Jinn, J., E. Conner and Jacobs, L.F. (2020). How ambient environment influences olfactory orientation in search and rescue dogs. Chemical Senses.

Under natural conditions, an animal orienting to an air-borne odor plume must contend with the
shifting influence of meteorological variables, such as air temperature, humidity and wind speed,
on the location and the detectability of the plume. Despite their importance, the natural statistics
of such variables are difficult to reproduce in the laboratory and hence few studies have
investigated strategies of olfactory orientation by mobile animals under different meteorological
conditions. Using trained search and rescue dogs, we quantified the olfactory orientation
behaviors of dogs searching for a trail (aged 1-3 hours) of a hidden human subject in a natural
landscape, under a range of meteorological conditions. Dogs were highly successful in locating
the human target hidden 800 m from the start location (93% success). Humidity and air
temperature had a significant effect on search strategy: as air conditions became cooler and more
humid, dogs searched significantly closer to the original trail. Dogs also modified their speed and
head position according to their search location distance from the original trail. When close to
the trail, dogs searched with their head up and ran quickly but when their search took them
farther from the trail, they were more likely to search with their nose to the ground, moving more
slowly. To our knowledge, this is the first study of a mammalian species responding to localized
shifts in ambient conditions and it lays the foundation for future studies of olfactory orientation,
and the development of a highly tractable mammalian species for such research.

Accepted manuscript:


Jacobs, Lucia F. (2019) The navigational nose: a new hypothesis for the function of the human external pyramid. Journal of Experimental Biology

There is currently only one hypothesis for the evolution of this structure, based on its role in respiration, yet the nose also has a sensory function. This paper proposes that the human external nose may also have evolved to enhance stereo olfaction during spatial orientation.

 PDF: 2019 JEB Jacobs

Delgado, M. M, and L. F Jacobs. How squirrels protect their caches: location,
conspicuousness during caching, and proximity to kin influence cache lifespan. BioRxiv 6
(August 16, 2019): 338–45.

By genotyping a subpopulation of Berkeley squirrels, Mikel showed that that scatter hoarding fox squirrels placed caches closer to caches made by their close relatives. This suggests that foraging decisions are influenced by kin selection, as the loss of caches to pilfering relatives would be offset by inclusive fitness. Her study also identified key strategic choices squirrels make while caching and how this influences how long the cache remains hidden before it is retrieved or pilfered.

PDF: Delgado & Jacobs 2019


Delgado, M.M. & Jacobs, L.F., 2017. Caching for where and what: evidence for a mnemonic strategy in a scatter-hoarder. Royal Society Open Science, 4(9), pp.170958–6.

PDF: R. Soc. open sci. 2017 Delgado

Squirrels organize their caches spatially by the species of nut. Such ‘spatial chunking’ increases the accuracy of memory recall in other species and probably reduces the cognitive load of building and managing an optimal cache map.



Delgado, M.M. and Jacobs, L.F. (2016). Inaccessibility of reinforcement increases persistence and signaling behavior in the fox squirrel, Sciurus niger. Journal of Comparative Psychology. 130(2): 128-137.

PDF: J Comp Psychol 2016 Delgado

Is frustration adaptive?  See the experiment here in this short film by KQED  http://”Watch Those Frustrated Squirrels”


Jacobs, L. F., Arter, J., Cook, A., & Sulloway, F. J. (2015). Olfactory orientation and navigation in humans. PLoS ONE, 10(6), e0129387.

PDF: PLoS ONE 2015 Jacobs

Given only one minute to sniff a location, humans will be able to return to that location, using only their sense of smell. This first study of olfactory mapping by humans suggests that olfaction is an under appreciated navigational device in our species spatial toolkit.


Jacobs, L. F., & Menzel, R. (2014). Navigation outside of the box: what the lab can learn from the field and what the field can learn from the lab. Movement Ecology, 2(1), 1–22.

PDF: Movement Ecology 2014 Jacobs

Delgado, M. M., Nicholas, M., Petrie, D. J., & Jacobs, L. F. (2014). Fox squirrels match food assessment and cache effort to value and scarcity. PLoS ONE, 9(3), e92892.

