Arnold Kriegstein, MD, PhD

Founding Director
Eli and Edythe Broad Center of Regeneration Medicine and Stem Cell Research
Department of Neurology

Dr. Kriegstein received BA from Yale University and his MD and PhD degrees from New York University in 1977 where his thesis advisor was Dr. Eric Kandel. He subsequently completed Residency training in Neurology at the Brigham and Women’s Hospital, Children’s Hospital, and Beth Israel Hospital in Boston. He has held academic appointments at Stanford University, Yale University, and Columbia University. In 2004 he joined the Neurology Department at the University of California, San Francisco. He founded the Eli and Edythe Broad Center of Regeneration Medicine and Stem Cell Research at UCSF and served as Director from 2004-2021. Dr. Kriegstein’s own research focuses on the way in which neural stem and progenitor cells in the embryonic brain produce neurons, and ways in which this information can be used for cell based therapies to treat diseases of the nervous system. His lab found that radial glial cells are neuronal stem cells in the developing brain, and also identified a second type of precursor cell produced by radial glial cells that is responsible for generating specific neuronal subtypes. He has more recently focused on human brain development and described a novel progenitor cell type, the outer radial glial cell, that contributes to the huge expansion of the cerebral cortex. He is using single cell transcriptomics and multi-omics as well as spatial transcriptomics to understand how neuronal diversity is produced during brain development. Organoid models are also employed for functional studies as well as insights into disease mechanisms.

Research Summary: 
Neural Stem Cells and Embryonic Cortical Development
Mentorship Development: 

4/29/19    Sharpening your Mentoring Skills (SyMS) with Sharon Milgram (Parnassus)    
9/11/20    Mentoring Across Differences
2/16/21    Three Truths and Three Tries: Facing and Overcoming Critical Social Justice Challenges at the Micro, Mezzo, and Macro Levels    

Websites

Publications: 

Protracted neuronal recruitment in the temporal lobe of young children.

Nature

Nascimento MA, Biagiotti S, Herranz-Pérez V, Santiago S, Bueno R, Ye CJ, Abel TJ, Zhang Z, Rubio-Moll JS, Kriegstein AR, Yang Z, Garcia-Verdugo JM, Huang EJ, Alvarez-Buylla A, Sorrells SF

Single-cell genomics reveals region-specific developmental trajectories underlying neuronal diversity in the human hypothalamus.

Science advances

Herb BR, Glover HJ, Bhaduri A, Colantuoni C, Bale TL, Siletti K, Hodge R, Lein E, Kriegstein AR, Doege CA, Ament SA

Single-cell analysis of prenatal and postnatal human cortical development.

Science (New York, N.Y.)

Velmeshev D, Perez Y, Yan Z, Valencia JE, Castaneda-Castellanos DR, Wang L, Schirmer L, Mayer S, Wick B, Wang S, Nowakowski TJ, Paredes M, Huang EJ, Kriegstein AR

Human pallial MGE-type GABAergic interneuron cell therapy for chronic focal epilepsy.

Cell stem cell

Bershteyn M, Bröer S, Parekh M, Maury Y, Havlicek S, Kriks S, Fuentealba L, Lee S, Zhou R, Subramanyam G, Sezan M, Sevilla ES, Blankenberger W, Spatazza J, Zhou L, Nethercott H, Traver D, Hampel P, Kim H, Watson M, Salter N, Nesterova A, Au W, Kriegstein A, Alvarez-Buylla A, Rubenstein J, Banik G, Bulfone A, Priest C, Nicholas CR

Single cell analysis of dup15q syndrome reveals developmental and postnatal molecular changes in autism.

bioRxiv : the preprint server for biology

Perez Y, Velmeshev D, Wang L, White M, Siebert C, Baltazar J, Dutton NG, Wang S, Haeussler M, Chamberlain S, Kriegstein A

A cross-species proteomic map reveals neoteny of human synapse development.

Nature

Wang L, Pang K, Zhou L, Cebrián-Silla A, González-Granero S, Wang S, Bi Q, White ML, Ho B, Li J, Li T, Perez Y, Huang EJ, Winkler EA, Paredes MF, Kovner R, Sestan N, Pollen AA, Liu P, Li J, Piao X, García-Verdugo JM, Alvarez-Buylla A, Liu Z, Kriegstein AR

LIF signaling regulates outer radial glial to interneuron fate during human cortical development.

Cell stem cell

Andrews MG, Siebert C, Wang L, White ML, Ross J, Morales R, Donnay M, Bamfonga G, Mukhtar T, McKinney AA, Gemenes K, Wang S, Bi Q, Crouch EE, Parikshak N, Panagiotakos G, Huang E, Bhaduri A, Kriegstein AR

Osteopontin drives retinal ganglion cell resiliency in glaucomatous optic neuropathy.

Cell reports

Zhao M, Toma K, Kinde B, Li L, Patel AK, Wu KY, Lum MR, Tan C, Hooper JE, Kriegstein AR, La Torre A, Liao YJ, Welsbie DS, Hu Y, Han Y, Duan X

Astroglial toxicity promotes synaptic degeneration in the thalamocortical circuit in frontotemporal dementia with GRN mutations.

The Journal of clinical investigation

Marsan E, Velmeshev D, Ramsey A, Patel RK, Zhang J, Koontz M, Andrews MG, de Majo M, Mora C, Blumenfeld J, Li AN, Spina S, Grinberg LT, Seeley W, Miller BL, Ullian EM, Krummel MF, Kriegstein A, Huang EJ

Machine learning dissection of human accelerated regions in primate neurodevelopment.

Neuron

Whalen S, Inoue F, Ryu H, Fair T, Markenscoff-Papadimitriou E, Keough K, Kircher M, Martin B, Alvarado B, Elor O, Laboy Cintron D, Williams A, Hassan Samee MA, Thomas S, Krencik R, Ullian EM, Kriegstein A, Rubenstein JL, Shendure J, Pollen AA, Ahituv N, Pollard KS

How mechanisms of stem cell polarity shape the human cerebral cortex.

Nature reviews. Neuroscience

Andrews MG, Subramanian L, Salma J, Kriegstein AR

Ensembles of endothelial and mural cells promote angiogenesis in prenatal human brain.

