Daniel Shain Research

Daniel H. Shain
Professor of Zoology
Biology Department


  • University of New Hampshire, BS/MS (1988)
  • Colorado State University, Ph.D. (1994)
  • University of California, Berkelely, Post-doc (1995-1998)

Research interests: We are interested in all aspects of worm biology, from the bioenergetic adaptation of glacier ice worms to the secretion of leech cocoons and production of embryonic stem cells. We also have ongoing projects exploring the evolution and adaptation of extremophilic animals (e.g., Rotifera, Tardigrada) that occupy glaciers and hot springs worldwide, including from Iceland, New Zealand, Norway and Tibet.

Selected Publications

*student author

  1. Shain DH, Salvadore C and Denis CL (1992) Evolution of the alcohol dehydrogenase (ADH) genes in yeast:  characterization of a fourth ADH in Kluyveromyces lactis. Gen. Genet. 232, 479-488.
  2. Shain DH, Yoo J, Slaughter RG, Hayes SE and Ji TH (1992) Electrofractionation:  a technique for detecting and recovering biomolecules.  Biochem. 200, 47-51.
  3. Neuman T, Keen A, Knapik E, Shain D, Ross M, Nornes HO and Zuber MX (1993) ME1 and GE1: basic helix-loop-helix transcription factors expressed at high levels in the developing nervous system and in morphogenetically active regions. Eur. J. Neuroscience 5, 311-318.
  4. Shain DH, Neuman T and Zuber MX (1995) A novel initiator regulates expression of the nontissue-specific helix-loop-helix gene ME1.  Nucl. Acids Res. 23, 1696-1703.
  5. Shain DH, Haile, DT, Verrastro TA and Zuber MX (1995) Cloning and embryonic expression of Xenopus laevis GAP-43 (XGAP-43).  Brain Res. 697, 241-246.
  6. Shain DH and Zuber MX (1995) Identification of nontissue-specific helix-loop-helix genes in Xenopus laevis.  Gene 165, 319-320.
  7. Shain DH, Stone RT, Yoo J and Zuber MX (1996) Characterization of a single-locus minisattelite DNA in Xenopus laevis.  Genome 39, 230-233.
  8. Shain DH and Zuber MX (1996) SDS-based whole-mount in situ hybridization of Xenopus laevis J. Biochem. Biophys. Methods 31, 185-188.
  9. Shain DH, Neuman T and Zuber MX (1997) Embryonic expression and evolution of duplicated E-protein genes in Xenopus laevis: parallels with ancestral E-proteins. Genetics 146, 345-353.
  10. Shain DH, Zuber MX and Neuman T (1998) Transcription initiation from a poly d(A) tract. Acids Res. 26(4), 1019-1025.
  11. Shain DH, Ramirez FA, Hsu J and Weisblat DA (1998) Gangliogenesis in leech:  morphogenetic processes leading to segmentation in the central nervous system. Dev. Genes Evol. 208, 28-36.
  12. Shain DH, Zuber MX, Yoo J, Norris J and Neuman T (1998) Selective conservation of an E-protein gene promoter during vertebrate evolution. FEBS Letters 440, 332-336.
  13. Shain DH, Stuart D and Weisblat DA (2000) Segmentation of the central nervous system in leech.  Development 127, 735-744.
  14. Shain DH, Carter MR, Murray KP, Maleski KA, Smith NR, *McBride TR, *Michalewicz LA and Saidel, WM (2000) Morphologic characterization of the ice worm Mesenchytraeus solifugus.  J. Morphology 246,192-197.
  15. Shain DH, *Mason TA, *Farrell AH, and *Michalewicz LA (2001) Distribution and behavior of ice worms (Mesenchytraeus solifugus) in south-central Alaska. Can. J. Zoology 79, 1813-1821.
  16. Duchovnay B and Shain DH (2002) Worms on Ice.  The Science Teacher 69, 20-25.
  17. *Dimitriu C and Shain DH (2004) Ultrastructural properties of the Theromyzon (Annelida: Hirudinea) cocoon membrane.  Micron 35, 281-285.
