Stark Lab

Neurobiology & Behavior



Here is a list of our laboratory publications. If you do not have access to any of the PDF reprints linked here, please feel free to email me and I will send you a copy via e-mail.

2020 and in press

Stark SM, Frithsen A, & Stark CEL (in press). Age-related alterations in functional connectivity along the longitudinal axis of the hippocampus and its subfields. Hippocampus

Ventakesh A, Stark SM, Stark CE, Bennett IJ (2020). Age- and Memory- Related Differences in Hippocampal Gray Matter Integrity Are Better Captured by NODDI Compared to Single-Tensor Diffusion Imaging. Neurobiology of Aging, doi:10.1016/j.neurobiolaging.2020.08.004

Papp KV, Rentz DM, Maruff P, Sun CK, Raman R, Donohue MC, Schembri A, Stark C, Yassa MA, Wessels AM, Yaari R, Holdridge KC, Aisen PS, Sperling RA on behaof of the A4 Study Team (2020) The Computerized Cognitive Composite (C3) in an Alzheimer’s Disease Secondary Prevention TrialThe Journal of Prevention of Alzheimer’s Disease.

Kolarik, Stark SM, & Stark CEL (2020) Enriching hippocampal memory function in older adults through real-world exploration. Frontiers in Aging Neuroscience, doi: 10.3389/fnagi.2020.00158

Clemenson GD, Stark SM, Rutledge SM, & Stark CEL (2020) Enriching hippocampal memory function in older adults through video games. Behavioral and Brain Research, doi: 10.1016/j.bbr.2020.112667

Klippenstein JL, Stark SM, Stark CEL, & Bennett IJ (2020) Neural substrates of mnemonic discrimination: A whole-brain fMRI investigation. Brain & Behavior, doi: 10.1002/brb3.1560

Radhakrishnana H, Stark SM, & Stark CEL (2020) Microstructural alterations in hippocampal subfields mediate age-related memory decline in humans. Frontiers in Aging Neuroscience,


Levine L, Lench H, Stark C, Carlson S, Carpenter Z, Stark S, & Frithsen A (2019). Predicted and remembered emotion: Tomorrow’s vividness trumps yesterday’s accuracy. Memory, 28(1): 128-140.

Olsen RK, Carr VA, … Stark CEL … Wisse LM on behalf of the Hippocampal Subfields Group (2019). Progress update from the hippocampal subfields group. Alzheimer’s & Dementia: Diagnosis, Assessment & Disease Monitoring, 11, 439-499.

Stark SM, Kirwan CB, & Stark CEL (2019). Mnemonic Similarity Task: A Tool for Assessing Hippocampal Integrity. Trends in Cognitive Science,

Acharya MM, Baulch JE, Klein PM, Baddour AAD, Apodaca LA, Kramár EA, Alikhani L, Garcia C, Angulo MC, Batra RS, Fallgren CM,  Borak TB, Stark CEL, Wood MA, Britten RA, Soltesz I, & Limoli CL (2019). New concerns for neurocognitive function during deep space exposures to chronic, low dose rate, neutron radiation. eNeuro August 5, 2019.

Clemenson GD, Henningfield CM, & Stark C (2019). Improving hippocampal memory through the experience of a rich Minecraft environment. Frontiers in Behavioral Neuroscience. doi: 10.3389/fnbeh.2019.00057

Stark SM, & Stark CEL (2019). Excitatory/inhibitory imbalance in anterior lateral occipital cortex can impair hippocampal mnemonic discrimination. Neuron, 101: 360-362.


Frithsen A, Stark SM, & Stark CEL (2018). Response bias, recollection, and familiarity in individuals with Highly Superior Autobiographical Memory (HSAM). Memory,

Stark SM, Frithsen A, Mattfeld AT, & Stark CEL (2018) Modulation of associative learning in the hippocampal-striatal circuit based on item-set similarity. Cortex, 109: 60-73.

Wais PE, Montgomery O, Stark CEL, & Gazzaley A (2018) Evidence of a causal role for mid-ventrolateral prefrontal cortex based functional networks in retrieving high-fidelity memory. Scientific Reports, 8(1): 14877. doi: 10.1038/s41598-018-33164-w.