Cell

Crouch EE, Bhaduri A, Andrews MG, Cebrian-Silla A, Diafos LN, Birrueta JO, Wedderburn-Pugh K, Valenzuela EJ, Bennett NK, Eze UC, Sandoval-Espinosa C, Chen J, Mora C, Ross JM, Howard CE, Gonzalez-Granero S, Lozano JF, Vento M, Haeussler M, Paredes MF, Nakamura K, Garcia-Verdugo JM, Alvarez-Buylla A, Kriegstein AR, Huang EJ

A nomenclature consensus for nervous system organoids and assembloids.

Nature

Pa?ca SP, Arlotta P, Bateup HS, Camp JG, Cappello S, Gage FH, Knoblich JA, Kriegstein AR, Lancaster MA, Ming GL, Muotri AR, Park IH, Reiner O, Song H, Studer L, Temple S, Testa G, Treutlein B, Vaccarino FM

Tropism of SARS-CoV-2 for human cortical astrocytes.

Proceedings of the National Academy of Sciences of the United States of America

Andrews MG, Mukhtar T, Eze UC, Simoneau CR, Ross J, Parikshak N, Wang S, Zhou L, Koontz M, Velmeshev D, Siebert CV, Gemenes KM, Tabata T, Perez Y, Wang L, Mostajo-Radji MA, de Majo M, Donohue KC, Shin D, Salma J, Pollen AA, Nowakowski TJ, Ullian E, Kumar GR, Winkler EA, Crouch EE, Ott M, Kriegstein AR

Zika virus alters centrosome organization to suppress the innate immune response.

EMBO reports

Kodani A, Knopp KA, Di Lullo E, Retallack H, Kriegstein AR, DeRisi JL, Reiter JF

Molecular profiling of stem cell-derived retinal pigment epithelial cell differentiation established for clinical translation.

Stem cell reports

Petrus-Reurer S, Lederer AR, Baqué-Vidal L, Douagi I, Pannagel B, Khven I, Aronsson M, Bartuma H, Wagner M, Wrona A, Efstathopoulos P, Jaberi E, Willenbrock H, Shimizu Y, Villaescusa JC, André H, Sundstr?m E, Bhaduri A, Kriegstein A, Kvanta A, La Manno G, Lanner F

An ACVR1 activating mutation causes neuropathic pain and sensory neuron hyperexcitability in humans.

Pain

Yu X, Ton AN, Niu Z, Morales BM, Chen J, Braz J, Lai MH, Barruet E, Liu H, Cheung K, Ali S, Chan T, Bigay K, Ho J, Nikolli I, Hansberry S, Wentworth K, Kriegstein A, Basbaum A, Hsiao EC

Diversifying stem cell debates: Including Muslim contexts and perspectives.

Stem cell reports

Dajani R, Jiwani B, Nanji A, Zoloth L, Ghaly M, Ilkiliç I, Raya Á, Patrão Neves M, de Melo H, Carvalho AS, Caulfield T, Carter R, Rendas A, Surani A, Rossant J, Kriegstein A, Lalani EN

Nests of dividing neuroblasts sustain interneuron production for the developing human brain.

Science (New York, N.Y.)

Paredes MF, Mora C, Flores-Ramirez Q, Cebrian-Silla A, Del Dosso A, Larimer P, Chen J, Kang G, Gonzalez Granero S, Garcia E, Chu J, Delgado R, Cotter JA, Tang V, Spatazza J, Obernier K, Ferrer Lozano J, Vento M, Scott J, Studholme C, Nowakowski TJ, Kriegstein AR, Oldham MC, Hasenstaub A, Garcia-Verdugo JM, Alvarez-Buylla A, Huang EJ

Challenges of Organoid Research.

Annual review of neuroscience

Andrews MG, Kriegstein AR

Identification of Lipid Heterogeneity and Diversity in the Developing Human Brain.

JACS Au

Bhaduri A, Neumann EK, Kriegstein AR, Sweedler JV

An atlas of cortical arealization identifies dynamic molecular signatures.

Nature

Bhaduri A, Sandoval-Espinosa C, Otero-Garcia M, Oh I, Yin R, Eze UC, Nowakowski TJ, Kriegstein AR

Phospholipid-flippase chaperone CDC50A is required for synapse maintenance by regulating phosphatidylserine exposure.

The EMBO journal

Li T, Yu D, Oak HC, Zhu B, Wang L, Jiang X, Molday RS, Kriegstein A, Piao X

A roadmap for the Human Developmental Cell Atlas.

Nature

Haniffa M, Taylor D, Linnarsson S, Aronow BJ, Bader GD, Barker RA, Camara PG, Camp JG, Chédotal A, Copp A, Etchevers HC, Giacobini P, Göttgens B, Guo G, Hupalowska A, James KR, Kirby E, Kriegstein A, Lundeberg J, Marioni JC, Meyer KB, Niakan KK, Nilsson M, Olabi B, Pe'er D, Regev A, Rood J, Rozenblatt-Rosen O, Satija R, Teichmann SA, Treutlein B, Vento-Tormo R, Webb S, Human Cell Atlas Developmental Biological Network

Distinct nuclear compartment-associated genome architecture in the developing mammalian brain.

Nature neuroscience

Ahanger SH, Delgado RN, Gil E, Cole MA, Zhao J, Hong SJ, Kriegstein AR, Nowakowski TJ, Pollen AA, Lim DA

Human intermediate progenitor diversity during cortical development.

Proceedings of the National Academy of Sciences of the United States of America

Pebworth MP, Ross J, Andrews M, Bhaduri A, Kriegstein AR

Tropism of SARS-CoV-2 for Developing Human Cortical Astrocytes.

bioRxiv : the preprint server for biology

Andrews MG, Mukhtar T, Eze UC, Simoneau CR, Perez Y, Mostajo-Radji MA, Wang S, Velmeshev D, Salma J, Kumar GR, Pollen AA, Crouch EE, Ott M, Kriegstein AR

Mitochondria Control Cortical Cell Fate after Mitosis.

Developmental cell

Wang L, Kriegstein A

Cell-type-specific 3D epigenomes in the developing human cortex.

