Goldstein Lab
some reprints
(pdf files and links to freely available articles)

Some of our published research articles and reviews: 

Papell LD, Coke AN, de Jesus BN, Harry CJ, Zhang P, Goldstein B (2026). Chromosomes remain individualized through interphase in embryos of the tardigrade Hypsibius exemplaris. Molecular Biology of the Cell 37(1):br2, doi.org/10.1091/mbc.E25-07-0356​.

Zhang P, Medwig-Kinney T, Breiner EA, Perez JM, Song AN, Goldstein B (2025). Cell signaling facilitates apical constriction by basolaterally recruiting Arp2/3 via Rac and WAVE. Journal of Cell Biology 224(5):e202409133.

Clark-Hachtel CM, Hibshman JD, De Buysscher T, Stair ER, Hicks LM, Goldstein B (2024). The tardigrade Hypsibius exemplaris dramatically upregulates DNA repair pathway genes in response to ionizing radiation. Current Biology 34:1819-1830.

Harry CJ*, Hibshman JD*, Damatac A, Davidson PL, Estermann MA, Flores-Flores M, Holmes CM, Lázaro J, Legere EA, Leyhr J, Thendral SB, Vincent BA, Goldstein B (2024). Protocol for fluorescent live-cell staining of tardigrades. STAR Protocols 5:103232.

Hibshman JD, Clark-Hachtel CM, Bloom KS, Goldstein B (2024). A bacterial expression cloning screen reveals single-stranded DNA-binding proteins as potent desicco-protectants. Cell Reports 43:114956. doi 10.1016/j.celrep.2024.114956.

Zhang P*, Medwig-Kinney TN*, Goldstein B (2023). Architecture of the cortical actomyosin network driving apical constriction in C. elegans. Journal of Cell Biology 222(9):e202302102. doi.org/10.1083/jcb.202302102

Slabodnick MM*, Tintori SC*, Prakash M, Zhang P, Higgins CD, Chen AH, Cupp TD, Wong T, Bowie E, Jug F, Goldstein B (2023). Zyxin contributes to coupling between cell junctions and contractile actomyosin networks during apical constriction. PLoS Genetics 19(3): e1010319. doi.org/10.1371/journal.pgen.1010319

Heikes KL, Game M, Smith FW, Goldstein B (2023). The Embryonic Origin of Primordial Germ Cells in the Tardigrade Hypsibius exemplaris. Developmental Biology 497:42-58.

Hibshman JD, Carra S, Goldstein B (2023). Tardigrade small heat shock proteins can limit desiccation-induced protein aggregation. Communications Biology, 6:121 doi:10.1038/s42003-023-04512-y.

Goldstein B (2022). Tardigrades and their emergence as model organisms, in B Goldstein and M Srivastava, Emerging Model Systems in Developmental Biology (pp 173-198), Academic Press.

Goldstein B (2022) Tardigrades. Nature Methods 19:904-905.

Pani AM, Gibney TV, Medwig-Kinney TN, Matus DQ, Goldstein B (2022). A new toolkit to visualize and perturb endogenous LIN-12/Notch signaling in C. elegans. microPublication Biology https://doi.org/10.17912/micropub.biology.000603.

Goldstein B, Avasthi P (2021). A Guide to Setting Up and Managing a Lab at a Research-Intensive Institution. BMC Proceedings 15(Suppl 2):8. https://doi.org/10.1186/s12919-021-00214-7

Hibshman JD, Goldstein B (2021). LEA motifs promote desiccation tolerance in vivo. BMC Biology, 19:263. https://doi.org/10.1186/s12915-021-01176-0.

Tintori SC, Golden P, Goldstein B (2020). Differential Expression Gene Explorer (DrEdGE): A tool for generating interactive online data visualizations for exploration of quantitative transcript abundance datasets. Submitted. Preprint: bioRxiv doi 10.1101/618439

Goldstein B, Nance J (2020). C. elegans gastrulation: a model for understanding how cells polarize, change shape, and journey toward the center of an embryo. Genetics 214:265-277.

Hibshman JD, Clegg JS, Goldstein B (2020). Mechanisms of Desiccation Tolerance: Themes and Variations in Brine Shrimp, Roundworms, and Tardigrades. Frontiers in Physiology 11:592016.

