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According to evolutionary theory, the development of sex-differences in spatial ability can be predicted in species in which one sex’s range is different from the other sex’s range. This model holds for polygynous rodent species (including the laboratory rat), primates, and humans, to name a few. In such ancestrally polygynous species such as our own, males evolved to have higher working memory in order to find and remember the moving and depleting resources over larger ranges that are associated with hunting while females evolved to have a higher reference memory that allowed them to find static resources associated with gathering.

Discussion questions

  1. Why do scientists believe that spatial learning is more developed in males than females? How can we test this?
  2. Can you think of some selective reasons why females perform better in the CEC memory retention, most operant tasks, and spatial memory tasks?


  • classical eyeblink conditioning (CEC) paradigm – a procedure in which an animal is exposed to a noise which is immediately followed by an aversive stimulation to the eyelid which causes the animal to blink; the animal eventually becomes conditioned to blink when it hears the noise, with or without the aversive stimulation (Dalla, 2009)
  • denominated taxon – a learning strategy that involves repetition, animals remember and repeat the same set of movements to reach a goal
  • estrus - The periodic state of sexual excitement in the female of most mammals, excluding humans, that immediately precedes ovulation and during which the female is most receptive to mating; heat.
  • learned helplessness – this occurs when a subject learns that escape from an negative stimulus is impossible; the subject will move less and will have difficulty learning new escape strategies in new tasks
  • local strategies – a learning strategy in which animals learn how to reach a goal through environmental, visual cues that surround the test site (for example the MWM)
  • Morris water-maze (MWM) – most commonly used method to study spatial behavior, subjects are put into a pool of water and learn to find a non-visible platform using spatial cues around the platform; the water is typically made opaque with dye or beads so the animal cannot see the escape platform
  • neurogenesis –the production of new neurons
  • operant conditioning – a task in which an animal must make a response to learn; often they learn how to avoid or escape an aversive stimulus, such as a mild foot shock
  • polygynous - a mating pattern in which a male mates with more than one female in a single breeding season
  • proestrus - The period immediately before estrus in most female mammals, characterized by development of the endometrial and ovarian follicles
  • thigmotaxis – a behavioral response when introduced to a new, possibly dangerous, environment that is best described as “wall-hugging”


  • Aganostaras, SG, Maren, S, DeCola, JP, Lane, NI, Gale, GD, Schlinger, BA, Fanselow, MS. 1998. Testicular hormones do not regulate sexually dimorphic Pavlovian fear conditioning or perforant-path long-term potentiation in adult male rats. Behav Brain Res, 92(1), 1-9.

    Males show greater contextual fear conditioning which depends on the hippocampus and is not regulated by testicular hormones.

  • Beatty, WW. 1979. Gonadal hormones and sex differences in nonreproductive behaviors in rodents: Organizational and activational influences. Hormones&Behav, 12(2): 112-163.

    Review of role of gonadal hormones in sex differences in rodents.

  • Chang, YJ, Yang, CH, Liang, YC, Yeh, CM, Huang, CC, Hsu, KS. 2009. Estrogen modulates sexually dimorphic contextual fear extinction in rats through estrogen receptor beta. Hippocampus, 19(11), 1142-1150.

    Male rats exhibited significantly higher levels of contextual fear memory than female rats. Female rats subjected to conditioning in the proestrus and estrus stage exhibited an enhancement of fear extinction than male rats. Estrogen-mediated enhancement of fear extinction involves the activation of estrogen receptor beta.

  • Cimadevilla, JM, Gonzalez-Pardo, H, Lopez, L, Diaz, F, Cueto, EG, Garcia-Moreno, LM, Arias, JL. 1999. Sex-related differences in spatial learning during the early postnatal development of the rat. Behav Processes, 46(2): 159-171.
  • Cinan, S, et. al. 2007. Memory for object locations: priority effect and sex differences in associative spatial learning. Learn&Motiv, 38(4), 326-341.

    The results indicated that object location memory is affected by priority effects, similar to verbal memory. Experiment 1 demonstrated a female advantage in object location memory.

  • Dalla, C, Edgecomb, C, Whetstone, AS, Shores, TJ. 2008. Females do not express learned helplessness like males do. Neuropsychopharmacology, 33(7), 1559-69.

    Learned helplessness was found to be absent in adult female rats. These sex differences were not dependent on the presence of gonadal sex hormones in adulthood or on testosterone exposure during perinatal development.

  • Dalla, C, Papachristos, EB, Whetstone, AS, Shores, TJ. 2009a. Female rats learn trace memories better than male rats and consequently retain a greater proportion of new neurons in their hippocampi. PNAS, 106(8): 2927-32.
  • Dalla, C, Shores, TJ. 2009b. Sex differences in learning processes of classical and operant conditioning. Phys&Behav, 97(2), 229-238.