Nature

Song M, Pebworth MP, Yang X, Abnousi A, Fan C, Wen J, Rosen JD, Choudhary MNK, Cui X, Jones IR, Bergenholtz S, Eze UC, Juric I, Li B, Maliskova L, Lee J, Liu W, Pollen AA, Li Y, Wang T, Hu M, Kriegstein AR, Shen Y

Single-Cell Analyses Identify Brain Mural Cells Expressing CD19 as Potential Off-Tumor Targets for CAR-T Immunotherapies.

Cell

Parker KR, Migliorini D, Perkey E, Yost KE, Bhaduri A, Bagga P, Haris M, Wilson NE, Liu F, Gabunia K, Scholler J, Montine TJ, Bhoj VG, Reddy R, Mohan S, Maillard I, Kriegstein AR, June CH, Chang HY, Posey AD, Satpathy AT

Are Organoids Ready for Prime Time?

Cell stem cell

Bhaduri A, Andrews MG, Kriegstein AR, Nowakowski TJ

Neurotoxic microglia promote TDP-43 proteinopathy in progranulin deficiency.

Nature

Zhang J, Velmeshev D, Hashimoto K, Huang YH, Hofmann JW, Shi X, Chen J, Leidal AM, Dishart JG, Cahill MK, Kelley KW, Liddelow SA, Seeley WW, Miller BL, Walther TC, Farese RV, Taylor JP, Ullian EM, Huang B, Debnath J, Wittmann T, Kriegstein AR, Huang EJ

Origins and Proliferative States of Human Oligodendrocyte Precursor Cells.

Cell

Huang W, Bhaduri A, Velmeshev D, Wang S, Wang L, Rottkamp CA, Alvarez-Buylla A, Rowitch DH, Kriegstein AR

A Chromatin Accessibility Atlas of the Developing Human Telencephalon.

Cell

Markenscoff-Papadimitriou E, Whalen S, Przytycki P, Thomas R, Binyameen F, Nowakowski TJ, Kriegstein AR, Sanders SJ, State MW, Pollard KS, Rubenstein JL

Identification of amygdala-expressed genes associated with autism spectrum disorder.

Molecular autism

Herrero MJ, Velmeshev D, Hernandez-Pineda D, Sethi S, Sorrells S, Banerjee P, Sullivan C, Gupta AR, Kriegstein AR, Corbin JG

SMART-Q:  An Integrative Pipeline Quantifying Cell Type-Specific RNA Transcription.

PloS one

Yang X, Bergenholtz S, Maliskova L, Pebworth MP, Kriegstein AR, Li Y, Shen Y

Medulloblastoma Arises from the Persistence of a Rare and Transient Sox2+ Granule Neuron Precursor.

Cell reports

Selvadurai HJ, Luis E, Desai K, Lan X, Vladoiu MC, Whitley O, Galvin C, Vanner RJ, Lee L, Whetstone H, Kushida M, Nowakowski T, Diamandis P, Hawkins C, Bader G, Kriegstein A, Taylor MD, Dirks PB

Cortical Neural Stem Cell Lineage Progression Is Regulated by Extrinsic Signaling Molecule Sonic Hedgehog.

Cell reports

Zhang Y, Liu G, Guo T, Liang XG, Du H, Yang L, Bhaduri A, Li X, Xu Z, Zhang Z, Li Z, He M, Tsyporin J, Kriegstein AR, Rubenstein JL, Yang Z, Chen B

Cell stress in cortical organoids impairs molecular subtype specification.

Nature

Bhaduri A, Andrews MG, Mancia Leon W, Jung D, Shin D, Allen D, Jung D, Schmunk G, Haeussler M, Salma J, Pollen AA, Nowakowski TJ, Kriegstein AR

Outer Radial Glia-like Cancer Stem Cells Contribute to Heterogeneity of Glioblastoma.

Cell stem cell

Bhaduri A, Di Lullo E, Jung D, Müller S, Crouch EE, Espinosa CS, Ozawa T, Alvarado B, Spatazza J, Cadwell CR, Wilkins G, Velmeshev D, Liu SJ, Malatesta M, Andrews MG, Mostajo-Radji MA, Huang EJ, Nowakowski TJ, Lim DA, Diaz A, Raleigh DR, Kriegstein AR

Neuroglial stem cell-derived inflammatory pseudotumor (n-SCIPT): clinicopathologic characterization of a novel lesion of the lumbosacral spinal cord and nerve roots following intrathecal allogeneic stem cell intervention.

Acta neuropathologica

Sloan EA, Sampognaro PJ, Junn JC, Chin C, Jacques L, Ramachandran PS, DeRisi JL, Wilson MR, Kriegstein AR, Bollen AW, Solomon DA, Margeta M, Engstrom JW

Oligodendrocyte Death in Pelizaeus-Merzbacher Disease Is Rescued by Iron Chelation.

Cell stem cell

Nobuta H, Yang N, Ng YH, Marro SG, Sabeur K, Chavali M, Stockley JH, Killilea DW, Walter PB, Zhao C, Huie P, Goldman SA, Kriegstein AR, Franklin RJM, Rowitch DH, Wernig M

The phenotypes of proliferating glioblastoma cells reside on a single axis of variation.

Cancer discovery

Wang L, Babikir H, Muller S, Yagnik G, Shamardani K, Catalan F, Kohanbash G, Alvarado B, Di Lullo E, Kriegstein A, Shah S, Wadhwa H, Chang SM, Philips JJ, Aghi MK, Diaz AA

New insights into the development of the human cerebral cortex.

Journal of anatomy

Molnár Z, Clowry GJ, Šestan N, Alzu'bi A, Bakken T, Hevner RF, Hüppi PS, Kostovic I, Rakic P, Anton ES, Edwards D, Garcez P, Hoerder-Suabedissen A, Kriegstein A

Neuronal vulnerability and multilineage diversity in multiple sclerosis.

Nature

Schirmer L, Velmeshev D, Holmqvist S, Kaufmann M, Werneburg S, Jung D, Vistnes S, Stockley JH, Young A, Steindel M, Tung B, Goyal N, Bhaduri A, Mayer S, Engler JB, Bayraktar OA, Franklin RJM, Haeussler M, Reynolds R, Schafer DP, Friese MA, Shiow LR, Kriegstein AR, Rowitch DH

Immature excitatory neurons develop during adolescence in the human amygdala.