Bennett IJ, Stark SM, & Stark CEL (2018) Recognition memory dysfunction relates to hippocampal subfield volume: A study of cognitively normal and mildly impaired older adults. Journal of Gerontology: Psychological Sciences,


Stark SM & Stark CEL (2017) Age-related deficits in the mnemonic similarity task for objects and scenes. Behavioral Brain Research,

Stark SM, Reagh ZM, Yassa MA, & Stark CEL (2017) What’s in a context? Cautions, limitations, and potential paths forward. Neuroscience Letters,

Huffman DJ & Stark CEL (2017). The influence of low-level stimulus features on the representation of contexts, items, and their mnemonic associations. Neuroimage,

Nikolova S, Stark SM, & Stark CEL (2017) 3T hippocampal glutamate-glutamine complex reflects verbal memory decline in aging. Neurobiology of Aging, DOI:

Wais PE, Jahanikia S, Steiner D, Stark CEL, Gazzaley A (2017) Retrieval of high-fidelity memory arises from distributed cortical networks. NeuroImage, 149: 178-189.

Huffman DJ & Stark CEL (2017) Age-related impairment on a forced-choice version of the Mnemonic Similarity Task. Behavioral Neuroscience, 131(1): 55-67.


Mesina L, Wilber AA, Clark BJ, Dube S, Demecha AJ, Stark CEL, & McNaughton BL (2016) A methodological pipeline for serial-section imaging and tissue realignment for whole-brain functional and connectivity assessment. Journal of Neuroscience Methods, 266: 151-160.

LePort AKR, Stark SM, McGaugh JL, & Stark CEL (2016).  A cognitive assessment of Highly Superior Autobiographical Memory.  Memory, doi: 10.1080/09658211.2016.1160126

Saletin JM, Goldstein-Piekarski AN, Greer SM, Stark S, Stark CEL, & Walker MP (2016). Human Hippocampal Structure: A Novel Biomarker Predicting Mnemonic Vulnerability to, and Recovery from, Sleep Deprivation. Journal of Neuroscience, 36(8): 2355-2363, doi: 10.1523/JNEUROSCI.3466-15.2016

LePort AKR, Stark SM, McGaugh JL, & Stark CEL (2016) Highly Superior Autobiographical Memory: Quality and quantity of retention over time. Frontiers in Psychology, 6:2017. doi:10.3389/fpsyg.2015.02017.

Wisse LEM, Daugherty, AM…..Stark, CEL…..Hippocampal Subfields Group (2016) A harmonized segmentation protocol for hippocampal and parahippocampal subregions: Why do we need one and what are the key goals? Hippocampus, DOI 10.1002/hipo.22671.


Clemenson GD & Stark CEL (2015). Virtual environmental enrichment through video games improves hippocampal-associated memory. Journal of Neuroscience, 35(49): 16116-16125; doi: 10.1523/JNEUROSCI.2580-15.2015.

Bennett IJ, & Stark CEL (2015). Mnemonic Discrimination Relates to Perforant Path Integrity: An Ultra-High Resolution Diffusion Tensor Imaging Study. Neurobiology of Learning and Memory, S1074-7427(15)00120-3 do: 10.1016/j.nlm.2015.06.014.

Stark SM, Stevenson R, Wu C, Rutledge S, & Stark CEL (2015). Stability of age-related deficits in the mnemonic similarity task across task variations. Behavioral Neuroscience, 129(3): 257-268.  [Supplemental data PDF]

Allen T, Morris AM, Stark SM, Fortin NJ, & Stark CEL (2015). Memory for sequences of events impaired in typical aging. Learning and Memory, DOI:10.1101/lm.036301.114

Mattfeld AM & Stark CEL (2015). Functional contributions and interactions between the human hippocampus and subregions of the striatum during arbitrary associative learning and memory. Hippocampus, 25: 900-911.

Yushkevich PA and the Hipppocampal Subfields Group (2015). Quantitative comparison of 21 protocols for labeling hippocampal subfields and parahippocampal subregions in in vivo MRI: Towards a harmonized segmentation protocol. Neuroimage, DOI: 10.1016/j.neuroimage.2015.01.004.


Das T, Ivlela EE, Wagner AD, Stark CEL, & Tammnga CA (2014). Loss of pattern separation performance in schizophrenia suggests dentate gyrus dysfunction. Schizophrenia Research, DOI: 10.1016/j.schres.2014.05.006

Huffman DJ & Stark CEL (2014) Multivariate pattern analysis of the human medial temporal lobe revealed representationally categorical cortex and representationally agnostic hippocampus. Hippocampus, DOI: 10.1002/hipo.22321

Allen TA, Morris, AM, Mattfeld AT, Stark CEL, & Fortin NJ (2014) A sequence of events model of episodic memory shows parallels in rats and humans. Hippocampus, DOI: 10.1002/hippo.22301.

