• S.B. Brain and Cognitive Sciences, S.B. Biology, Massachusetts Institute of Technology (2006)
• Ph.D. Biological and Biomedical Sciences, Harvard University (2013)
• Postdoctoral Fellowship, Division of Genetics and Genomics, Children’s Hospital Boston and Harvard Medical School (2016)
• Postdoctoral Fellowship, Department of Ophthalmology, Stanford University (2023)

Courses Taught

• FYS 505 STEM Scholars
• NRSC 160 Introduction to Neuroscience
• NRSC 311 The Hypothalamus
• NRSC 325 Neural Development
• NRSC 334 Medical Genetics
• NRSC 335 Degeneration and Regeneration of the Nervous System
• NRSC s22 Methods in Developmental Neuroscience

Research Interests

Vision loss is a devastating medical problem that leads to lower quality of life and loss of independence among those affected. Because the human retina has minimal or no regenerative ability, the death of retinal neurons due to injury or disease is generally irreversible, making this the most common cause of permanent visual impairment. If retinal neurons could be regenerated from progenitor cells that live within the adult human body, these patients could have their vision restored.

I study the development of retinal ganglion cells, the neurons that connect the eye with the brain. These neurons are vulnerable to injury in traumatic optic nerve injuries and to diseases such as glaucoma, and a deeper understanding of their development and regeneration could have significant implications for reversing visual impairment. I investigate retinal ganglion cells through the lens of development, by seeking to understand the way retinal ganglion cells normally develop and the ways these developmental pathways could be exploited to encourage regeneration in adult animals, using mice as a model system.

Selected Publications

1. Cameron EG, Nahmou M, Toth AB, Heo L, Tanasa B, Dalal R, Yan W, Nallagatla P, Xia X, Hay S, Knasel C, Stile TL, Douglas C, Atkins M, Sun C, Ashouri M, Bian M, Chang KC, Russano K, Shah S, Woodworth MB, Galvao J, Nair RV, Kapiloff MS, Goldberg JL. “A molecular switch for neuroprotective astrocyte reactivity.” (2024) Nature 626, 574-582. PMID: 38086421
2. Woodworth MB, Greig LC, Goldberg J. (2023) “Intrinsic and induced neuronal regeneration in the mammalian retina.” Antioxidants and Redox Signaling 39(16-18): 1039-1052. PMID: 37276181.
3. Woodworth MB, Girskis K, Walsh CA. (2017) “Building a lineage from single cells: Genetic techniques for cell lineage tracking.” Nature Reviews Genetics 18(4): 230-244. PMID: 28111472. With cover.
4. Greig LC*, Woodworth MB*, Greppi C, Macklis JD. (2016) “Ctip1 controls acquisition of sensory area identity and establishment of sensory input fields in the developing neocortex.” Neuron 90(2):261-277. PMID: 27100196. *equal contribution
5. Woodworth MB*, Greig LC*, Liu KX, Ippolito GC, Tucker HO, Macklis JD. (2016) “Ctip1 regulates the balance of projection neuron subtype specification in deep cortical layers.” Cell Reports 15(5): 999-1012. PMID: 27117402. With cover. *equal contribution
6. Lodato MA*, Woodworth MB*, Lee S*, Evrony GD, Mehta BK, Karger A, Lee S, Chittenden TW, D’Gama AM, Cai X, Luquette LJ, Lee E, Park PJ, Walsh CA. (2015) “Somatic mutation in single human neurons tracks developmental and transcriptional history.” Science 350(6256):94-8. PMID: 26430121. With cover. *equal contribution
7. Greig LC*, Woodworth MB*, Galazo MJ, Padmanabhan H, Macklis JD. (2013) “Molecular logic of neocortical projection neuron specification, development, and diversity.” Nature Reviews Neuroscience 14(11): 755-69. PMID: 24105342. *equal contribution
8. Woodworth MB*, Custo Greig L*, Kriegstein AR, Macklis JD. (2012) “Snapshot: Cortical development.” Cell 151(4): 918-918.e.1. PMID: 23141546. *equal contribution

Office hours: Tuesday, 10 – 11 a.m.