Nature communications

Sorrells SF, Paredes MF, Velmeshev D, Herranz-Pérez V, Sandoval K, Mayer S, Chang EF, Insausti R, Kriegstein AR, Rubenstein JL, Manuel Garcia-Verdugo J, Huang EJ, Alvarez-Buylla A

Single-cell genomics identifies cell type-specific molecular changes in autism.

Science (New York, N.Y.)

Velmeshev D, Schirmer L, Jung D, Haeussler M, Perez Y, Mayer S, Bhaduri A, Goyal N, Rowitch DH, Kriegstein AR

Multimodal Single-Cell Analysis Reveals Physiological Maturation in the Developing Human Neocortex.

Neuron

Mayer S, Chen J, Velmeshev D, Mayer A, Eze UC, Bhaduri A, Cunha CE, Jung D, Arjun A, Li E, Alvarado B, Wang S, Lovegren N, Gonzales ML, Szpankowski L, Leyrat A, West JAA, Panagiotakos G, Alvarez-Buylla A, Paredes MF, Nowakowski TJ, Pollen AA, Kriegstein AR

Establishing Cerebral Organoids as Models of Human-Specific Brain Evolution.

Cell

Pollen AA, Bhaduri A, Andrews MG, Nowakowski TJ, Meyerson OS, Mostajo-Radji MA, Di Lullo E, Alvarado B, Bedolli M, Dougherty ML, Fiddes IT, Kronenberg ZN, Shuga J, Leyrat AA, West JA, Bershteyn M, Lowe CB, Pavlovic BJ, Salama SR, Haussler D, Eichler EE, Kriegstein AR

Shared and derived features of cellular diversity in the human cerebral cortex.

Current opinion in neurobiology

Miller DJ, Bhaduri A, Sestan N, Kriegstein A

Neuroserpin expression during human brain development and in adult brain revealed by immunohistochemistry and single cell RNA sequencing.

Journal of anatomy

Adorjan I, Tyler T, Bhaduri A, Demharter S, Finszter CK, Bako M, Sebok OM, Nowakowski TJ, Khodosevich K, Møllgård K, Kriegstein AR, Shi L, Hoerder-Suabedissen A, Ansorge O, Molnár Z

Does Adult Neurogenesis Persist in the Human Hippocampus?

Cell stem cell

Paredes MF, Sorrells SF, Cebrian-Silla A, Sandoval K, Qi D, Kelley KW, James D, Mayer S, Chang J, Auguste KI, Chang EF, Gutierrez Martin AJ, Kriegstein AR, Mathern GW, Oldham MC, Huang EJ, Garcia-Verdugo JM, Yang Z, Alvarez-Buylla A

Regulation of cell-type-specific transcriptomes by microRNA networks during human brain development.

Nature neuroscience

Nowakowski TJ, Rani N, Golkaram M, Zhou HR, Alvarado B, Huch K, West JA, Leyrat A, Pollen AA, Kriegstein AR, Petzold LR, Kosik KS

Identification of cell types in a mouse brain single-cell atlas using low sampling coverage.

BMC biology

Bhaduri A, Nowakowski TJ, Pollen AA, Kriegstein AR

Secretagogin is Expressed by Developing Neocortical GABAergic Neurons in Humans but not Mice and Increases Neurite Arbor Size and Complexity.

Cerebral cortex (New York, N.Y. : 1991)

Raju CS, Spatazza J, Stanco A, Larimer P, Sorrells SF, Kelley KW, Nicholas CR, Paredes MF, Lui JH, Hasenstaub AR, Kriegstein AR, Alvarez-Buylla A, Rubenstein JL, Oldham MC

Human-Specific NOTCH2NL Genes Affect Notch Signaling and Cortical Neurogenesis.

Cell

Fiddes IT, Lodewijk GA, Mooring M, Bosworth CM, Ewing AD, Mantalas GL, Novak AM, van den Bout A, Bishara A, Rosenkrantz JL, Lorig-Roach R, Field AR, Haeussler M, Russo L, Bhaduri A, Nowakowski TJ, Pollen AA, Dougherty ML, Nuttle X, Addor MC, Zwolinski S, Katzman S, Kriegstein A, Eichler EE, Salama SR, Jacobs FMJ, Haussler D

An analytical framework for whole-genome sequence association studies and its implications for autism spectrum disorder.

Nature genetics

Werling DM, Brand H, An JY, Stone MR, Zhu L, Glessner JT, Collins RL, Dong S, Layer RM, Markenscoff-Papadimitriou E, Farrell A, Schwartz GB, Wang HZ, Currall BB, Zhao X, Dea J, Duhn C, Erdman CA, Gilson MC, Yadav R, Handsaker RE, Kashin S, Klei L, Mandell JD, Nowakowski TJ, Liu Y, Pochareddy S, Smith L, Walker MF, Waterman MJ, He X, Kriegstein AR, Rubenstein JL, Sestan N, McCarroll SA, Neale BM, Coon H, Willsey AJ, Buxbaum JD, Daly MJ, State MW, Quinlan AR, Marth GT, Roeder K, Devlin B, Talkowski ME, Sanders SJ

Human hippocampal neurogenesis drops sharply in children to undetectable levels in adults.

Nature

Sorrells SF, Paredes MF, Cebrian-Silla A, Sandoval K, Qi D, Kelley KW, James D, Mayer S, Chang J, Auguste KI, Chang EF, Gutierrez AJ, Kriegstein AR, Mathern GW, Oldham MC, Huang EJ, Garcia-Verdugo JM, Yang Z, Alvarez-Buylla A

Single-cell profiling of human gliomas reveals macrophage ontogeny as a basis for regional differences in macrophage activation in the tumor microenvironment.

Genome biology

Müller S, Kohanbash G, Liu SJ, Alvarado B, Carrera D, Bhaduri A, Watchmaker PB, Yagnik G, Di Lullo E, Malatesta M, Amankulor NM, Kriegstein AR, Lim DA, Aghi M, Okada H, Diaz A

Spatiotemporal gene expression trajectories reveal developmental hierarchies of the human cortex.

Science (New York, N.Y.)

