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 adaptation of glacier ice worms to the secretion of leech cocoons and production of embryonic stem cells. We also have an ongoing project exploring the adaptation of rotifers that occupy Icelandic glaciers and hot springs. 

Selected Publications

*student author

  • Marotta R and Shain DH (2007) Irregular helicoids in leech cocoon membranes.  Journal of Structural Biology 158, 336-343.
  • 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.
  • *Morrison BA and Shain DH (2008) An AMP nucleosidase gene knockout in Escherichia coli elevates intracellular ATP levels and increases cold tolerance. Biol. Lett. 4, 53-56.
  • *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.
  • *Tartaglia L and Shain DH (2008) Cold adapted tubulins in the ice worm, Mesenchytraeus solifugus. Gene 423, 135-141.
  • *Sayers CW, *Coleman J and Shain DH (2009) Cell dynamics during cocoon secretion in the aquatic leech Theromyzon tessulatum. Tissue & Cell  41, 35-42.
  • 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.
  • *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.
  • *Hohenstein KA, *Lang SA, *Nuthulaganti T and Shain DH (2010) A glutamine-rich factor affects stem cell genesis in leech. Stem Cellls International doi:10.4061/2010/145183
  • *Purzycki C and Shain DH (2010) Fungal toxins and multiple sclerosis: A compelling connection. Brain Res Bull. 82, 4-6.
  • *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.
  • Shain DH and Gagliardi LJ (2011) Can molecular cell biology explain chromosome motions? Theoretical Biology and Medical Modeling 8, 15.
  • *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.
  • 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.
  • *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.
  • Gagliardi LJ and Shain DH (2013) Is intracellular pH a clock for mitosis? Theortical Biology and Medical Modeling 10, 8-9.
  • *Rossi A, Marotta R, Saglam N, Shain DH (2013) Operculum ultrastructure in leech cocoons. Journal of Morphology 274, 940-946.
  • Gagliardi LJ and Shain DH (2014) Chromosome congression explained by nanoscale electrostatics. Theoretical Biology and Medical Modeling 11, 12-19.
  • Gagliardi LJ and Shain DH (2014) Polar electrostatic forces drive poleward chromosome motions. Cell Division 9, 191-198.
  • *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
  • Gagliardi LJ, Shain DH (2015) Intracellular pH as an electrostatic regulator of the spindle assembly checkpoint. Medical Research Archives 2, 12-18.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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
  • Gagliardi LJ, Shain DH (2016) Electrostatic forces drive poleward chromosome motions at kinetochore. Cell Division 11, 1-8.
  • *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.
  • *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
  • 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.
  • 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.
  • Gagliardi LJ, Shain DH (2018) Emergent mitotic chromosome motions from a changing intracellular pH. Open Journal of Biophysics 8, 1-13.
  • *Saunders R, Saglam N, Shain DH (2018) Morphologic and phylogenetic resolution of the Helobdella stagnalis complex (Annelida: Hirudinidae). Zootaxa (in press)
  • *Lang SA, Shain DH. Dynamic Evolution of the F1F0 ATP Synthase Regulatory ATP6 subunit in Glacier Ice Worms (Annelida: Clitellata: Mesenchytraeus). Evolutionary Bioinformatics (in press).