  18. *Mason TA, McIlroy PJ and Shain DH (2004) A cysteine-rich protein in the Theromyzon(Annelida: Hirudinea) cocoon membrane. FEBS Lett. 561, 167-172.
  19. *Napolitano MJ, Nagele RO and Shain DH (2004) The ice worm, Mesenchytraeus solifugus,elevates adenylate levels at low physiological temperature.  Comp. Biochem. Physiol. Part A 137, 227-235.
  20. *Hohenstein KA and Shain DH (2004) Changes in gene expression at the precursor  stem cell transition in leech.  Stem Cells 22, 84-91.
  21. *Napolitano MJ and Shain DH (2004) Four kingdoms on glacier ice: convergent energetic processes boost energy levels as temperatures fall. Proc. R. Soc. B. (Suppl.) 271, S273-S276.
  22. Shain DH (2004) The ice worm’s secret? Alaska Park Science 3, 31.
  23. Shain DH, Stuart D, Huang, FZ and Weisblat DA.  (2004) Cell interactions affecting axonogenesis in the leech, Theromyzon rude. Development 131, 4143-4153.
  24. *Farrell AH, *Hohenstein KA and Shain DH (2004) Molecular adaptation of the ice worm, Mesenchytraeus solifugus: divergence of energetic-associated genes. J. Mol. Evol. 56, 666-673.
  25. *Napolitano MJ and Shain DH (2005) Distinctions in adenylate metabolism among organisms inhabiting temperature extremes.  Extremophiles 9, 93-98.
  26. *Mason TA, *Sayers CW, *Paulson TL, *Coleman J and Shain DH (2005) Cocoon deposition and hatching in the aquatic leech, Theromyzon tessulatumAm. Midl. Nat. 154, 78-87.
  27. *Napolitano MJ and Shain DH (2005) Quantitating adenylate levels in diverse organisms. J. Biochem. Biophys. Methods 63, 69-77.
  28. Hartzell PL, *Nghiem JV, Richio KJ and Shain DH (2005) Distribution and phylogeny of glacier ice worms (Mesenchytraeus solifugus and M. solifugus rainierensis).  Can. J. Zool. 83, 1206-1213.
  29. *Hohenstein KA and Shain DH (2006) Divergence of F1 ATP synthase subunits in the ice worm, Mesenchytraeus solifugus. Hydrobiologia 564, 51-58.
  30. *Dimitriu C, *Sayers CW, *Coleman J and Shain DH (2006) Two-dimensional ultrastructural elements lead to three-dimensional reconstruction of protuberances on the cocoon membrane of the leech, Theromyzon tessulatum. Tissue & Cell 38, 35-41.
  31. *Mason TA, McIlroy PJ and Shain DH (2006) Structural model of an antistasin/notch-like fusion protein in the cocoon wall of the aquatic leech, Theromyzon tessulatum. Journal of Molecular Modeling 12, 829-834.
  32. Hong D, Man S, Shain DH and Martin JV (2006) Development of maximized total capillary surface: the quarter power scaling law from a developmental perspective. Proc. Int. Conf. Bioinform. Comp. Biol.
  33. Marotta R and Shain DH (2007) Irregular helicoids in leech cocoon membranes.  Journal of Structural Biology 158, 336-343.
  34. Shain DH, *Aldea Guevara MI, *Lopez Coral A, Crampton WGR and Albert JS (2007) A survey of freshwater annelids in the Peruvian Amazon. ACTA Hydrobiologia Sinica 31, 47-51.
  35. *Morrison BA and Shain DH (2008) An AMP nucleosidase gene knockout in Escherichia colielevates intracellular ATP levels and increases cold tolerance. Biol. Lett. 4, 53-56.
  36. *Coleman J, Marotta R and Shain DH (2008) Surface topology and structural integrity of the Theromyzon tessulatum (Annelida: Hirudinea: Glossiphoniidae) cocoon. Journal of Morphology 269, 812-819.
