVX rats	High E2 < no E2	High E2 < no E2	Wang et al., 2009
Task switching
	Humans
	Premenopause	GnRHa = GnRHa + E/P GnRHa: ↓ DLPFC GnRHa + E/P: ↑ DLPFC		Berman et al., 1997
	Postmenopause (∼60 years)		ERT > no ERT	Schmidt et al., 1996
	Postmenopause (55–65 years)		HRT > no HRT	Castonguay et al., 2015
	Early postmenopause (∼52 years)		↑DLPFC in HRT users; correlated with performance	Girard et al., 2017
	Postmenopause (∼65 years)		3-mo ERT = no ERT	Duka et al., 2000
	Animals
	OVX macaques (∼20 years)		HRT > no HRT	Voytko et al., 2009
	OVX macaques (∼24 years)		EE2 = no EE2	Lacreuse et al., 2004
	Young OVX rats	E2 > no E2		Lipatova et al., 2016
reversal learning	OVX marmosets (3–5 years)		High E2 < No E2	Lacreuse et al., 2014
reversal learning	OVX rats (21–22 mo)		E2 < no E2	Gibbs et al., 2011
Response inhibition
	Humans
Go/No-Go, Stop-Signal	Premenopause	F < L		Colzato et al., 2010; Reimers et al., 2014
Stroop Interference	Postmenopause (50–65 years)		3 days ERT > no ERT	Krug et al., 2006
Stroop Interference	Postmenopause		HRT = no HRT	Alhola et al., 2006; Baker et al., 2012; Castonguay et al., 2015; Duka et al., 2000; Polo-Kantola et al., 1998b; Wolf et al., 2005
	Animals
Stop Signal	Premenopausal baboons	O = L		Lacreuse et al., 2016
DRL	OVX rats	High E2 < no E2		Wang et al., 2008
DRL	OVX rats	High E2 < no E2	E2 = no E2	Wang et al., 2011
Decision making
	Humans
	Premenopause & postmenopause (∼55 years)	Menses = mid L	ERT = no ERT	Reavis & Overman, 2001
	Premenopause	F > menses		Smith et al., 2014
	Animals
	OVX rats	High EB > no EB		Uban et al., 2012
Spatial Working Memory
	Humans
	Premenopause	High E2 > Low E2 E2 correlates with performance		Hampson & Morley 2013
	Postmenopause (∼56 years)		HRT > no HRT	Duff & Hampson, 2000
	Postmenopause (∼71 years)		3-mo ERT = no ERT	Schiff et al., 2005
	Animals
	Premenopause	O < F=L		Lacreuse et al., 2001
	OVX macaques (6–9 years)	EE2 = no EE2		Lacreuse et al., 2003
	OVX macaques (7–16 years)	E2 = Vehicle		Lacreuse et al., 2009
	Macaques (6–10 years)	OVX = intact E2 = no E2		Voytko, 2000
	Macaques (∼ 10 years)	E2 = no E2		Hao et al., 2007
	Macaques (20–27 years)		Premenopause > peri/postmenopause	Roberts et al., 1997
	OVX macaques (∼22 years)		E2 > no E2	Rapp et al., 2003
	OVX macaques (18–25 years)		E2 > E2/P > No E2	Kohama et al., 2016
	OVX macaques (21–24 years)		EE2 > no EE2	Lacreuse et al., 2002
	OVX macaques (∼20 years)		E2 = E2/P = no E2	Voytko et al., 2008
	OVX marmosets (3–5 years)		High E2 < No E2	Lacreuse et al., 2014
	OVX macaques (19–27 years)		Long-term OVX > intact	Lacreuse et al., 2000
	Intact rats	Proestrous > estrous		Pompili et al., 2010
	OVX rats	E2 > no E2		Gibbs, 2007; Holmes et al., 2002; Luine et al., 1998; Sandstrom & Williams, 2001; Velàzquez-Zamora et al., 2012
	OVX rats	Win-stay: E2 = no E2 Win-shift: E2 < no E2		Galea et al., 2001
	OVX rats	Intra-PFC/HPC E2 > vehicle		Sinopoli et al., 2006
	12-mo old OVX rats		E2 > no E2	Daniel et al., 2006
	21–25-mo old OVX rats		E2+P > no E2+P	Gibbs, 2000
	13–22-mo old OVX rats		OVX < intact E2 > no E2	Markowska & Savonenko, 2002
Spatial reference memory