    This study reviews sex and age differences that occur in laboratory rodent species. The study focuses on eyeblink conditioning, fear-conditioning, active avoidance and conditioned taste aversion. Sex differences in the brain are responsible for the differences in learning.

  • Denti, A, Epstein, A. 1972. Sex differences in the acquisition of two kinds of avoidance behavior in rats. Phys&Behav, 8(4), 611-615.

    Female rats acquired an active avoidance response more quickly than males, while males were more efficient in passive-avoidance situation.

  • D’Hooge,R, De Deyn, PP. 2001. Applications of the Morris water maze in the study of learning and memory. Brain Res Rev, 36(1): 60-90.
  • Gaulin, SJ. 1992a. Evolution of sex-differences in spatial ability. Yearbook of Phys Anth, 35: 125-151.
  • Gaulin, SJ. 1992b. How and why sex differences evolve, with spatial ability as a paradigm example. In M. Huag (Ed.), The development of sex differences and similarities in behavior (pp. 111- 128). Norwell, MA: Kluwer Academic Publishers.
  • Gaulin, SJ, FitzGerald, RW. 1986. Sex differences in spatial ability: an evolutionary hypothesis and test. Am Naturalist, 127(1): 74-88.
  • Gresack, JE, Schafe, GE, Orr, PT, Frick, KM. 2009. Sex differences in contextual fear conditioning are associated with differential ventral hippocampal extracellular signal-regulated kinase activation. Neurosci, 159(2), 451-467.

    Male rats exhibit more long-term retention of contextual fear conditioning than females.

  • Grimshaw, GM, Sitarenos, G, Finnegan, JK. 1995. Mental rotation at 7 years: relations with prenatal testosterone levels and spatial play experiences. Brain&Cognition, 29: 85-100.
  • Gupta, RR, Sen, S, Diepenhorst, LL, Rudick, CN, Maren, S. 2000. Estrogen modulates sexually dimorphic contextual fear conditioning and hippocampal long-term potentiation (LTP) in rats. Brain Rsrch, 888: 356-65.

    Showed that ovariectomized female rats performed similarly to males in contextual fear conditioning, and both froze more than intact females. Estrogen replacement reduced fear conditioning.

  • Gurzu, C, Artenie, V, Hritcu, L, Ciobica, A. 2008. Prenatal testosterone improves the spatial learning and memory by protein synthesis in different lobes of the brain in the male and female rat. Cent Euro J Biol, 3(1), 39-47.

    This study shows that steroid hormones play an important role in the spatial learning and memory formation by means of protein synthesis in different lobes of the brain.

  • Hajszan, T, Szigeti-Buck, K, Sallam, NL, Bober, J, Parducz, A, MacLusky, NJ, Leranth, C, Duman, RS. 2010. Effects of estradiol on learned helplessness and associated remodeling of hippocampal spine synapses in female rats. Bio Psych, 67(2), 168-174.

    Development of helpless behavior is associated with a severe loss of hippocampal spine synapses, which is reversed by treatment with the antidepressant desipramine.

  • Hetherington, M, Ross, LE. 1963. Effect of sex of subject, sex of experimenter, and reinforcement condition on serial verbal learning. J Exp Psych, 65(6), 572-575.

    Females learned significantly more rapidly than males only under the punishment condition.

  • Hines, M, Fane, BA, Pasterski, VL, Mathews, GA, Conway, GS, Brook, C. 2003. Spatial abilities following prenatal androgen abnormality: targeting and mental rotations performance in individuals with congenital adrenal hyperplasia. Psychoneuroendocrinology, 28(8): 1010-1026.
  • Hyde, JF, Jerussi, TP. 1983. Sexual dimorphism in rats with respect to locomotor activity and circling behavior. Pharmacol Biochem Behav, 18(5): 725-29.

    Female rats are more active than males.

  • Jonasson, Z. 2005. Meta-analysis of sex-differences in rodent models of learning and memory: a review of behavioral and biological data. Neurosci&Behav Rev, 28(8), 811-825.

    Findings indicate reliable male advantages for rats in radial and water maze protocols. Mouse studies exhibited a different model of sex effects.

  • Kanit, L, Koylu, EO, Erdogan, O,&Pogun, S. 2005. Effects of laterality and sex on cognitive strategy in a water maze place learning task and modification by nicotine and nitric oxidase synthase inhibition in rats. Brain Res Bul, 66(3), 189-202.

    Naïve rats of either sex could acquire place learning using response-learning or visual cues in a water maze. When given the choice between response learning versus visually cued learning, response learning was preferred. Drug treatments impaired learning.