PDF: PLoS ONE 2014 Delgado et al

Evan L. MacLean, Brian A. Hare, Charles L. Nunn, Elsa Addessi, Federica Amici, Rindy C. Anderson, Filippo Aureli, Joseph M. Baker, Amanda E. Bania, Allison M. Barnard, Neeltje J. Boogert, Elizabeth M. Brannon, Emily E. Bray, Joel Bray, Lauren J. N. Brent, Judith M. Burkart, Josep Call, Jessica F. Cantlon, Lucy G. Cheke, Nicola S. Clayton, Mikel M. Delgado, Louis J. DiVincenti, Kazuo Fujita, Chihiro Hiramatsu, Lucia F. Jacobs, Kerry E. Jordan, Jennifer R. Laude, Kristin L. Leimgruber, Emily J. E. Messer, Antonio C. de A. Moura, Ljerka Ostojić, Alejandra Picard, Michael L. Platt, Joshua M. Plotnik, Friederike Range, Simon M. Reader, Rachna B. Reddy, Aaron A. Sandel, Laurie R. Santos, Katrin Schumann, Amanda M. Seed, Kendra B. Sewall, Rachael C. Shaw, Katie E Slocombe, Yanjie Su, Ayaka Takimoto, Jingzhi Tan, Ruoting Tao, Carel P. van Schaik, Zsófia Virányi, Elisabetta Visalberghi, Jordan C. Wade, Arii Watanabe, Jane Widness, Thomas R. Zentall, Yini Zhao. (2014) The evolution of self-control. Proceedings of the National Academy of Sciences, 111(20), E2140–8. doi:10.1073/pnas.1323533111.

PDF: Proc Natl Acad Sci USA 2014 MacLean


Cook, A., Arter, J., & Jacobs, L. F. (2013). My owner, right or wrong: the effect of familiarity on the domestic dog’s behavior in a food-choice task. Animal Cognition, 17(2), 461–470.

PDF: Anim Cogn 2013 Cook

Waismeyer, A. S., & Jacobs, L. F. (2013). The emergence of flexible spatial strategies in young children. Developmental Psychology, 49(2), 232–242.

PDF: Dev Psych 2013 Waismeyer


Jacobs, L. F. (2012). From chemotaxis to the cognitive map: the function of olfaction. Proc Natl Acad Sci, 109, 10693–10700.

PDF: Proc Natl Acad Sci USA 2012 Jacobs

Bettis, T., & Jacobs, L. F. (2012). Sex differences in object recognition are modulated by object similarity. Behavioural Brain Research, 233(2), 288–292.

PDF: Behav Brain Res 2012 Bettis

Chai, X. J., & Jacobs, L. F. (2012). Digit ratio predicts sense of direction in women. PLoS ONE, 7(2), e32816.

PDF: PLoS ONE 2012 Chai

Kleschevnikov, A. M., Belichenko, P. V., Faizi, M., Jacobs, L. F., Htun, K., Shamloo, M., & Mobley, W. C. (2012). Deficits in cognition and synaptic plasticity in a mouse model of Down Syndrome ameliorated by GABAB receptor antagonists. Journal of Neuroscience, 32(27), 9217–9227.

PDF: J Neurosci 2012 Kleschevnikov

MacLean, E.L., Matthews, L.J., Hare, B.A., Nunn, C.L., Anderson, R.C., Aureli, F., Brannon, E.M., Call, J., Drea, C.M., Emery, N.J., Haun, D.B., Herrmann, E., Jacobs, L.F., Platt, M.L., Rosati, A.G., Sandel, A.A., Schroepfer, K.K., Seed, A.M., Tan, J., van Schaik, C.P., and Wobber, V. (2012). How does cognition evolve? Phylogenetic comparative psychology. Animal Cognition 15, 223-238.

PDF: Anim Cogn 2012 Maclean

Delgado, M.M. (2012).

Jacobs, L. F. (2012). “Do Squirrels Remember Where They Bury Their Nuts?” “What do Honeybees Say When they Dance?” “What Does “Chickadee” Mean to a Chickadee?”

Essays in: M. Lamothe, J. Rothman, & J. Volvovski (Eds.), The Where, the Why and the How: 75 Artists Illustrate the Wondrous Mysteries of Science. San Francisco: Chronicle Books.


Wiener, J., Shettleworth, S., Bingman, V.P., Cheng, K., Healy, S., Jacobs, L.F., Jeffery, K.J., Mallot, H.A., Menzel, R., and Newcombe, N.S. (2011). Animal Navigation – A Synthesis. In Animal Thinking: Contemporary Issues in Comparative Cognition, R. Menzel, and J. Fischer, eds. (Cambridge, MA, MIT Press).

PDF: 2011 Wiener

Waisman, A. S., Lucas, C. G., Griffiths, T. L., & Jacobs, L. F. (2011). A Bayesian model of navigation in squirrels. Presented at the Proc 33rd Annual Conf Cog Sci Society.

PDF: CogSci 2011 Waisman


Chai, X. J., & Jacobs, L. F. (2010). Effects of cue types on sex differences in human spatial memory. Behavioural Brain Research, 208(2), 336–342.