Pani AM and B Goldstein (2018). Direct visualization of a native Wnt in vivo reveals that a long-range Wnt gradient forms by extracellular dispersal.  eLife 7:e38325.

Smith FW, Cumming M, Goldstein B (2018). Analyses of nervous system patterning genes in the tardigrade Hypsibius exemplaris illuminate the evolution of panarthropod brains. EvoDevo 9:19.

Heppert JK, Pani AM, Roberts AM, Dickinson DJ, and B Goldstein (2018). A CRISPR tagging-based screen reveals localized players in Wnt-directed asymmetric cell division. Genetics 208:1147-1164.

Goldstein B (2018). The emergence of the tardigrade Hypsibius exemplaris as a model system. CSH Protocols, 10:859-866.

McGreevy KM, Heikes KL, Kult S, Tharp ME, and B Goldstein (2018). Fluorescent cell staining methods for living Hypsibius exemplaris embryos. CSH Protocols, 10:878-884.

Heikes KL and B Goldstein (2018). Live imaging of tardigrade embryonic development by differential interference contrast microscopy. CSH Protocols 10:974-877.

Fielmich LE, Schmidt R, Dickinson DJ, Goldstein B, Akhmanova A, and S Van den Heuvel (2018). Optogenetic dissection of mitotic spindle positioning in vivo. eLife 7:e38199.

Dickinson DJ, Slabodnick MM, Chen AH, and Goldstein B (2018). SapTrap assembly of repair templates for Cas9-triggered homologous recombination with a self-excising cassette. Micropublication: biology. Dataset. https://doi.org/10.17912/W2KT0N

Yumerefendi H, Wang H, Dickinson DJ, Lerner A, Malkus P, Goldstein B, Hahn K, and Kuhlman B (2018). Light-dependent cytoplasmic recruitment enhances the dynamic range of a nuclear import photoswitch. Chembiochem. doi: 10.1002/cbic.201700681.

Fadero TC, Gerbich TM, Rana K, Suzuki A, DiSalvo M, Schaefer KN, Heppert JK, Boothby TC, Goldstein B, Peifer M, Allbritton NL, Gladfelter AS, Maddox AS and Maddox PS (2018). LITE microscopy: Tilted light-sheet excitation of model organisms offers high resolution and low photobleaching. Journal of Cell Biology 217:1869-1882.

Martinez P, Allsman LA, Brakke KA, Hoyt C, Hayes J, Liang H, Neher W, Rui Y, Roberts AM, Moradifam A, Goldstein B, Anderson CT, Rasmussen C (2018). Predicting division planes of three-dimensional cells by soap-film minimization. Plant Cell pii: tpc.00401.2018. doi: 10.1105/tpc.18.00401.

Dickinson DJ, Schwaeger F, Pintard L, Gotta M, and B Goldstein (2017). A Single-Cell Biochemistry Approach Reveals PAR Complex Dynamics during Cell Polarization. Developmental Cell 42:416-434.

Boothby TC, Tapia H, Brozena AH, Piszkiewicz S, Smith AE, Giovanninni I, Rebecchi L, Pielak GJ, Koshland D, and B Goldstein (2017). Tardigrades use intrinsically disordered proteins to survive desiccation. Molecular Cell 65:975-984.

Smith FW, Bartels PJ, and Goldstein B (2017). A hypothesis for the composition of the tardigrade brain and its implications for panarthropod brain evolution. Integrative and Comparative Biology 57:546-559.

Russell JJ, Theriot JA, Sood P, Marshall WF, Landweber LF, Fritz-Laylin L, Polka JK, Oliferenko S, Gerbich T, Gladfelter A, Umen J, Bezanilla M, Lancaster MA, He S, Gibson MC, Goldstein B, Tanaka EM, Hu C-K, and Brunet A (2017). Non-model model organisms. BMC Biology 15:55.

Smith FW and B Goldstein (2017). Segmentation in Tardigrada and diversification of segmental patterns in Panarthropoda. Arthropod Structure & Development 46:328-340.

Zallen JA and Goldstein B (2017). Cellular mechanisms of morphogenesis. Semin Cell Dev Biol 67:101-102.

Goldstein B and Zallen JA (2017). Cell Polarity and Morphogenesis: new technologies and new findings. Molecular Biology of the Cell, 28:699-700.