Bennett IJ, Huffman DJ & Stark CEL (2014) Limbic tract integrity contributes to pattern separation performance across the lifespan. Cerebral Cortex, DOI: 10.1093/cercor/bhu093.

Patihis L, Frenda SJ, LePort AKR, Petersen N, Nichols RM, Stark CEL, McGaugh JL, & Loftus EF (2013). False memories in highly superior autobiographical memory individuals. Proceedings of the National Academy of Sciences, 110(52): 20947-20952.

Azab M, Stark SM, & Stark CEL (2013). A domain agnostic role for the dentate gyrus in pattern separation: Evidence from a temporal and spatial associative learning task using high-resolution fMRI. Hippocampus, 24(3):293-302.

Lacy JW & Stark CEL (2013). The neuroscience of memory: implications for the courtroom. Nature Reviews Neuroscience, 14: 1-10.  Public pre-print available

Stark SM, Yassa MA, Lacy JW, & Stark CEL (2013). A task to assess behavioral pattern separation (BPS) in humans: Data from healthy aging and mild cognitive impairment. Neuropsychologia, doi: 10.1016/j.neuropsychologia.2012.12.014

Kirwan CB, Hartshorn A, Stark SM, Goodrich-Hunsaker NJ, Hopkins RO, & Stark CEL (2012). Pattern separation deficits following damage to the hippocampus. Neuropsychologia, 50(10), 2408-2414.

LePort, AKR, Mattfeld, AT, Dickinson-Anson, H, Fallon JH, Stark, CEL, Kruggel, FR, Cahill, L, & McGaugh, JL (2012) A behavioral and neuroanatomical investigation of Highly Superior Autobiographical Memory (HSAM). Neurobiology of Learning and Memory, 98(1), 78-92.

Lacy, JW, & Stark, CEL (2012). Intrinsic functional connectivity of the medial temporal lobe suggest a distinction between adjacent MTL cortices and hippocampus. Hippocampus. DOI: 10.1002/hipo.22047

Stark CEL (2012) It’s time to fill gaps left by simple dissociations. Cognitive Neuroscience, 3, 215-216.

Yassa MA, Hazlett RL, Stark CEL, & Hoehn-Saric R (2012) Functional MRI of the amygdala and bed nucleus of the stria terminals during prolonged stress in generalized anxiety disorder. Journal of Psychiatric Research, 46(8):1045-52

Hargreaves EL, Mattfeld MA, Stark CEL, & Suzuki WA (2012) Equivalent fMRI and LFP signals underlying new associative learning in the human and macaque monkey medial temporal lobe. Neuron, 74(4), 743-752.

Bakker A., Krauss G, Albert MA, Speck CL, Jones LR, Stark CEL, Yassa MA, Bassett SS, Shelton AL, & Gallagher M (2012) Levetiracetam treatment attenuates excess hippocampal activity with cognitive benefit in patients with amnestic mild cognitive impairment. Neuron, 74(3): 467-474.

Segal SK, Stark SM, Kattan D, Stark CEL, & Yassa MA (2012) Norepinephrine-mediated emotional arousal facilitates subsequent pattern separation. Neurobiology of Learning and Memory, 97(4): 465-469.

Mattfeld AT, Gluck MA, & Stark CEL (2011) Functional specialization within the striatum along both the dorsal/ventral and anterior/posterior axes during associative learning via reward and punishment. Learning and Memory, 18:703-711.

Yassa MA & Stark CEL (2011) Pattern separation in the hippocampus. Trends in Neuroscience, 34(10): 515-525.

Yassa MA, Mattfeld, AT, Stark SM, & Stark CEL (2011) Age-related memory deficits linked to circuit-specific disruptions in the hippocampus. Proceedings of the National Academy of Sciences USA, 108(21), 8873-8878.

Lacy JW, Yassa MA, Stark SM, Muftuler LT, & Stark CEL (2011) Distinct pattern separation related transfer functions in human CA1 and CA3/DG revealed using high-resolution fMRI and mnemonic similarity. Learning and Memory, 18: 15-18.

Yassa MA, Wu J, Stark SM, Albert MS, Gallagher M, & Stark CEL (2011) Pattern separation deficits associated with increased hippocampal CA3 and dentate gyrus activity in nondemented older adults. Hippocampus, 21(9):968-979.