PDF: Behav Brain Res 2010 Chai

Chai, X. J., Ofen, N., Jacobs, L. F., & Gabrieli, J. D. E. (2010). Scene complexity: influence on perception, memory, and development in the medial temporal lobe. Frontiers in Human Neuroscience, 4, 21–10.

PDF: Front Hum Neurosci 2010 Chai


Chai, X. J., & Jacobs, L. F. (2009). Sex differences in directional cue use in a virtual landscape. Behavioral Neuroscience, 123(2), 276–283.

PDF: Behav Neurosci 2009 Chai

Bettis, T. J., & Jacobs, L. F. (2009). Sex-specific strategies in spatial orientation in C57BL/6J mice. Behavioural Processes.

PDF:  Behav Proc 2009 Bettis

Preston, S. D., & Jacobs, L. F. (2009). Mechanisms of cache decision making in fox squirrels (Sciurus niger). Journal of Mammalogy, 90(4), 787–795.

PDF: Journal of Mammalogy 2009 Preston

Jacobs, L. F. (2009). The role of social selection in the evolution of hippocampal specialization. In L. Tommasi, M. A. Peterson, & L. Nadel (Eds.), Cognitive Biology: Evolutionary and Developmental Perspectives on Mind. Cambridge, MA: MIT Press.

PDF pending


Waisman, A. S., & Jacobs, L. F. (2008). Flexibility of cue use in the fox squirrel (Sciurus niger). Animal Cognition, 11(4), 625–636.

PDF: Anim Cogn 2008 Waisman


Barkley, C., & Jacobs, L. F. (2007). Sex and species differences in spatial memory in food-storing kangaroo rats. Animal Behaviour, 73(2), 321–329.

PDF: Anim Behav 2007 Barkley

Gibbs, S. E. B., Lea, S. E. G., & Jacobs, L. F. (2007). Flexible use of spatial cues in the southern flying squirrel (Glaucomys volans). Animal Cognition, 10(2), 203–209.

PDF: Anim Cogn 2007 Gibbs

Adams-Hunt, M.M., and Jacobs, L.F. (2007). Learning to forage. In: Foraging: Behavior and Ecology, D.W. Stephens, J.S. Brown, and R.C. Ydenberg, eds. (Chicago, IL, University of Chicago), p. 576.

PDF: 2007 Adams-Hunt


Jacobs, L. F. (2006). From movement to transitivity: the role of hippocampal parallel maps in configural learning. Reviews in the Neurosciences, 17(1-2), 99–109.

PDF:  Reviews in the Neurosciences 2006 Jacobs


Hermes, G. L., Jacobs, L. F., & McClintock, M. (2005). The sectored foraging field: A novel design to quantify spatial strategies, learning, memory, and emotion. Neurobiology of Learning and Memory, 84(1), 69–73.

PDF:  Neurobiol Learn Mem 2005 Hermes

Preston, S. D., & Jacobs, L. F. (2005). Cache decision making: the effects of competition on cache decisions in Merriam’s kangaroo rat (Dipodomys merriami). Journal of Comparative Psychology, 119(2), 187.

PDF:   J Comp Psych 2005 Preston


Jacobs, L. F. (2003). The evolution of the cognitive map. Brain, Behavior and Evolution, 62(2), 128–139.

PDF:Brain Behav Evol 2003 Jacobs

Jacobs, L. F., & Schenk, F. (2003). Unpacking the cognitive map: the parallel map theory of hippocampal function. Psychological Review, 110(2), 285–315.

PDF:  Psych Rev 2003 Jacobs


Preston, S. D., & Jacobs, L. F. (2001). Conspecific pilferage but not presence affects Merriam’s kangaroo rat cache strategy. Behavioral Ecology, 12(5), 517–523.

PDF:  Behav Ecol 2001 Preston


Jacobs, L.F. (2000). Sexual differentiation and cognitive function. In: Gender and Society: the 1995 Herbert Spencer Lectures, S. Iversen, ed. (Oxford, UK, Oxford University Press).

PDF:  Spencer Lecture 2000 Jacobs

Lavenex, P., Steele, M. A., & Jacobs, L. F. (2000). Sex differences, but no seasonal variations in the hippocampus of food‐caching squirrels: A stereological study. Journal of Comparative Neurology, 425(1), 152–166.

PDF:  J Comp Neurol 2000 Lavenex

Lavenex, P., Steele, M. A., & Jacobs, L. F. (2000). The seasonal pattern of cell proliferation and neuron number in the dentate gyrus of wild adult eastern grey squirrels. European Journal of Neuroscience, 12(2), 643–648.

PDF:  Eur J Neurosci 2000 Lavenex


Jacobs, L. F., & Shiflett, M. W. (1999). Spatial orientation on a vertical maze in free-ranging fox squirrels ( Sciurus niger). Journal of Comparative Psychology, 113(2), 116–127.