Nowakowski TJ, Bhaduri A, Pollen AA, Alvarado B, Mostajo-Radji MA, Di Lullo E, Haeussler M, Sandoval-Espinosa C, Liu SJ, Velmeshev D, Ounadjela JR, Shuga J, Wang X, Lim DA, West JA, Leyrat AA, Kent WJ, Kriegstein AR

The Human Cell Atlas.

eLife

Regev A, Teichmann SA, Lander ES, Amit I, Benoist C, Birney E, Bodenmiller B, Campbell P, Carninci P, Clatworthy M, Clevers H, Deplancke B, Dunham I, Eberwine J, Eils R, Enard W, Farmer A, Fugger L, Göttgens B, Hacohen N, Haniffa M, Hemberg M, Kim S, Klenerman P, Kriegstein A, Lein E, Linnarsson S, Lundberg E, Lundeberg J, Majumder P, Marioni JC, Merad M, Mhlanga M, Nawijn M, Netea M, Nolan G, Pe'er D, Phillipakis A, Ponting CP, Quake S, Reik W, Rozenblatt-Rosen O, Sanes J, Satija R, Schumacher TN, Shalek A, Shapiro E, Sharma P, Shin JW, Stegle O, Stratton M, Stubbington MJT, Theis FJ, Uhlen M, van Oudenaarden A, Wagner A, Watt F, Weissman J, Wold B, Xavier R, Yosef N

The BRAIN Initiative Cell Census Consortium: Lessons Learned toward Generating a Comprehensive Brain Cell Atlas.

Neuron

Ecker JR, Geschwind DH, Kriegstein AR, Ngai J, Osten P, Polioudakis D, Regev A, Sestan N, Wickersham IR, Zeng H

The use of brain organoids to investigate neural development and disease.

Nature reviews. Neuroscience

Di Lullo E, Kriegstein AR

Dynamic behaviour of human neuroepithelial cells in the developing forebrain.

Nature communications

Subramanian L, Bershteyn M, Paredes MF, Kriegstein AR

Human iPSC-Derived Cerebral Organoids Model Cellular Features of Lissencephaly and Reveal Prolonged Mitosis of Outer Radial Glia.

Cell stem cell

Bershteyn M, Nowakowski TJ, Pollen AA, Di Lullo E, Nene A, Wynshaw-Boris A, Kriegstein AR

oRGs and mitotic somal translocation - a role in development and disease.

Current opinion in neurobiology

Ostrem B, Di Lullo E, Kriegstein A

Zika virus cell tropism in the developing human brain and inhibition by azithromycin.

Proceedings of the National Academy of Sciences of the United States of America

Retallack H, Di Lullo E, Arias C, Knopp KA, Laurie MT, Sandoval-Espinosa C, Mancia Leon WR, Krencik R, Ullian EM, Spatazza J, Pollen AA, Mandel-Brehm C, Nowakowski TJ, Kriegstein AR, DeRisi JL

Single-cell sequencing maps gene expression to mutational phylogenies in PDGF- and EGF-driven gliomas.

Molecular systems biology

Müller S, Liu SJ, Di Lullo E, Malatesta M, Pollen AA, Nowakowski TJ, Kohanbash G, Aghi M, Kriegstein AR, Lim DA, Diaz A

Primate Neurons Flex Their Musclin.

Neuron

Pollen AA, Kriegstein AR

Transplanted Human Stem Cell-Derived Interneuron Precursors Mitigate Mouse Bladder Dysfunction and Central Neuropathic Pain after Spinal Cord Injury.

Cell stem cell

Fandel TM, Trivedi A, Nicholas CR, Zhang H, Chen J, Martinez AF, Noble-Haeusslein LJ, Kriegstein AR

Transformation of the Radial Glia Scaffold Demarcates Two Stages of Human Cerebral Cortex Development.

Neuron

Nowakowski TJ, Pollen AA, Sandoval-Espinosa C, Kriegstein AR

A Primate lncRNA Mediates Notch Signaling during Neuronal Development by Sequestering miRNA.

Neuron

Rani N, Nowakowski TJ, Zhou H, Godshalk SE, Lisi V, Kriegstein AR, Kosik KS

SCell: integrated analysis of single-cell RNA-seq data.

Bioinformatics (Oxford, England)

Diaz A, Liu SJ, Sandoval C, Pollen A, Nowakowski TJ, Lim DA, Kriegstein A

Single-cell analysis of long non-coding RNAs in the developing human neocortex.

Genome biology

Liu SJ, Nowakowski TJ, Pollen AA, Lui JH, Horlbeck MA, Attenello FJ, He D, Weissman JS, Kriegstein AR, Diaz AA, Lim DA

Expression Analysis Highlights AXL as a Candidate Zika Virus Entry Receptor in Neural Stem Cells.

Cell stem cell

Nowakowski TJ, Pollen AA, Di Lullo E, Sandoval-Espinosa C, Bershteyn M, Kriegstein AR

Neuronal Migration Dynamics in the Developing Ferret Cortex.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Gertz CC, Kriegstein AR

Molecular identity of human outer radial glia during cortical development.

Cell

Pollen AA, Nowakowski TJ, Chen J, Retallack H, Sandoval-Espinosa C, Nicholas CR, Shuga J, Liu SJ, Oldham MC, Diaz A, Lim DA, Leyrat AA, West JA, Kriegstein AR

Wide Dispersion and Diversity of Clonally Related Inhibitory Interneurons.

Neuron

Harwell CC, Fuentealba LC, Gonzalez-Cerrillo A, Parker PR, Gertz CC, Mazzola E, Garcia MT, Alvarez-Buylla A, Cepko CL, Kriegstein AR

The long noncoding RNA Pnky regulates neuronal differentiation of embryonic and postnatal neural stem cells.

Cell stem cell

Ramos AD, Andersen RE, Liu SJ, Nowakowski TJ, Hong SJ, Gertz C, Salinas RD, Zarabi H, Kriegstein AR, Lim DA

Radial glia in the proliferative ventricular zone of the embryonic and adult turtle, Trachemys scripta elegans.

Neurogenesis (Austin, Tex.)

Clinton BK, Cunningham CL, Kriegstein AR, Noctor SC, Martínez-Cerdeño V

Radial glia require PDGFD-PDGFRß signalling in human but not mouse neocortex.

Nature

Lui JH, Nowakowski TJ, Pollen AA, Javaherian A, Kriegstein AR, Oldham MC

Yoshiki Sasai (1962–2014).