Stark CEL, Okado Y, & Loftus, EF (2010) Imaging the reconstruction of true and false memories using sensory reactivation and the misinformation paradigm. Learning and Memory, 17: 485-488.

Mattfeld AM & Stark CEL (2010) Striatal and medial temporal lobe functional interactions during visuomotor associative learning. Cerebral Cortex, 21(3):647-658.

Yassa MA, Muftuler LT, & Stark CEL (2010) Ultrahigh-resolution microstructural diffusion tensor imaging (msDTI) elucidates perforant path degradation in aged humans in vivo. Proceedings of the National Academy of Sciences, 107(28), 12687-12691.

Stark SM, Yassa MA, & Stark CEL (2010) Individual differences in spatial pattern separation performance associated with healthy aging in humans. Learning and Memory, 17(6), 284-288.

Yassa MA, Stark SM, Bakker A, Albert MS, Gallagher M, & Stark CEL (2010) High-resolution functional MRI of hippocampal CA3 and dentate gyrus in patients with amnestic mild cognitive impairment. NeuroImage, 51(3), 1242-1252.

Gallagher M, Bakker A, Yassa MA, & Stark CEL (2010) Bridging neurocognitive aging and disease modification: targeting functional mechanisms of impairment. Current Alzheimer’s Research, 7(3), 1567-2050.

Yassa MA & Stark CEL (2009). A quantitative evaluation of cross-participant alignment techniques for MRI studies of the medial temporal lobe. Neuroimage 44(2) 319-327.

Yassa MA & Stark CEL (2008). Multiple Recognition-related signals in the medial temporal lobe. Hippocampus 18(9): 945-54.

Bakker A, Kirwan CB, Miller MI, & Stark CEL (2008). Pattern separation in the human hippocampal CA3 and dentate gyrus. Science, 319(5870), 1640-1642.

Stark SM, Gordon B & Stark CEL (2008). Does priming hinder subsequent recognition performance?, Memory 16(2), 157-173.

Stark S, Gordon B, & Stark CEL (2008). A case study of amnesia: Exploring a paradigm for new semantic learning and generalization. Brain Injury, 22(3), 283-292.

Kirwan CB, & Stark CEL (2007). Overcoming interference: An fMRI investigation of pattern separation in the medial temporal lobe. Learning and Memory, 14, 625-633.

Kirwan CB, Jones, C, Miller MI, & Stark CEL (2007). High-resolution fMRI investigation of the medial temporal lobe. Human Brain Mapping, 28. 959-966.

Gold JJ, Smith CN, Bayley PJ, Shrager Y, Brewer JB, Stark CEL, Hopkins RO, & Squire LR. (2006). Item memory, source memory, and the medial temporal lobe: Concordant findings from fMRI and memory-impaired patients. Proceedings of the National Academy of Sciences, 103, 9351-9356.

Miller MI, Beg MF, Ceritoglu C, & Stark CEL (2005). Increasing the power of functional maps of the medial temporal lobe using large deformation metric mapping. Proceedings of the National Academy of Sciences of the USA, 102, 9685-9690.

Law JR, Flanery MA, Wirth S, Yanike M, Smith AC, Frank LM, Suzuki WA, Brown EN, & Stark CEL (2005). fMRI activity during the gradual acquisition and expression of paired-associate memory. The Journal of Neuroscience, 25, 5720-5729.

Okado Y, & Stark CEL (2005). Neural activity during encoding predicts false memories created by misinformation. Learning and Memory, 12, 3-11. (Download stimuli)

Stark CEL, Stark SM, & Gordon BG (2005). New semantic learning and generalization in an amnesic patient. Neuropsychology, 19, 139-151.

Levy DA, Stark CEL, & Squire LR (2004). Intact conceptual priming in the absence of declarative memory. Psychological Science, 15, 680-686.

Kirwan BC, & Stark CEL (2004). Medial temporal lobe activation during encoding and retrieval of novel face-name pairs. Hippocampus, 14, 910-930.

Squire LR, Stark CEL, & Clark RE (2004). The medial temporal lobe. Annual Review of Neuroscience, 27: 279-306.

Okado Y & Stark CEL (2003). Neural processing associated with true and false memory retrieval. Cognitive, Affective, and Behavioral Neuroscience, 3, 323-334.