  • Lamberty, Y, Gower, AJ. 1988. Investigation into sex-related differences in locomotor activity, place learning, and passive avoidance responding in NMRI mice. Phys and Behav, 44(6), 787-790.
  • Leuner, B, Gould, E, Shors, TJ. 2006. Is there a link between adult neurogenesis and learning? Hippocampus, 16: 216-24.

    Review of hippocampal activity and learning which includes a discussion of the contradiction among literature.

  • Leuner, B, Mendolia-Loffredo, S, Shors TJ. 2004. High levels of estrogen enhance associative memory formation in ovariectomized females. Psychoneuroendocrinology, 29(7):883–90.
  • Levy, LJ, Astur, RS, Frick, KM. 2005. Men and women differ in object memory but not performance of a virtual radial maze. Behav Neurosci, 119(4), 853-862.

    Women significantly outperformed men in recalling the locations and identities of objects.

  • Maren, S, De Oca, B, Fanselow, MS. 1994. Sex differences in hippocampal long-term potentiation (LTP) and Pavlovian fear conditioning in rats: positive correlation between LTP and contextual learning. Brain Research, 661: 25-34.
  • Mendez-Lopez, M, Mendez, M, Lopez, L, Arias, JL. 2009. Spatial working memory learning in young male and female rats: Involvement of different limbic system regions revealed by cytochrome oxidase activity. Neurosci Res, 65(1): 28-34.
  • Neunmeister, A, Wood, S, Bonne, O, Nugent, AC, Luckenbaugh, DA, Young, T, Bain, EE, Charney, DS, Drevets, WC. 2005. Reduced hippocampal volume in unmedicated, remitted patient with major depression versus control subjects. Biol Psychiatry, 57(8): 935-37.
  • Newhouse, P, Newhouse, C, Astur, RS. 2007. Sex differences in visual-spatial learning using a virtual water maze in pre-pubertal children. Behav Brain Res, 183(1), 1-7.

    Males consistently show better performance on the Morris water task than females. This study showed that pre-pubertal boys showed superior performance than similar-aged girls. This shows that the sex differences do not appear to require the effects of sex hormones at puberty.

  • Perrot-Sinal, TS, Kostenuik, MA, Ossenkopp, KP, Kavaliers, M. 1996. Sex differences in performance in the Morris Water Maze and the effects of initial nonstationary hidden platform training. Behav Neurosci, 110(6): 1309-20.

    This study introduces different conditioning paradigms and explains the sex differences observed for each.

  • Pryce, CR, Lehmann, J, Feldon, J. 1999. Effect of sex on fear conditioning is similar for context and discrete CS in Wistar, Lewis, and Fischer rat strains. Pharmacol Biochem&Behav, 64(4):753-759.

    This study explored fear conditioning in rats and discovered that female rats react less frequently to aversive stimuli than male rats. Males are able to learn more successfully from aversive stimuli than females.

  • Puts, DA, Gaulin, SJ, Breedlove, SM. 2007. Sex differences in spatial ability: evolution, hormones, and the brain. In SM Plateck (Ed.), Evolutionary cognitive sciences (pp. 329-379). Cambridge, MA: MIT Press.
  • Randall-Thompson, JF, Riley, AL. 2003. Morphine-induced conditioned taste aversions: assessment of sexual dimorphism. Pharmacol Biochem&Behav, 76(2): 373-81.

    The extinction of conditioned taste aversion showed sex differences for LiCL but not morphine.

  • Ross, J, Roeltgen, D, Zinn, A. 2006. Cognition and the sex chromosomes: studies in Turner syndrome. Hormone Research, 65(1): 47-56.
  • Schulze, I. 1976. Sex differences in the acquisition of appetitively motivated learning in rats. Phys&Behav, 17(1), 19-22.

    No sex difference was observed for learning tasks at 70 days old, but at 100 days old males learned discrimination tasks with chain schedules better than females. In a stimulus discrimination task females suppressed bar-pressing when punished with shock while males only reduced their response rates. Males and females failed to learn active avoidance tasks.

  • Shores, TJ. 2006. Stressful experience and learning across the lifespan. Annu. Rev. Psychol. 57: 55-85.
  • Shores, TJ, Mathew, J, Sisti, HM, Edgecomb, C, Beckoff, S, Dalla C. 2007. Neurogenesis and helplessness are mediated by controllability in males but not in females. Biol Psychiatry, 62: 487-95.

    Controllability of the stress reduced neurogenesis activity and the expression of helplessness behavior in males but not females.

  • Tarpley, JW, Shlifer, IG, Birnbaum, MS, Halladay, LR, Blair, HT. 2009. Bilateral phosphorylation of ERK in the lateral and centrolateral amygdala during unilateral storage of fear memories. Neurosci, 164(3), 908-917.