Neuron

Kriegstein AR

The Ink4a/Arf locus is a barrier to direct neuronal transdifferentiation.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Price JD, Park KY, Chen J, Salinas RD, Cho MJ, Kriegstein AR, Lim DA

Yoshiki Sasai (1962–2014).

Cell stem cell

Kriegstein AR

Low-coverage single-cell mRNA sequencing reveals cellular heterogeneity and activated signaling pathways in developing cerebral cortex.

Nature biotechnology

Pollen AA, Nowakowski TJ, Shuga J, Wang X, Leyrat AA, Lui JH, Li N, Szpankowski L, Fowler B, Chen P, Ramalingam N, Sun G, Thu M, Norris M, Lebofsky R, Toppani D, Kemp DW, Wong M, Clerkson B, Jones BN, Wu S, Knutsson L, Alvarado B, Wang J, Weaver LS, May AP, Jones RC, Unger MA, Kriegstein AR, West JA

Control of outer radial glial stem cell mitosis in the human brain.

Cell reports

Ostrem BE, Lui JH, Gertz CC, Kriegstein AR

Interneurons from embryonic development to cell-based therapy.

Science (New York, N.Y.)

Southwell DG, Nicholas CR, Basbaum AI, Stryker MP, Kriegstein AR, Rubenstein JL, Alvarez-Buylla A

Axonal control of the adult neural stem cell niche.

Cell stem cell

Tong CK, Chen J, Cebrián-Silla A, Mirzadeh Z, Obernier K, Guinto CD, Tecott LH, García-Verdugo JM, Kriegstein A, Alvarez-Buylla A

Diverse behaviors of outer radial glia in developing ferret and human cortex.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Gertz CC, Lui JH, LaMonica BE, Wang X, Kriegstein AR

Cell-autonomous correction of ring chromosomes in human induced pluripotent stem cells.

Nature

Bershteyn M, Hayashi Y, Desachy G, Hsiao EC, Sami S, Tsang KM, Weiss LA, Kriegstein AR, Yamanaka S, Wynshaw-Boris A

The dynamics of neuronal migration.

Advances in experimental medicine and biology

Wu Q, Liu J, Fang A, Li R, Bai Y, Kriegstein AR, Wang X

Non-epithelial stem cells and cortical interneuron production in the human ganglionic eminences.

Nature neuroscience

Hansen DV, Lui JH, Flandin P, Yoshikawa K, Rubenstein JL, Alvarez-Buylla A, Kriegstein AR

Functional maturation of hPSC-derived forebrain interneurons requires an extended timeline and mimics human neural development.

Cell stem cell

Nicholas CR, Chen J, Tang Y, Southwell DG, Chalmers N, Vogt D, Arnold CM, Chen YJ, Stanley EG, Elefanty AG, Sasai Y, Alvarez-Buylla A, Rubenstein JL, Kriegstein AR

Use of "MGE enhancers" for labeling and selection of embryonic stem cell-derived medial ganglionic eminence (MGE) progenitors and neurons.

PloS one

Chen YJ, Vogt D, Wang Y, Visel A, Silberberg SN, Nicholas CR, Danjo T, Pollack JL, Pennacchio LA, Anderson S, Sasai Y, Baraban SC, Kriegstein AR, Alvarez-Buylla A, Rubenstein JL

A high-resolution enhancer atlas of the developing telencephalon.

Cell

Visel A, Taher L, Girgis H, May D, Golonzhka O, Hoch RV, McKinsey GL, Pattabiraman K, Silberberg SN, Blow MJ, Hansen DV, Nord AS, Akiyama JA, Holt A, Hosseini R, Phouanenavong S, Plajzer-Frick I, Shoukry M, Afzal V, Kaplan T, Kriegstein AR, Rubin EM, Ovcharenko I, Pennacchio LA, Rubenstein JL

Mitotic spindle orientation predicts outer radial glial cell generation in human neocortex.

Nature communications

LaMonica BE, Lui JH, Hansen DV, Kriegstein AR

Regulation of microtubule stability and organization by mammalian Par3 in specifying neuronal polarity.

Developmental cell

Chen S, Chen J, Shi H, Wei M, Castaneda-Castellanos DR, Bultje RS, Pei X, Kriegstein AR, Zhang M, Shi SH

SnapShot: cortical development.

Cell

Woodworth MB, Custo Greig L, Kriegstein AR, Macklis JD

Stem cells in the land of the rising sun: ISSCR 2012.

Cell stem cell

Plath K, Srivastava D, Alvarez-Buylla A, Tanaka EM, Kriegstein AR

Forebrain GABAergic neuron precursors integrate into adult spinal cord and reduce injury-induced neuropathic pain.

Neuron

Bráz JM, Sharif-Naeini R, Vogt D, Kriegstein A, Alvarez-Buylla A, Rubenstein JL, Basbaum AI

OSVZ progenitors in the human cortex: an updated perspective on neurodevelopmental disease.

Current opinion in neurobiology

LaMonica BE, Lui JH, Wang X, Kriegstein AR

Sonic hedgehog expression in corticofugal projection neurons directs cortical microcircuit formation.

Neuron

Harwell CC, Parker PR, Gee SM, Okada A, McConnell SK, Kreitzer AC, Kriegstein AR

Persistent sonic hedgehog signaling in adult brain determines neural stem cell positional identity.

Neuron

Ihrie RA, Shah JK, Harwell CC, Levine JH, Guinto CD, Lezameta M, Kriegstein AR, Alvarez-Buylla A

Development and evolution of the human neocortex.

Cell

Lui JH, Hansen DV, Kriegstein AR

A new subtype of progenitor cell in the mouse embryonic neocortex.

Nature neuroscience

Wang X, Tsai JW, LaMonica B, Kriegstein AR

Kinesin 3 and cytoplasmic dynein mediate interkinetic nuclear migration in neural stem cells.

Nature neuroscience

Tsai JW, Lian WN, Kemal S, Kriegstein AR, Vallee RB

Connexin 43 mediates the tangential to radial migratory switch in ventrally derived cortical interneurons.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Elias LA, Turmaine M, Parnavelas JG, Kriegstein AR

Neurogenic radial glia in the outer subventricular zone of human neocortex.