I am currently not accepting new requests for letters of recommendation.

Research Interests

I am broadly interested in how plants on a cell-to-cell level process external information that affects their development. More specifically, I want to understand how this information in the form of abiotic stresses such as heat or drought is interpreted on a genomic level at various stages of development. My research primarily focuses on understanding how this information is stored and encoded in the context of chromatin and 3-dimensional contacts of DNA and its associated proteins inside the nucleus so that plants “learn” how to adapt to recurring stress situations. In my lab, we use next-generation sequencing, plant physiological, genetic, molecular and cell biology approaches to tackle these questions. Especially in light of climate change, it will be important to understand how plants react to changing environmental conditions.

Publications

Chen H, Xiong F, Wangler AM, Bischoff T, Wang K, Miao Y, Slane D, Schwab R, Laux T, and Bayer M. Phosphorylation-Dependent Activation of the bHLH Transcription Factor ICE1/SCRM Promotes Polarization of the Arabidopsis Zygote. New Phytologist. 2025 Feb;245(3):1029-1039. doi: 10.1111/nph.20265. Epub 2024 Nov 14. PMID: 39543803.

Berendzen KW, Grefen Christopher, Sakamoto Takuya, and Slane D. Analysis of Chromatin Accessibility, Histone Modifications, and Transcriptional States in Specific Cell Types Using Flow Cytometry. Methods in Molecular Biology. 2023, 2698:57-73.

Sakamoto T, Sakamoto Y, Grob S, Slane D, Yamashita T, Ito N, Oko Y, Sugiyama T, Higaki T, Hasezawa S, Tanaka M, Matsui A, Seki M, Suzuki T, Grossniklaus U, and Matsunaga S. Two-step regulation of centromere distribution by condensin II and the nuclear envelope proteins. Nature Plants. 2022 Aug;8(8):940-953.

Slane D, Lee CH, Kolb M, Dent C, Miao Y, Franz-Wachtel M, Lau S, Maček B, Balasubramanian S, Bayer M, and Jürgens G. The integral spliceosomal component CWC15 is required for development in Arabidopsis. Scientific Reports. 2020 Aug 7;10(1):13336.

Smit ME, Llavata-Peris CI, Roosjen M, van Beijnum H, Novikova D, Levitsky V, Sevilem I, Roszak P, Slane D, Jürgens G, Mironova V, Brady SM, and Weijers D. Specification and regulation of vascular tissue identity in the Arabidopsis embryo. Development. 2020 Apr 20;147(8):dev186130.

Slane D, Berendzen KW, Witthöft J, and Jürgens G. Transcriptomic Profiling of the Arabidopsis Embryonic Epidermis Using FANS in Combination with RNAseq. Methods in Molecular Biology. 2020;2122:151-164.

Neu A, Eilbert E, Asseck LY, Slane D, Henschen A, Wang K, Bürgel P, Hildebrandt M, Musielak TJ, Kolb M, Lukowitz W, Grefen C, and Bayer M. Constitutive signaling activity of a receptor-associated protein links fertilization with embryonic patterning in Arabidopsis thaliana. Proceedings of the National Academy of Sciences of the United States of America. 2019 Mar 19;116(12):5795-5804.

Wallmeroth N, Jeschke D, Slane D, Nägele J, Veerabagu M, Mira-Rodado V, and Berendzen KW. ARR22 overexpression can suppress plant Two-Component Regulatory Systems. PLoS ONE. 2019 Feb 11;14(2):e0212056.

Slane D, Reichardt I, El Kasmi F, Bayer M, and Jürgens G. Evolutionarily diverse SYP1 Qa-SNAREs jointly sustain pollen tube growth in Arabidopsis. The Plant Journal. 2017 Nov;92(3):375-385.

Slane D, Bürgel P, and Bayer M. Staining and Clearing of Arabidopsis Reproductive Tissue for Imaging of Fluorescent Proteins. Methods in Molecular Biology. 2017;1669:87-94.