  37. *Tartaglia L and Shain DH (2008) Cold adapted tubulins in the ice worm, Mesenchytraeus solifugus. Gene 423, 135-141.
  38. *Sayers CW, *Coleman J and Shain DH (2009) Cell dynamics during cocoon secretion in the aquatic leech Theromyzon tessulatum. Tissue & Cell 41, 35-42.
  39. Marotta R, *Parry BR and Shain DH (2009) Divergence of AMP deaminase in the ice worm Mesenchytraeus solifugus (Annelida, Clitellata, Enchytraeidae). Int. J. Evol. Biol., ID 715086, doi:10.4061/2009/715086.
  40. *Wirchansky BA and Shain DH (2010) A new species of Haemopis (Annelida: Hirudinea): Evolution of North American terrestrial leeches. Mol. Phylogenet. Evol. 54, 226-234.
  41. *Hohenstein KA, *Lang SA, *Nuthulaganti T and Shain DH (2010) A glutamine-rich factor affects stem cell genesis in leech. Stem Cells International doi:10.4061/2010/145183
  42. *Purzycki C and Shain DH (2010) Fungal toxins and multiple sclerosis: A compelling connection. Brain Res Bull. 82, 4-6.
  43. *Parry BR and Shain DH (2011) Manipulations of AMP metabolic genes increase growth rate and cold tolerance in Escherichia coli: Implications for psychrophilic evolution. Mol. Biol Evol. 28, 2139-2145.
  44. Shain DH and Gagliardi LJ (2011) Can molecular cell biology explain chromosome motions? Theoretical Biology and Medical Modeling 8, 15.
  45. *Lang SA, Garcia MV, James SW, *Sayers CW, Shain DH (2012) Phylogeny and clitellar morphology of the giant Amazonian earthworm Rhinodrilus priollii (Oligochaeta: Glossoscolecidae). American Midland Naturalist 167:384-395.
  46. Dial CR, Dial RJ, *Saunders R, *Lang SA, Tetreau MD, Lee B, Wimberger P, *Dinapoli MS, *Egiazarov AS, *Gipple SL, *Maghirang MR, *Swartley-McArdle DJ, *Yudkovitz SR, Shain DH(2012) Historical biogeography of the North American glacier ice worm, Mesenchytraeus solifugus(Annelida: Oligochaeta: Enchytraeidae). Mol. Phylogenet. Evol. 63, 577-584.
  47. *Choudhari S, Smith S, Owens S, Gilbert J, Shain DH, Dial RJ, Grigoriev A (2013) Metagenome Sequencing of Prokaryotic Microbiota Collected from Byron Glacier, Alaska. Genome Announcements 1, 9-13.
  48. Gagliardi LJ and Shain DH (2013) Is intracellular pH a clock for mitosis? Theortical Biology and Medical Modeling 10, 8-9.
  49. *Rossi A, Marotta R, Saglam N, Shain DH (2013) Operculum ultrastructure in leech cocoons. Journal of Morphology 274, 940-946.
  50. Gagliardi LJ and Shain DH (2014) Chromosome congression explained by nanoscale Electrostatics. Theoretical Biology and Medical Modeling 11, 12-19.
  51. Gagliardi LJ and Shain DH (2014) Polar electrostatic forces drive poleward chromosome motions. Cell Division 9, 191-198.
  52. *Choudhari S, Dial RJ, Kumar D, Shain DH, Grigoriev A. (2014) Sequence composition diversity in Alaskan glacier and other metagenomes. PeerJ PrePrints 2:e734v1 https://dx.doi.org/10.7287/peerj.preprints.734v1
  53. Gagliardi LJ, Shain DH (2015) Intracellular pH as an electrostatic regulator of the spindle assembly checkpoint. Medical Research Archives 2, 12-18.
  54. Kotchoni SO, Gachomo EW, *Slobodenko K, Shain DH (2016) AMP DEAMINASE suppression increases biomass, cold tolerance and oil content in green algae. Algal Research 16, 473-480.
  55. Saltmarsh DM, Bowser ML, Morton JM, *Lang SA, Shain DH, Dial RJ (2016) Distribution and abundance of exotic earthworms within a boreal forest system in southcentral Alaska. NeoBiota 28, 67-86.