Nature

Hansen DV, Lui JH, Parker PR, Kriegstein AR

Embryonic MGE precursor cells grafted into adult rat striatum integrate and ameliorate motor symptoms in 6-OHDA-lesioned rats.

Cell stem cell

Martínez-Cerdeño V, Noctor SC, Espinosa A, Ariza J, Parker P, Orasji S, Daadi MM, Bankiewicz K, Alvarez-Buylla A, Kriegstein AR

Research ethics. NIH guidelines for stem cell research and gamete donors.

Science (New York, N.Y.)

Lo B, Parham L, Cedars M, Fisher S, Gates E, Giudice L, Halme DG, Hershon W, Kriegstein A, Rao R, Roberts C, Wagner R

Cloning mice and men: prohibiting the use of iPS cells for human reproductive cloning.

Cell stem cell

Lo B, Parham L, Alvarez-Buylla A, Cedars M, Conklin B, Fisher S, Gates E, Giudice L, Halme DG, Hershon W, Kriegstein A, Kwok PY, Wagner R

Mammalian Par3 regulates progenitor cell asymmetric division via notch signaling in the developing neocortex.

Neuron

Bultje RS, Castaneda-Castellanos DR, Jan LY, Jan YN, Kriegstein AR, Shi SH

A stem cell niche for intermediate progenitor cells of the embryonic cortex.

Cerebral cortex (New York, N.Y. : 1991)

Javaherian A, Kriegstein A

Commentary: the prospect of cell-based therapy for epilepsy.

Neurotherapeutics : the journal of the American Society for Experimental NeuroTherapeutics

Kriegstein AR, Pitkänen A

Importing human pluripotent stem cell lines derived at another institution: tailoring review to ethical concerns.

Cell stem cell

Lo B, Parham L, Broom C, Cedars M, Gates E, Giudice L, Halme DG, Hershon W, Kriegstein A, Kwok PY, Oberman M, Roberts C, Wagner R

Defining the role of GABA in cortical development.

The Journal of physiology

Wang DD, Kriegstein AR

The glial nature of embryonic and adult neural stem cells.

Annual review of neuroscience

Kriegstein A, Alvarez-Buylla A

Differential trafficking of AMPA and NMDA receptors by SAP102 and PSD-95 underlies synapse development.

Proceedings of the National Academy of Sciences of the United States of America

Elias GM, Elias LA, Apostolides PF, Kriegstein AR, Nicoll RA

GABA regulates stem cell proliferation before nervous system formation.

Epilepsy currents

Wang DD, Kriegstein AR, Ben-Ari Y

GABA regulates excitatory synapse formation in the neocortex via NMDA receptor activation.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Wang DD, Kriegstein AR

Manipulating midbrain stem cell self-renewal.

Cell stem cell

LoTurco JJ, Kriegstein AR

Distinct behaviors of neural stem and progenitor cells underlie cortical neurogenesis.

The Journal of comparative neurology

Noctor SC, Martínez-Cerdeño V, Kriegstein AR

Gap junctions: multifaceted regulators of embryonic cortical development.

Trends in neurosciences

Elias LA, Kriegstein AR

Clinical trials in stem cell transplantation: guidelines for scientific and ethical review.

Clinical trials (London, England)

Lo B, Kriegstein A, Grady D

In utero intraventricular injection and electroporation of E15 mouse embryos.

Journal of visualized experiments : JoVE

Walantus W, Castaneda D, Elias L, Kriegstein A

In utero intraventricular injection and electroporation of E16 rat embryos.

Journal of visualized experiments : JoVE

Walantus W, Elias L, Kriegstein A

Organotypic slice culture of E18 rat brains.

Journal of visualized experiments : JoVE

Elias L, Kriegstein A

Contribution of intermediate progenitor cells to cortical histogenesis.

Archives of neurology

Noctor SC, Martínez-Cerdeño V, Kriegstein AR

Neural stem and progenitor cells in cortical development.

Novartis Foundation symposium

Noctor SC, Martinez-Cerdeño V, Kriegstein AR

Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex.

The European journal of neuroscience

Martínez-Cerdeño V, Noctor SC, Kriegstein AR

Patterns of neural stem and progenitor cell division may underlie evolutionary cortical expansion.

Nature reviews. Neuroscience

Kriegstein A, Noctor S, Martínez-Cerdeño V

Characterization of mice with targeted deletion of glycine receptor alpha 2.

Molecular and cellular biology

Young-Pearse TL, Ivic L, Kriegstein AR, Cepko CL

Progress in corticogenesis.

Cerebral cortex (New York, N.Y. : 1991)

Kriegstein A, Parnavelas JG

The role of intermediate progenitor cells in the evolutionary expansion of the cerebral cortex.

Cerebral cortex (New York, N.Y. : 1991)

Martínez-Cerdeño V, Noctor SC, Kriegstein AR

Blind patch clamp recordings in embryonic and adult mammalian brain slices.

Nature protocols

Castañeda-Castellanos DR, Flint AC, Kriegstein AR

The temporal and spatial origins of cortical interneurons predict their physiological subtype.

Neuron

Butt SJ, Fuccillo M, Nery S, Noctor S, Kriegstein A, Corbin JG, Fishell G

A new era in the ethics of human embryonic stem cell research.

Stem cells (Dayton, Ohio)

Lo B, Zettler P, Cedars MI, Gates E, Kriegstein AR, Oberman M, Reijo Pera R, Wagner RM, Wuerth MT, Wolf LE, Yamamoto KR

LIS1 RNA interference blocks neural stem cell division, morphogenesis, and motility at multiple stages.

The Journal of cell biology

Tsai JW, Chen Y, Kriegstein AR, Vallee RB

GABA puts the brake on stem cells.

Nature neuroscience

Kriegstein AR

Cortical development: new concepts.

Neuron

Fishell G, Kriegstein A

Controlling neuron number: does Numb do the math?

Nature neuroscience

Castañeda-Castellanos DR, Kriegstein AR

Patterns of neuronal migration in the embryonic cortex.

Trends in neurosciences

Kriegstein AR, Noctor SC

Cortical neurons arise in symmetric and asymmetric division zones and migrate through specific phases.

Nature neuroscience

Noctor SC, Martínez-Cerdeño V, Ivic L, Kriegstein AR

Terpene trilactones from Ginkgo biloba are antagonists of cortical glycine and GABA(A) receptors.