Slane D and Bayer M. Cell Type-Specific Gene Expression Profiling Using Fluorescence-Activated Nuclear Sorting. Methods in Molecular Biology. 2017;1629:27-35.

Bayer M, Slane D, and Jürgens G. Early plant embryogenesis-dark ages or dark matter? Current Opinion in Plant Biology. 2017 Feb;35:30-36.

Musielak TJ, Slane D, Liebig C, and Bayer M. A Versatile Optical Clearing Protocol for Deep Tissue Imaging of Fluorescent Proteins in Arabidopsis thaliana. PLoS ONE. 2016 Aug 12;11(8):e0161107.

Murphy E, Vu LD, Van den Broeck L, Lin Z, Ramakrishna P, van de Cotte B, Gaudinier A, Goh T, Slane D, Beeckman T, Inzé D, Brady SM, Fukaki H, De Smet I. RALFL34 regulates formative cell divisions in Arabidopsis pericycle during lateral root initiation. Journal of Experimental Botany. 2016 Aug;67(16):4863-4875.

Slane D, Kong J, Schmid M, Jürgens G, Bayer M. Profiling of embryonic nuclear vs. cellular RNA in Arabidopsis thaliana. Genomics Data. 2015 Apr 8;4:96-98.

Slane D, Kong J, Berendzen KW, Kilian J, Henschen A, Kolb M, Schmid M, Harter K, Mayer U, De Smet I, Bayer M, Jürgens G. Cell type-specific transcriptome analysis in the early Arabidopsis thaliana embryo. Development. 2014 Dec;141(24):4831-4840.

Lau S, Slane D, Herud O, Kong J, Jürgens G. Early Embryogenesis in Flowering Plants: Setting Up the Basic Body Pattern. Annual Review of Plant Biology. 2012;63:483-506.

Reichardt I, Slane D, El Kasmi F, Knöll C, Fuchs R, Mayer U, Lipka V, Jürgens G. Mechanisms of Functional Specificity Among Plasma-Membrane Syntaxins in Arabidopsis.��Traffic.��2011 Sep;12(9):1269-1280.

Ph.D. Human Genetics, University of Houston

Research interests
My research is focused on studying the evolution of conserved gene regulatory networks. In my lab, we are particularly interested in how changes to genomes and gene regulatory networks have led to the diversity of animal forms and functions. We use marine and freshwater sponges as model systems to ask questions about the genetics and development of animal evolution and symbioses. Sponges are ancient animals that retain characteristics of an early and successful experiment in multicellularity while also sharing some highly conserved features (e.g., stem cells) and molecular blueprints with all other animals, including humans. These unique features provide a system where we can explore hypotheses about the evolution of all animals.

Student Research Opportunities
Research projects in my lab focus on the role of conserved developmental control genes and gene regulatory networks that are uniquely animal, but may have originated prior to the advent of adaptations such as nervous and immune systems, muscles, or eyes. We also study gene networks and the molecular linkages between intracellular algal symbionts and sponges as we try to understand how the host:symbiont interaction is important for animal development and function.  Along these lines, some students also study the unique features of the symbiotic microalgae that inhabit sponge cells. We employ molecular, cellular, developmental, and functional genomic approaches and frequently collaborate with field biologists and computational scientists to address our questions. The research projects in my lab have implications for basic biomedical research as well as environmental and climate change studies.

Selected Publications

(*denotes student co-author)

A.L. Horton, H. Neighmond*, A. Neighmond*, R. Anderson*, M. Lessard, V. Price, S.P. Leys, A. Riesgo. (2026) Molecular and spatial integration of algal endosymbionts of the freshwater sponge, Ephydatia muelleri, throughout development in light and dark conditions. BMC Genomics. 27, 763. https://doi.org/10.1186/s12864-026-12618-w

R. Cassidy, L. de la Cruz, K. Mitsi, C. Galià-Camps, A. Benítez-López, C. Gracia-Sancha, J. Lorente-Sorolla, A. Álvarez, R. Mozo, S. Kolomyjec, S. Nichols, R. Manconi, R. Pereira, K. Evans, V. Itskovitch, A.L. Horton, S.P. Leys, S. Taboada, A. Riesgo. (2026) Genomic Connectivity and Adaptation Signals of the Freshwater Sponge Ephydatia muelleri across its distribution. Journal of Biogeography. 53:e70142.