Tintori SC, Osborne Nishimura E, Golden P, Lieb JD, Goldstein B (2016). A Transcriptional Lineage of the Early C. elegans Embryo. Dev Cell 38:430-44.

Goldstein B and King N (2016). The Future of Cell Biology: Emerging Model Organisms. Trends in Cell Biology 11:818-824.

Goldstein B (2016). Sydney Brenner on the Genetics of Caenorhabditis elegans. Genetics 204:1-2.

Heppert JK, Dickinson DJ, Pani AM, Higgins CD, Steward A, Ahringer J, Kuhn JR, and B Goldstein (2016). Comparative assessment of fluorescent proteins for in vivo imaging in an animal model system. Molecular Biology of the Cell 27:3385-3394 and Spectrum Viewer

Marston DJ*, Higgins CD*, Peters KA, Cupp TD, Dickinson DJ, Pani AM, Moore RP, Cox AH, Kiehart DP, and Goldstein B (2016). MRCK-1 drives apical constriction in C. elegans by linking developmental patterning to force generation. Current Biology 26:2079-2089.

Dickinson DJ and Goldstein B (2016). CRISPR-based methods for Caenorhabditis elegans genome engineering. Genetics 202:885-901.

Smith FW, Boothby TC, Giovannini I, Rebecchi L, Jockusch EL, and B Goldstein (2016). The compact body plan of tardigrades evolved by the loss of a large body region. Current Biology 26:224-229.

Sullivan-Brown J, Tandon P, Bird KE, Dickinson DJ, Tintori SC, Heppert JK, Meserve JH, Trogden KP, Orlowski SK, Conlon FL, and Goldstein B (2016). Identifying regulators of morphogenesis common to vertebrate neural tube closure and C. elegans gastrulation. Genetics 202:123-139.

Dickinson, D.J., A.M. Pani, J.K. Heppert, C.D. Higgins, and B. Goldstein (2015). Streamlined Genome Engineering with a Self-Excising Drug Selection Cassette. Genetics 200:1035-1049.

Osborne Nishimura E, Zhang JC, Werts AD, Goldstein B, Lieb JD (2015). Asymmetric transcript discovery by RNA-seq in C. elegans blastomeres identifies neg-1, a gene important for anterior morphogenesis. PLoS Genet. 11(4):e1005117.

Goldstein, B and DP Kiehart (2015). Moving Inward: Establishing the Mammalian Inner Cell Mass. Developmental Cell 34:385-386.

Martin A.C. and B. Goldstein (2014). Apical constriction: themes and variations on a cellular mechanism driving morphogenesis. Development 141:1987-98.

Dickinson, D.J., J.D. Ward, D.J. Reiner and B. Goldstein (2013). Engineering the Caenorhabditis elegans genome using Cas9-triggered homologous recombination. Nature Methods 10:1028-34.

Peters EC, Gossett AJ, Goldstein B, Der CJ, Reiner DJ (2013). Redundant Canonical and Noncanonical Caenorhabditis elegans p21-Activated Kinase Signaling Governs Distal Tip Cell Migrations. G3 3:181-95.

Tenlen, J.R., S. McCaskill and B. Goldstein (2013). RNA interference can be used to disrupt gene function in tardigrades. Development Genes and Evolution 223:171-81.

Gao, L., L. Shao, C.D. Higgins, J.S. Poulton, M. Peifer, M.W. Davidson,  X. Wu, B. Goldstein and E. Betzig (2012). Noninvasive imaging of 3D dynamics in thickly fluorescent specimens beyond the diffraction limit. Cell 151:1370-85.

Roh-Johnson, M., G. Shemer., C.D. Higgins, J.H. McClellan, A.D. Werts, U.S. Tulu, L. Gao, E. Betzig, D.P. Kiehart and B. Goldstein (2012) Triggering a Cell Shape Change by Exploiting Pre-Existing Actomyosin Contractions. Science 335:1232-1235.

Sullivan-Brown, J. and B. Goldstein (2012) Neural tube closure: The curious case of shrinking junctions. Current Biology 22:R574-R576.


Edgar, L.G. and B. Goldstein (2012). Culture and Manipulation of Embryonic Cells, in Caenorhabditis elegans: Cell Biology and Physiology, eds. Joel H. Rothman and Andrew Singson. Methods in Cell Biology, 107:151-17.