The Journal of biological chemistry

Ivic L, Sands TT, Fishkin N, Nakanishi K, Kriegstein AR, Strømgaard K

Changing concepts of cortical development.

Cerebral cortex (New York, N.Y. : 1991)

Kriegstein A, Parnavelas JG

Neurogenic radial glial cells in reptile, rodent and human: from mitosis to migration.

Cerebral cortex (New York, N.Y. : 1991)

Weissman T, Noctor SC, Clinton BK, Honig LS, Kriegstein AR

Neurons from radial glia: the consequences of asymmetric inheritance.

Current opinion in neurobiology

Fishell G, Kriegstein AR

Developmental neurotransmitters?

Neuron

Owens DF, Kriegstein AR

Is there more to GABA than synaptic inhibition?

Nature reviews. Neuroscience

Owens DF, Kriegstein AR

Dividing precursor cells of the embryonic cortical ventricular zone have morphological and molecular characteristics of radial glia.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Noctor SC, Flint AC, Weissman TA, Wong WS, Clinton BK, Kriegstein AR

GABA may act as a self-limiting trophic factor at developing synapses.

Science's STKE : signal transduction knowledge environment

Kriegstein AR, Owens DF

Neurons derived from radial glial cells establish radial units in neocortex.

Nature

Noctor SC, Flint AC, Weissman TA, Dammerman RS, Kriegstein AR

Maturation of channels and receptors: consequences for excitability.

International review of neurobiology

Owens DF, Kriegstein AR

Extrinsic GABAergic innervation of developing neocortical layer 1 in organotypic slice co-cultures.

The Journal of comparative neurology

Dammerman RS, Noctor SC, Kriegstein AR

An excitatory GABAergic plexus in developing neocortical layer 1.

Journal of neurophysiology

Dammerman RS, Flint AC, Noctor S, Kriegstein AR

Calcium dynamics of neocortical ventricular zone cells.

Developmental neuroscience

Owens DF, Flint AC, Dammerman RS, Kriegstein AR

Leukoencephalopathy and raised brain lactate from heroin vapor inhalation ("chasing the dragon")

Neurology

Kriegstein AR, Shungu DC, Millar WS, Armitage BA, Brust JC, Chillrud S, Goldman J, Lynch T

Endogenous activation of metabotropic glutamate receptors in neocortical development causes neuronal calcium oscillations.

Proceedings of the National Academy of Sciences of the United States of America

Flint AC, Dammerman RS, Kriegstein AR

Ontogeny of channels, transmitters and epileptogenesis.

Advances in neurology

Kriegstein AR, Owens DF, Avoli M

Patterns of intracellular calcium fluctuation in precursor cells of the neocortical ventricular zone.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Owens DF, Kriegstein AR

Postnatal development of low [Mg2+] oscillations in neocortex.

Journal of neurophysiology

Flint AC, Maisch US, Kriegstein AR

Cell coupling and uncoupling in the ventricular zone of developing neocortex.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Bittman K, Owens DF, Kriegstein AR, LoTurco JJ

Mechanisms underlying neuronal migration disorders and epilepsy.

Current opinion in neurology

Flint AC, Kriegstein AR

NR2A subunit expression shortens NMDA receptor synaptic currents in developing neocortex.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Flint AC, Maisch US, Weishaupt JH, Kriegstein AR, Monyer H

Heroin inhalation and progressive spongiform leukoencephalopathy.

The New England journal of medicine

Kriegstein AR, Armitage BA, Kim PY

Excitatory GABA responses in embryonic and neonatal cortical slices demonstrated by gramicidin perforated-patch recordings and calcium imaging.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Owens DF, Boyce LH, Davis MB, Kriegstein AR

Cortical neurogenesis and its disorders.

Current opinion in neurology

Kriegstein AR

GABA and glutamate depolarize cortical progenitor cells and inhibit DNA synthesis.

Neuron

LoTurco JJ, Owens DF, Heath MJ, Davis MB, Kriegstein AR

Spontaneous action potential activity and synaptic currents in the embryonic turtle cerebral cortex.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Blanton MG, Kriegstein AR

Morphological differentiation of distinct neuronal classes in embryonic turtle cerebral cortex.

The Journal of comparative neurology

Blanton MG, Kriegstein AR

Clusters of coupled neuroblasts in embryonic neocortex.

Science (New York, N.Y.)

Lo Turco JJ, Kriegstein AR

Initial expression and endogenous activation of NMDA channels in early neocortical development.

The Journal of neuroscience : the official journal of the Society for Neuroscience

LoTurco JJ, Blanton MG, Kriegstein AR

Endogenous neurotransmitter activates N-methyl-D-aspartate receptors on differentiating neurons in embryonic cortex.

Proceedings of the National Academy of Sciences of the United States of America

Blanton MG, Lo Turco JJ, Kriegstein AR

Whole cell recording from neurons in slices of reptilian and mammalian cerebral cortex.

Journal of neuroscience methods

Blanton MG, Lo Turco JJ, Kriegstein AR

Development of GABA responsiveness in embryonic turtle cortical neurons.

Neuroscience letters

Shen JM, Huguenard JR, Kriegstein AR

Synaptic responses of cortical pyramidal neurons to light stimulation in the isolated turtle visual system.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Kriegstein AR

Glutamate neurotoxicity in cortical cell culture.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Choi DW, Maulucci-Gedde M, Kriegstein AR

Cellular physiology of the turtle visual cortex: distinctive properties of pyramidal and stellate neurons.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Connors BW, Kriegstein AR

Cellular physiology of the turtle visual cortex: synaptic properties and intrinsic circuitry.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Kriegstein AR, Connors BW

Morphological classification of rat cortical neurons in cell culture.

The Journal of neuroscience : the official journal of the Society for Neuroscience

Kriegstein AR, Dichter MA

Stages in the post-hatching development of Aplysia californica.

The Journal of experimental zoology

Kriegstein AR

Development of the nervous system of Aplysia californica.

Proceedings of the National Academy of Sciences of the United States of America

Kriegstein AR

Metamorphosis of Aplysia californica in laboratory culture.

Proceedings of the National Academy of Sciences of the United States of America

Kriegstein AR, Castellucci V, Kandel ER