  56. Dial RJ, Becker M, Hope AG, Dial CR, Thomas J, *Slobodenko K, Golden TS, Shain DH (2016) The role of temperature in the distribution of the glacier ice worm, Mesenchytraeus solifugus(Annelida: Oligochaeta: Enchytraeidae). Arctic, Antarctic, and Alpine Res. 48, 199-211.
  57. Shain DH, Halldórsdóttir K, Pálsson F, Aðalgeirsdóttir G, Gunnarsson A, Jónsson T, *Lang SA, Pálsson HS, Steinþórssson S, Arnason E (2016) Colonization of Maritime Glacier Ice by Bdelloid Rotifera. Mol. Phylog. Evol. 98, 280–287.
  58. Saglam N, *Saunders R, *Lang S, Shain DH (2016) A new species of Hirudo (Annelida: Hirudinidae): biogeographical history of Eurasian medicinal leeches. BMC Zoology 1:5, DOI 10.1186/s40850-016-0002
  59. Gagliardi LJ, Shain DH (2016) Electrostatic forces drive poleward chromosome motions at kinetochore. Cell Division 11, 1-8.
  60. *Rossi A, Saidel WM, *Gravante C, Sayers CW, Shain DH (2016) Mechanics of cocoon secretion in a segmented worm (Annelida: Hirudinae). Micron 86, 30-35.
  61. *Lang SA, Saglam N, *Kawash J, Shain DH (2017) Punctuated invasion of water, ice, snow and terrestrial ecozones by segmented worms (Oligochaeta: Enchytraeidae: Mesenchytraeus). Proc. Roy. Soc. B 284, 20171081. http://dx.doi.org/10.1098/rspb.2017.1081
  62. Saidel WM, Saglam N, Salas-de la Cruz D, *Saunders R, Shain DH (2018) Elaborate Ultrastructure of the Hirudo (Annelida: Hirudinae) Cocoon Surface. J Morphology 279, 545-553.
  63. Saglam N, *Saunders R, *Lang SA, Shain DH (2018) Phylogeny and cocoon production in the parasitic leech Myzobdella lugubris Leidy, 1851 (Hirudinidae, Piscicolidae). Acta Parasitologica 63, 15-26.
  64. Gagliardi LJ, Shain DH(2018) Emergent mitotic chromosome motions from a changing intracellular pH. Open Journal of Biophysics 8, 1-13.
  65. Saglam N, Kutschera U, *Saunders R, Saidel WM, *Balombini KLW, Shain DH(2018) Morphologic and phylogenetic resolution of the Helobdella stagnaliscomplex (Annelida: Hirudinidae). Zootaxa 4403(1) 061-086. https://doi.org/10.11646/zootaxa.4403.1.3
  66. *Lang SA, Shain DH (2018) Atypical Evolution of the F1F0 ATP Synthase Regulatory ATP6subunit in Glacier Ice Worms (Annelida: Clitellata: Mesenchytraeus). Evolutionary Bioinformatics 14, 1-4. doi: 10.1177/1176934318788076
  67. Zawierucha K and Shain DH (2019) Disappearing Kilimanjaro snow – are we the last generation to explore equatorial glacier biodiversity? Ecology and Evolution 9, 8911–8918. DOI:10.1002/ece3.5327
  68. Kutschera U and Shain DH (2019) Hirudinea Lamarck 1818: Evolutionary origin and taxonomy of the six medicinal leeches (genus Hirudo) known today. Biomedical Research and Reviews 3, 1-4.
  69. Hotaling S, Shain DH, Lang SA, Bagley RK, Tronstad LM, Weisrock DW and Kelley JL (2019) Long-distance dispersal, ice sheet dynamics, and mountaintop isolation underlie the genetic structure of glacier ice worms. Proc. Roy. Soc. B 286, 20190983. http://dx.doi.org/10.1098/rspb.2019.0983
  70. Gagliardi LJ, Shain DH(2019) Electrostatic mechanism for depolymerization-based poleward force generation at kinetochores. Open Journal of Biophysics 9, 198-203.