Tse, Y.C., M. Werner, K.M. Longhini, J.-C. Labbé, B. Goldstein and M. Glotzer (2012) RhoA activation during polarization and cytokinesis of the early C. elegans embryo are differentially dependent on NOP-1 and CYK-4. Molecular Biology of the Cell 23:4020-4031.

Werts, A.D., M. Roh-Johnson and B. Goldstein (2011). Dynamic localization of C. elegans TPR-GoLoco proteins mediates mitotic spindle orientation by extrinsic signaling. Development 138:4411-22.

Werts, A.D. and B. Goldstein (2011). How signaling between cells can orient a mitotic spindle. Seminars in Cell and Developmental Biology, 22:842-9.

Sawyer, J.M., S. Glass, T. Li, G. Shemer, N.D. White, N.G. Starostina, E.T. Kipreos, C.D. Jones, and B. Goldstein (2011). Overcoming Redundancy: an RNAi Enhancer Screen for Morphogenesis Genes in C. elegans. Genetics 188:549-564.

Harrell, J.R. and B. Goldstein (2011). Internalization of multiple cells during C. elegans gastrulation depends on common cytoskeletal mechanisms but different cell polarity and cell fate regulators. Developmental Biology 350:1-12.

Higgins C.D. and B. Goldstein (2010) Asymmetric Cell Division: A New Way to Divide Unequally. Current Biology R1029-31.

Roh-Johnson, M., J. Sullivan-Brown and B. Goldstein. (2010) Roles for Actin Dynamics in Cell Movements during Development. Chapter in Actin-Based Motility, Cellular, Molecular and Physical Aspects. ed. M.-F. Carlier, Springer-London.

Sawyer, J.M., J.R. Harrell, G. Shemer, J. Sullivan-Brown, M. Roh-Johnson and B. Goldstein (2010). Apical constriction: A cell shape change that can drive morphogenesis 341:5-19.

Arata, Y., J.-Y. Lee, B. Goldstein and H. Sawa (2010) Extracellular control of PAR protein localization during asymmetric cell division in the C. elegans embryo. Development 137:3337-3345.

Roh-Johnson, M. and B. Goldstein (2009). In vivo roles for Arp2/3 in cortical actin organization during C. elegans gastrulation. J Cell Sci 122: 3983-93.

McCarthy Campbell, E.K., A.D. Werts and B. Goldstein (2009). A Cell Cycle Timer for Asymmetric Spindle Positioning. PLoS Biology 7(4):e88.

Goldstein, B. and H. Hamada (2009). Shape Meets Polarity in Japan. Development 136:2487-92.

Marston, D.J., M. Roh, A. Mikels, R. Nusse, and B. Goldstein (2008). Wnt signaling during Caenorhabditis elegans embryonic development.  Methods in Molecular Biology 469:103-111.

Goldstein, B. and I. Macara (2007). The PAR Proteins: Fundamental Players in Animal Cell Polarization 13: 609-622.

Gabriel, W.N., R. McNuff, S.K. Patel, T.R. Gregory, W.R. Jeck, C.D. Jones and B. Goldstein (2007). The Tardigrade Hypsibius dujardini, a New Model for Studying the Evolution of Development.  Developmental Biology 312:545-559.

Gabriel, W.N. and B. Goldstein (2007). Segmental Expression of Pax3/7 and Engrailed Homologs in Tardigrade Development. Development Genes and Evolution 217:421-433.

Lee, J.-Y., D.J. Marston, T. Walston, J. Hardin, A. Halberstadt and B. Goldstein. (2006) Wnt/Frizzled Signaling Controls C. elegans Gastrulation by Activating Actomyosin Contractility. Current Biology, 16: 1986-1997.

Goldstein B., H. Takeshita, K. Mizumoto and H. Sawa (2006). Wnt Signals Can Function as Positional Cues in Establishing Cell Polarity. Developmental Cell 10: 391-396.

Marston, D.J. and B. Goldstein (2006). Symmetry Breaking in C. elegans: Another Gift from the Sperm. Developmental Cell 11: 273-274.

Marston, D.J. and B. Goldstein (2006). Actin-based forces driving embryonic morphogenesis in C. elegans. Current Opinion in Genetics and Development 16: 392-398.