PDF:  J Comp Psych 1999 Jacobs

Barkley, C., & Jacobs, L. F. (1998). Visual environment and delay affect cache retrieval accuracy in a food-storing rodent. Learning & Behavior, 26(4), 439–447.

PDF:  Learn Behav 1998 Barkley

Lavenex, P., Shiflett, M. W., Lee, R. K., & Jacobs, L. F. (1998). Spatial versus nonspatial relational learning in free-ranging fox squirrels (Sciurus niger). Journal of Comparative Psychology, 112(2), 127–136.

PDF:   J Comp Psych 1998 Lavenex

Feener, D. H., Jacobs, L. F., & Schmidt, J. O. (1996). Specialized parasitoid attracted to a pheromone of ants. Animal Behaviour, 51(1), 61–66.

PDF:   Anim Behav 1996 Feener

Jacobs, L. F. (1996). Sexual selection and the brain. Trends Ecol Evol, 11(2), 82–86.

PDF:  Trends Ecol Evol 1996 Jacobs

Jacobs, L. F. (1996). The economy of winter: phenotypic plasticity in behavior and brain structure. The Biological Bulletin, 191(1), 92–100.

PDF:  Biol Bull 1996 Jacobs

Jacobs, L. F. (1995). The ecology of spatial cognition:  adaptive patterns of hippocampal size and space use in wild rodents. In E. Alleva, A. Fasolo, H.-P. Lipp, & L. Nadel (Eds.), (pp. 301–322). Dordrecht: Behavioural Brain Research in Naturalistic and Semi-Naturalistic Setting.

PDF:Jacobs 1995 eco of spatnav

Jacobs, L. F., & Spencer, W. D. (1994). Natural space-use patterns and hippocampal size in kangaroo rats. Brain, Behavior and Evolution, 44(3), 125–132.

PDF:   Brain Behav Evol 1994 Jacobs

Daly, M., Behrends, P., Wilson, M. I., & Jacobs, L. F. (1992). Behavioural modulation of predation risk: moonlight avoidance and crepuscular compensation in a nocturnal desert rodent, Dipodomys merriami. Animal Behaviour, 44(1), 1–9.

PDF:   Anim Behav 1992 Daly

Daly, M., Jacobs, L. F., Wilson, M. I., & Behrends, P. (1992). Scatter hoarding by kangaroo rats (Dipodomys merriami) and pilferage from their caches. Behavioral Ecology, 3(2), 102–111.

PDF:  Behav Ecol 1992 Daly

Daly, M., Wilson, M. I., Behrends, P., & Jacobs, L. F. (1992). Sexually differentiated effects of radio transmitters on predation risk and behaviour in kangaroo rats (Dipodomys merriami). Canadian Journal of Zoology, 70(9), 1851–1855.

PDF:  Can J Zool 1992 Daly

Jacobs, L. F. (1992). Memory for cache locations in Merriam’s kangaroo rats. Animal Behaviour, 43(4), 585–593.

PDF: Anim Behav 1992 Jacobs

Jacobs, L. F. (1992). The effect of handling time on the decision to cache by grey squirrels. Animal Behaviour, 43, 533–524.

PDF: Anim Behav 1992 Jacobs

Sherry, D. F., Jacobs, L. F., & Gaulin, S. J. C. (1992). Spatial memory and adaptive specialization of the hippocampus. Trends in Neurosciences, 15(8), 298–303.

PDF: Trends Neurosci 1992 Sherry

Jacobs, L. F., & Liman, E. R. (1991). Grey squirrels remember the locations of buried nuts. Animal Behaviour, 41(1), 103–110.

PDF: Animal Behaviour 1991 Jacobs

Jacobs, L. F., Gaulin, S. J. C., Sherry, D. F., & Hoffman, G. E. (1990). Evolution of spatial cognition: sex-specific patterns of spatial behavior predict hippocampal size. Proc Natl Acad Sci USA, 87(16), 6349–6352.

PDF: Proc Natl Acad Sci USA 1990 Jacobs

Daly, M., Wilson, M., Behrends, P., & Jacobs, L. F. (1990). Characteristics of kangaroo rats, Dipodomys merriami, associated with differential predation risk. Animal Behaviour, 40(2), 380–389.

PDF: Anim Behav 1990 Daly

Sengelaub, D. R., Jacobs, L. F., & Finlay, B. L. (1985). Regional differences in normally occurring cell death in the developing hamster lateral geniculate nuclei. Neuroscience Letters, 55(2), 103–108.

PDFNeurosci Lett 1985 Sengelaub