“…Even though spatial navigation seems effortless at the behavioral level, it is a multimodal activity that draws upon a multitude of cognitive and neural resources (Moffat, 2009 , 2016 ; Wolbers and Hegarty, 2010 ; Wolbers, 2015 ; Zhong and Kozhevnikov, 2016 ; Lester et al, 2017 ). Numerous behavioral studies of spatial navigation in the cognitive aging literature have identified age-related declines or deficits in navigation strategies (Moffat and Resnick, 2002 ; Bohbot et al, 2012 ; Harris et al, 2012 ; Wiener et al, 2013 ; Harris and Wolbers, 2014 ; Colombo et al, 2017 ; Zhong et al, 2017 ), associative learning/memory (Head and Isom, 2010 ; Liu et al, 2011 ; Wiener et al, 2012 , 2013 ; Zhong and Moffat, 2016 ; Allison and Head, 2017 ; O’Malley et al, 2018 ), and working memory (Mahmood et al, 2009 ; Taillade et al, 2013a , b , 2016 ; Ariel and Moffat, 2018 ). Complementary neuroimaging studies that investigated age-related declines in spatial navigation performance and memory have largely linked them to age-related reduction in the volume or activation of the hippocampus (e.g., Driscoll et al, 2003 , 2005 ; Astur et al, 2006 ; Moffat et al, 2006 , 2007 ; Antonova et al, 2009 ; Yuan et al, 2014 ; Daugherty et al, 2015 , 2016 ), a region that has long been proposed as the neural basis of a “cognitive map” (O’Keefe and Dostrovsky, 1971 ; O’Keefe and Nadel, 1978 ).…”