    This study shows that fear memories are consolidated by the contralateral amygdala. These findings suggest that associative plasticity may occur in both amygdala hemispheres even when only one hemisphere is involved in freezing behavior.

  • Tomilin, MI, Stone, CP. 1933. Sex differences in learning abilities of albino rats. J Comp Psych, 16(2), 207-229.

    The animals learned the problems and were then required to learn a reverse pattern. This study showed no sex differences in the making or breaking of the habits investigated.

  • Vanhaaren, F, Vanhesta, A, Heinsbroek, RPW. 1990. Behavioral-differences between male and female rats – effects of gonadal-hormones on learning and memory. Neurosci&Biobehav Rev, 14(1), 23-33.

    This paper provides an overview of the different experimental procedures, summarizes the most important findings and discusses some of the variables which determine the effects of manipulations in gonadal hormones on learning and memory in male and female rats.

  • VanMeer, P, Raber, J. 2005. Mouse behavioral analysis in systems biology. Biochem J, 389: 593-610.
  • Wood, GE, Shors, TJ. 1998. Stress facilitates classical conditioning in males, but impairs classical conditioning in females through activational effects of ovarian hormones. Proc Natl Acad Sci, 95(7): 4066–71.

About the author

a photo of the author

Vanessa Lippay was born on March 23, 1988 in Shamokin, Pennsylvania. There, she lives with her two parents, her younger brother, and three dogs. She received her undergraduate degree in 2010 from Rice University in Houston, Texas where she studied Ecology and Evolutionary biology and earned a B.A. in Biosciences. She plans on attending graduate school in the fall to obtain a Masters Degree in either Biology or Biomedical Sciences and eventually plans on going to medical school to become a physician. Her interests include film, her dog Bruce, dance, and travel.

Questions & Answers

where we get a research paper on Nano chemistry....?
Maira Reply
nanopartical of organic/inorganic / physical chemistry , pdf / thesis / review
what are the products of Nano chemistry?
Maira Reply
There are lots of products of nano chemistry... Like nano coatings.....carbon fiber.. And lots of others..
Even nanotechnology is pretty much all about chemistry... Its the chemistry on quantum or atomic level
no nanotechnology is also a part of physics and maths it requires angle formulas and some pressure regarding concepts
Preparation and Applications of Nanomaterial for Drug Delivery
Hafiz Reply
Application of nanotechnology in medicine
what is variations in raman spectra for nanomaterials
Jyoti Reply
ya I also want to know the raman spectra
I only see partial conversation and what's the question here!
Crow Reply
what about nanotechnology for water purification
RAW Reply
please someone correct me if I'm wrong but I think one can use nanoparticles, specially silver nanoparticles for water treatment.
yes that's correct
I think
Nasa has use it in the 60's, copper as water purification in the moon travel.
nanocopper obvius
what is the stm
Brian Reply
is there industrial application of fullrenes. What is the method to prepare fullrene on large scale.?
industrial application...? mmm I think on the medical side as drug carrier, but you should go deeper on your research, I may be wrong
How we are making nano material?
what is a peer
What is meant by 'nano scale'?
What is STMs full form?
scanning tunneling microscope
how nano science is used for hydrophobicity
Do u think that Graphene and Fullrene fiber can be used to make Air Plane body structure the lightest and strongest. Rafiq
what is differents between GO and RGO?
what is simplest way to understand the applications of nano robots used to detect the cancer affected cell of human body.? How this robot is carried to required site of body cell.? what will be the carrier material and how can be detected that correct delivery of drug is done Rafiq
analytical skills graphene is prepared to kill any type viruses .
Any one who tell me about Preparation and application of Nanomaterial for drug Delivery
what is Nano technology ?
Bob Reply
write examples of Nano molecule?
The nanotechnology is as new science, to scale nanometric
nanotechnology is the study, desing, synthesis, manipulation and application of materials and functional systems through control of matter at nanoscale
Is there any normative that regulates the use of silver nanoparticles?
Damian Reply
what king of growth are you checking .?
What fields keep nano created devices from performing or assimulating ? Magnetic fields ? Are do they assimilate ?
Stoney Reply
why we need to study biomolecules, molecular biology in nanotechnology?
Adin Reply
yes I'm doing my masters in nanotechnology, we are being studying all these domains as well..
what school?
biomolecules are e building blocks of every organics and inorganic materials.
how did you get the value of 2000N.What calculations are needed to arrive at it
Smarajit Reply
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Source:  OpenStax, Mockingbird tales: readings in animal behavior. OpenStax CNX. Jan 12, 2011 Download for free at http://cnx.org/content/col11211/1.5
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