Stark CEL & Okado Y (2003). Making memories without trying: Medial temporal lobe activity associated with incidental memory formation during recognition. The Journal of Neuroscience, 23, 6748-6753.

Stark CEL & Squire LR (2003). Hippocampal damage equally impairs memory for single items and memory for conjunctions. Hippocampus, 13, 281-292.

Stark CEL, Bayley PJ, & Squire LR (2002). Recognition memory for single items and for associations is similarly impaired following damage to the hippocampal region. Learning and Memory, 9, 238-242.

Stark CEL & Squire LR (2001). When zero is not zero: The problem of ambiguous baseline conditions in fMRI. Proceedings of the National Academy of Sciences of the USA, 98, 12760-12766.

Stark CEL & Squire LR (2001). Simple and associative recognition memory in the hippocampal region. Learning and Memory, 8, 190-197.

Stark CEL & Squire LR (2000). Recognition memory and familiarity judgments in severe amnesia: No evidence for a contribution of repetition priming. Behavioral Neuroscience, 114, 459-467.

Stark CEL & Squire LR (2000). fMRI activity in the medial temporal lobe during recognition memory as a function of study-test interval. Hippocampus, 10, 329-337.

Stark CEL & Squire LR (2000). Functional magnetic resonance imaging (fMRI) activity in the hippocampal region during recognition memory. The Journal of Neuroscience, 20(20), 7776-7781.

Stark CEL & Squire LR (2000). Intact visual perceptual discrimination in humans in the absence of perirhinal cortex. Learning and Memory, 7(5): 273-278.

Manns J, Stark CEL & Squire LR (2000). The visual paired-comparison task as a measure of declarative memory. Proceedings of the National Academy of Sciences, 97(22):12375-12379.

Stark CEL & McClelland JL (2000). Repetition priming of words, pseudowords, and nonwords. Journal of Experimental Psychology: Learning, Memory, and Cognition, 26(4), 945-972.

Reber PJ, Stark CEL & Squire LR (1998). Cortical areas supporting category learning identified using functional MRI. Proceedings of the National Academy of Sciences of the USA, 95, 747-750.

Reber PJ, Stark CEL, & Squire LR (1998). Contrasting cortical activity associated with category memory and recognition memory. Learning and Memory, 5, 420-428.

Book Chapters

Stark SM & Stark CEL (2015). Introduction to Memory. In: Neurobiology of Language. Eds: G Hickok & S Small. Elsevier. 841-854.

Stark CEL (2010). The Neural Basis of Declarative Memory. In Encyclopedia of Behavioral Neuroscience. G. Koob, M. LeMoal, R. Thompson (Eds). Elsevier.

Squire LR & Stark CEL (2008). Memory Systems. In Topics in Integrative Neuroscience: From Cells to Cognition. J Pomerantz & M. Crair (Eds.) Oxford University Press

Stark CEL (2007). The Functional Role of the Human Hippocampus. In The Hippocampus. P. Anderson, R. Morris, D. Amaral, T. Bliss, & J. O’Keefe (Eds). Oxford University Press

Stark CEL & Squire LR (2002). Functional magnetic resonance imaging (fMRI). In McGraw-Hill 2002 Yearbook of Science and Technology. McGraw-Hill, New York.

Video and Radio appearances

Highly Superior Autobiographical Memory: What Would it Mean to Remember Every Day of your Life? LA Sunday Assembly, March 13, 2016

3D Videogames and Memory on “Science Friday” with Ira Flatow. National Public Radio, December 25, 2015

Is That What Happened?, LA Sunday Assembly, March 8 2015

The Neuroscience of True and False Memories. European Molecular Biology Laboratory Forum on Science and Society. Heidelberg, Germany. March 24, 2014.

Forget with the Program: Skeptic Check with Gary Niederhoff, Big Picture Science Radio Show, May 15, 2012.

The Nature of Memory on “Midmorning” with Kerri Miller.  Minnesota Public Radio (MPR) January 12, 2012, 9:45AM CST.

The Cognitive Neuroscience of True and False Memories. Fred Kavli Public Symposium: The Brain on Trial: Neuroscience and the Law. 41st Annual Meeting for the Society for Neuroscience, Washington D.C., November 12, 2011.

Making False Memories on “Talk of the Nation: Science Friday” with Ira Flatow.  National Public Radio, February 4, 2005, 3:40PM EST

The Science of Memory on “All things Considered”, with Jon Hamilton.  National Public Radio, February 9, 2005, 4:40PM EST