	Animals
	Intact rats	Proestrous = estrous		Berry & McMahan, 1997;
	Intact rats	Watermaze: Proestrous < estrous RAM: Proestrous = estrous		Pompili et al., 2010
		Proestrous < estrous		Warren & Juraska, 1997
	OVX rats	Watermaze: E2 > no E2 Y-maze: E2 = no E2		McLaughlin, 2008
		E2 > no E2 Intra-HPC E2 > vehicle		Packard & Teather, 1997a,b
	OVX rats	E2 < no E2		Galea et al., 2001
	OVX rats	E2 = no E2		Fader et al., 1999; Holmes et al., 2002; Luine et al., 1998;
	4, 16, & 24-mo old OVX rats	E2 > no E2	E2 = no E2	Talboom et al., 2008
	13–22-mo old OVX rats		OVX = intact E2 = no E2	Markowska & Savonenko, 2002
Object placement/NOIP test
	Animals
	OVX rats	E2 > no E2		As reviewed by Tuscher et al., 2014; McLaughlin et al., 2008
Spatial strategy
	Humans
	Premenopause	Late L > Early F/O		Hussain et al., 2016
	Animals
	Rats	Proestrous > estrous E2 > no E2	E2 > no E2	As reviewed by Korol & Pisani, 2015; Korol et al., 2004
	OVX rats	Intra-HPC E2 > Vehicle		Zurkovsky et al., 2007
Verbal memory
	Humans
	Premenopause	Inconsistent reports, or F = L = Menses		As reviewed by Poromaa & Gingnell, 2014
	Premenopause	GnRHa < cycling		Craig et al., 2008
		GnRHa = cycling		Guerrieri et al., 2016; Owens et al., 2002
	Age: ∼45–52 years		Peri-/postmenopause/ < premenopause	Greendale et al., 2009; Weber et al., 2013
	Surgically menopausal women (36–45 years)		BL > post-surgery	Sherwin, 1988
	Postmenopause		HRT > no HRT	Baker et al., 2012; Castonguay et al., 2015; Jacobs et al., 1998; Kampen & Sherwin, 1994; Keenan et al., 2001; Maki, Zonderman, & Resnick, 2001; Phillips & Sherwin, 1992b; Resnick et al., 1998
	Postmenopause		HRT = no HRT	Baker et al., 2012; Gleason et al., 2015; Henderson et al., 2016
Visual recognition memory
	Humans
	Premenopause	GnRHa < cycling		Craig et al., 2010
	Postmenopause (∼65–67 years)		ERT > no ERT	Resnick et al., 1998
	Postmenopause (∼64 years)		HRT = no HRT	Alhola et al., 2006
	Older AD patients (∼74 years)		ERT = no ERT	Sundermann et al., 2006
	Animals
	Premenopause	No/brief retention delay: F = L 30-s retention delay: O < F=L		Kromrey, Czoty, & Nader, 2015; Lacreuse et al., 2001
	OVX macaques (6–9 years)	EE2 = no EE2		Lacreuse & Herndon, 2003
	OVX macaques (7–16 years)	E2 = Vehicle		Lacreuse et al., 2009
	OVX macaques (∼20–24 years)		HRT > no HRT	Lacreuse et al., 2002; Rapp et al., 2003; Voytko et al., 2008
	OVX rats with low E2	Intra-PRh E2 < vehicle		Gervais et al., 2013, 2016
Novel object preference
	Rats	Proestrous > estrous E2 > no E2 Intra-HPC > vehicle Intra-PRh > vehicle	Intra-HPC E2 = vehicle	As reviewed by: Galea, Frick, Hampson, Sohrabji, & Choleris, 2016; Luine, 2015; Tuscher et al., 2014

Note: AD = Alzheimer’s Disease; DLPFC = dorsolateral prefrontal cortex; DRST = delayed recognition span test; E2 = 17β-estradiol; EB = estradiol benzoate; EE2 = ethinyl estradiol; ERT = estrogen replacement therapy; F = follicular phase; GnRHa = gonadotropin releasing hormone agonist; HPC = hippocampus; HRT = hormone replacement therapy; L = luteal; mo = month; O = Ovulatory; P = progesterone; PRh = perirhinal cortex; OVX = ovariectomized.