McCarthy, E.K. and B. Goldstein (2006). Asymmetric Spindle Positioning. Current Opinion in Cell Biology 18: 79-85.

Marston, D.J. and B. Goldstein (2006). Symmetry Breaking in C. elegans: Another Gift from the Sperm. Developmental Cell 11: 273-274.

Nance, J., J.-Y. Lee and B. Goldstein (2005). Gastrulation in C. elegans, WormBook, ed. The C. elegans Research Community.

McCarthy, E.K. and B. Goldstein (2005). Asymmetric Division: A Kinesin for Spindle Positioning.  Current Biology 15:R591-593.

Dudley, N.R. and B. Goldstein (2005). RNA Interference in Caenorhabditis elegans. Methods in Molecular Biology 309:29-38.

Dudley, N.R., A.Z. Amin and B. Goldstein (2005). Genes Required for RNA Interference  (chapter in "RNA Interference: From Basic Science to Drug Development", Cambridge University Press, Cambridge, UK, ed. K. Appasani). (preprint)

Labbe, J.-C., E. McCarthy and B. Goldstein (2004). The forces that position a mitotic spindle asymmetrically are tethered until after the time of spindle assembly. The Journal of Cell Biology 167:245-256.

Cheeks, R.J., J.C. Canman, W.N. Gabriel, N. Meyer, S. Strome and B. Goldstein (2004). C. elegans PAR Proteins Function by Mobilizing and Stabilizing Asymmetrically Localized Protein Complexes.  Current Biology 14:851-862.

Labbe, J.-C., P.S. Maddox, E.D. Salmon, and B. Goldstein (2003). PAR proteins regulate microtubule dynamics at the cell cortex in C. elegans.  Current Biology 13:707-714.

Lee, J.-Y. and B. Goldstein (2003). Mechanisms of cell positioning during C. elegans gastrulation.  Development 130:307-320.

Goldstein, B. (2003). Asymmetric Division: AGS Proteins Position the Spindle.  Current Biology 13:R879-R880.

Dudley, N.R. and B. Goldstein (2003). RNA interference: Silencing in the cytoplasm and nucleus.  Curr Opin Mol Ther 5:113-117.

Labbé, J.-C. and B. Goldstein (2002). Embryonic Development: A New SPN on Cell Fate Specification.  Current Biology 12:R396-R398.

Goldstein, B. and M. Blaxter (2002). Tardigrades. Current Biology, 12:R475.

Dudley, N.R., J.-C. Labbé, and B. Goldstein (2002). Using RNA Interference to Identify Genes Required for RNA Interference.  PNAS 99:4191-4196.

Goldstein, B., M. Leviten and D. A. Weisblat (2001). Dorsal and Snail homologs in leech development.  Development Genes and Evolution 211:329-337.

Goldstein, B. (2001). On the Evolution of Early Development in the Nematoda. Phil. Trans. Roy. Soc. Lond. B. 356:1521-1531.

Goldstein, B. (2000). Embryonic polarity: A role for microtubules. Current Biology 10:R820-R822.

Goldstein, B. (2000). When cells tell their neighbors which direction to divide. Developmental Dynamics,218:23-29.

Goldstein, B., L. Frisse and W. K. Thomas (1998). Embryonic axis specification in nematodes: evolution of the first step in development. Current Biology 8:157-160.

Wittmann, C., O. Bossinger, B. Goldstein, M. Fleischmann, R. Kohler, K. Brunschwig, H. Tobler and F. Müller (1997). The expression of the C. elegans labial-like Hox gene ceh-13 during early embryogenesis relies on cell fate and on anteroposterior cell polarity. Development 124:4193-4200.

Goldstein, B. and S. N. Hird (1996). Specification of the anteroposterior axis in Caenorhabditis elegans. Development 122:1467-1474.

Goldstein, B. (1995). Cell contacts orient some cell division axes in the early C. elegans embryo. Journal of Cell Biology 129:1071-1080.

Goldstein, B. (1995). An analysis of the response to gut induction in the C. elegans embryo. Development121:1227-1236.

Goldstein, B., S. N. Hird, and J. G.White (1993). Cell polarity in early C. elegans development. Development 1993 Supplement:279-287.

Goldstein, B. (1993). Establishment of gut fate in the E lineage of C. elegans: the roles of lineage-dependent mechanisms and cell interactions. Development 118:1267-1277.

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