C. Cevallos*, A.L. Leigh White*, B. Fazio*, L. Wendt, J. Feng, D. Posfai, A.L. Horton, J. Warrick, O.A. Quintero-Carmona. (2025) Transcriptomic Analysis of CAD Cell Differentiation. microPublication Biology. 10.17912.

S.P. Leys, L Grombacher, D. Field, V. Ho, G.R.D. Elliot, A.S. Kahn, P. Reid, A. Riesgo, E. Lanna, Y. Bobkov, J.F. Ryan, A.L. Horton. (2025) A Morphological Cell Atlas of the Freshwater Sponge, Ephydatia muelleri, with Key Insights from Targeted Single-Cell Transcriptomes. EvoDevo 16:1.

H. Neighmond*, A. Quinn*, B. Schmandt*, K. Ettinger*, A.L. Hill, L. Williams. (2023) Developmental Bisphenol S Toxicity in Two Freshwater Animal Models. Environmental Toxicology and Pharmacology 104:104311..

M.S. Hill, B. Lawson, J.W. Cain, N. Rahman*, S. Toolsidass*, T. Wang*, S. Geraghty*, E. Raymundo*, A.L. Hill, (2023) Sustained Beneficial Infections: Priority Effects, Competition, and Specialization Drive Patterns of Association in Phototroph:Heterotroph Mutualisms. Frontiers in Ecology and Evolution 11.

K. Hustus, K. Mitsi, J. Nutakki*, V. Kering*, I. Nguyen*, M. Gomes Spencer*, S.P. Leys, M.S. Hill, A. Riesgo, A.L. Hill (2023) Algal Symbionts of the Freshwater Sponge Ephydatia muelleri. Symbiosis. https://doi.org/10.1007/s13199-023-00934-8.

S. Jones, A. Blake, L. Corado-Santiago, J Crenshaw, E. Goldman, F. Gomez, C. Hall, H. Hoke, S. Holmes, B. Kornegay, P. Kwarteng, B. Lawson, M. Leber, G. Leconte, E. Modeste, K. Nolin, M. Norris, J. Santinni Roma, A. Swackhammer, M. Torres, J. Wares, D. Williams, A. Hill, K. Hoke, C. Parish, BD Pierce. (2023) A SMART Decade: Outcomes of an Integrated, Inclusive, First-Year College-Level STEM Curricular Innovation. Frontiers in Education 8:1152339. doi: 10.3389/feduc.2023.1152339

S. Geraghty*, V. Koutsouveli, C. Hall, L. Chang*, O. Sacristan-Sorianob , M.S. Hill, A. Riesgo, A.L. Hill (2021) Establishment of host:algal endosymbioses: Genetic response to symbiont versus prey in a sponge host. Genome Biology and Evolution. 13: evab252..

C. Hall, S, Camilli, H. Dwaah, B. Kornegay, C. Lacy, M.S. Hill, A.L. Hill (2021) Freshwater sponge hosts and their green algae symbionts: a tractable model to understand intracellular symbiosis. PeerJ.��11:e10654. doi: 10.7717/peerj.10654

N.J. Kenny, W.R. Francis, R.E. Rivera-Vicéns, K. Jurval, A. de Mendoza, C. Díez-Vives, R. Lister, L. Bezares-Calderon, L. Grombacher, M. Roller, L.D. Barlow, S. Camilli*, J.F. Ryan, G. Wöheide, A.L. Hill, A. Riesgo, S.P. Leys (2020) Tracing animal genomic evolution with the chromosomal-level assembly of the freshwater sponge Ephydatia muelleri. Nature Communications. 11: 3676. https://rdcu.be/b5ROn

C. Hall, M. Rodriguez*, J. Garcia*, D. Posfai*, R. Dumez*, E. Wictor*, O. Quintero, M. Hill, A. Rivera, A. Hill (2019) Secreted frizzled related protein is a target of PaxB and plays a role in aquiferous system development in the freshwater sponge,��Ephydatia muelleri. PLOS ONE. https://doi.org/10.1371/journal. pone.0212005

O. Sacristán-Soriano, M. Winkler, P. Erwin, J. Weisz, O. Harriott, G. Heussler, E. Bauer, B. West Marsden*, A. Hill, M. Hill (2019) Ontogeny of symbiont community structure in two carotenoid-rich, viviparous marine sponges: comparison of microbiomes and analysis of culturable pigmented heterotrophic bacteria. Environmental Microbiology Reports.