  71. *Lang SA, McIlroy PJ, Shain DH (2020) Structural evolution of the glacier ice worm F1Fo ATP Synthase complex. The Protein Journal 39, 152-159. doi: 10.1007/s10930-020-09889-x
  72. Gagliardi LJ, Shain DH (2020) Electrostatic contributions to carcinogenesis. Open Journal of Biophysics 10, 27-45.
  73. Elser JJ, Wu C, González AL, Shain DH, Smith HJ, Sommaruga J, Williamson CE, Brahney J, Hotaling S, Vanderwall J, Yu J, Aizen V, Aizen E, Battin TJ, Camassa R, Feng X, Jiang H, Lu L, Qu JJ, Ren Z, Wen J, Wen L, Woods HA, Xiong X, Xu J, Yu G, Saros JE (2020) Rules of life and the fading cryosphere: impacts in alpine lakes and streams. Global Change Biology 26, 6644-6656.
  74. Zawierucha K, Porazinska DL, Ficetola GF, Ambrosini R, Baccolo G, Buda J, Ceballos JL, Devetter M, Dial R, Franzetti A, Fuglewicz U, Gielly L, Łokas E, Janko K, Jaromerska, Kościński A, Kozłowska A, Ono M, Parnikoza I, Pittino F, Poniecka E, Sommers P, Schmidt PK, Shain DH, Sikorska S, Uetake J, Takeuchi N (2020) A hole in the nematosphere: tardigrades and rotifers dominate the cryoconite hole environment, whereas nematodes are missing. Journal of Zoology313, 18-36.
  75. Saglam N, *Saunders R, Shain DH, Saidel WM (2020) Detailed ultrastructure of the Hirudosalivary gland. Micron 136. doi.org/10.1016/j.micron.2020.102887.
  76. Gagliardi LJ, Shain DH (2020) Biophysical mechanism for zinc as an anticancer agent. Medical Hypotheses 144, 110273.
  77. Shain DH, Novis P, Cridge A, Zawierucha K, Dearden PK (2021) Five animal phyla in glacier ice reveal unprecedented biodiversity in New Zealand’s Southern Alps. Scientific Reports 11, 3898 https://doi.org/10.1038/s41598-021-83256-3
  78. Shmakova L, Malavin S, Iakovenko N, Vishnivetskaya T, Shain D, Plewka M, Rivkina E (2021) A living bdelloid rotifer from 24,000-year-old Arctic permafrost. Current Biology, 31, R712-13. https://doi.org/10.1016/j.cub.2021.04.077
  79. Shain DH, Iakovenko N, Novis P, Cridge A, Plášek V, Dearden PK (2022) Microinvertebrate colonization of New Zealand’s thermally extreme environments. Evol. Biol. 49, 414-423
  80. Osorio T, Shain DH, Scoma ER, Melissaratos DS, Riggs LM, Hambardikar V, Solesio ME (2022) The Glacier Ice Worm, Mesenchytraeus solifugus, Elevates Mitochondrial Inorganic Polyphosphate (PolyP) Levels in Response to Stress. Biology 11, 1771.
  81. Zawierucha K, Stec D, Dearden PK, Shain DH (2023) Two New Tardigrade Genera from New Zealand’s Southern Alp Glaciers Display Morphological Stasis and Parallel Evolution. Molecular Phylogenetic & Evolution 178, 107634.
  82. Saglam N, *Melissaratos DA, Shain DH (2023) Biocontrol of Snail-Borne Parasites with the Glossiphoniid Leech, Helobdella austinensis. Biol. Lett. 19, 20220484. https://doi.org/10.1098/rsbl.2022.0484
  83. Shain DH, Irina Rogozhina, Naim Saglam, Atle Nesje, Jesamine Bartlett, Krzysztof Zawierucha, Einar Arnason, Jørgen Rosvold (2023) Glacial Rotifera Reveal Deep Metazoan Ancestries in the Nordic Cryosphere. (in revision).