P. Windsor-Reid, E. Matveev, A. McClymont, D. Posfai*, A. Hill, S.P. Leys (2018) Wnt signaling and polarity in freshwater  sponges. BMC Evolutionary Biology. 18:12.

J. Cramer, D. Pohlmann*, F. Gomez, L. Mark*, B. Kornegay*, C. Hall*, N. Walavalkar, S. Bilinovich, J. Prokop, A. Hill, D. Williams (2017) Methylation specific targeting of a chromatin remodeling complex from sponges to humans.��Scientific Reports. 7:40674.

Q. Schenkelaars, O. Quintero, C. Hall*, L. Fierro-Constain, E. Renard, C. Borchiellini, A. Hill (2016) ROCK inhibition abolishes the establishment of the aquiferous system in Ephydatia muelleri (Porifera, Demospongiae).��Developmental Biology. 412: 298-310.

Research Interests

Selected Publications

* denotes �ֲ���ý student

Williams LM, Bowsher AM, *Chrysovergi M-A, Ambrose WG Jr (2020). Bloodworm (Glycera dibranchiata Ehlers, 1868) populations in the Gulf of Maine are connected through gene flow.��Marine Science and Biology.��1:��1-4.

Sant KE, *Moreau HM,��Williams LM, Jacobs HM, Bowsher AM, *Boisvert JD, Smolowitz RM, *Pantazis J, Timme-Laragy A (2020). Embryonic exposures to mono-2-ethylhexyl phthalate induce larval steatosis in zebrafish independent of Nrf2a signaling.��Journal of Developmental Origins of Health and Disease.��doi: 10.1017/S2040174420000057.

*Ulin A, *Henderson J, *Pham M-T, *Meyo J, *Chen Y, Karchner SI, Goldstone JV, Hahn ME,��Williams LM (2019). Developmental regulation of nuclear factor erythroid-2 related factors (Nrfs) by AHR1b in zebrafish (Danio rerio).��Toxicological Sciences,��167(2): 536-545.

Jacobs HM, Sant KE, Basnet A,��Williams LM, Moss JB, Timme-Laragy A (2018). Embryonic exposure to Mono(2-ethylhexyl) phthalate (MEHP) disrupts pancreatic organogenesis in zebrafish (Danio rerio).�����𳾴Dz��������,��195:��498-507.

Sant KE, Hansen JM,��Williams LM, *Tran NL, Goldstone JV, Stegeman JJ, Hahn ME, Timme-Laragy A (2017). The role of Nrf1 and Nrf2 in the regulation of glutathione and redox dynamics in the developing embryos.��Redox Biology,��13:��207-218.

Lord JP,��Williams LM (2017). Northward Expansion of Genetically Diverse Invasive Asian Shore Crab (Hemigrapsus sanguineus) Populations.��Biological Invasions,��19(4): 1153-1168.

Williams LM, *Lago B, McArthur AG, Raphenya AR, *Pray N, *Saleem N, *Salas S, *Paulson K, *Mangar R, Liu Y, Vo AH, Shavit J (2016). The transcription factor, Nuclear factor, erythroid 2 (Nfe2), is a regulator of the oxidative stress response during Danio rerio development.��Aquatic Toxicology,��180:��141-154.

Williams LM, *Nivison CL, Ambrose WG Jr, *Dobbin R, Locke WL V (2015). Lack of adult novel northern lineages of invasive green crab Carcinus maenas along much of the northern US Atlantic coast.��Marine Ecology Progress Series,��532:��153-159.