Daniel Shain Research

Daniel H. Shain
Associate 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: Annelid development and evolution

Current Research Areas

  1. Cloning and expression of genes differentially expressed during stem cell formation.
  2. Genetic and ultrastructural analyses of leech cocoons
  3. Bioenergetics of ice worms and extremophilic microbiota.
  4. Ice worm phylogeny and phylogeography.

Selected Publications

*student author

Leech Development

  • *Hohenstein KA and Shain DH (2004) Changes in gene expression at the precursor -> stem cell transition in leech. Stem Cells (in press).
  • *Hohenstein KA, Canturk B* and Shain DH (2003) Genetic control of early development and stem cell formation in leech. Recent Res. Dev. Genet. 3, 17-30.
  • Shain DH, Stuart D and Weisblat DA (2000) Segmentation of the central nervous system in leech. Development 127, 735-744.
  • 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.

Leech Cocoons

  • *Mason TA, *Paulson T and Shain DH. Cocoon deposition and hatching in the aquatic leech, Theromyzon rude. (submitted).
  • *Dimitriu C and Shain DH (2004) Ultrastructural properties of the Theromyzon (Annelida: Hirudinea) cocoon. Micron (in press).
  • *Mason TA, McIlroy PJ and Shain DH (2004) A cysteine-rich protein in the Theromyzon (Annelida: Hirudinea) cocoon membrane. FEBS Lett. (in press).

Ice Worms

  • *Hohenstein KA and Shain DH. Divergence of F1 ATP synthase subunits in the ice worm, Mesenchytraeus solifugus. (submitted).
  • *Farrell AH, *Hohenstein KA and Shain DH. Molecular adaptation of the ice worm, Mesenchytraeus solifugus: divergence of energetic-associated genes (submitted).
  • *Napolitano MJ and Shain DH (2004) Four kingdoms on glacier ice: convergent energetic processes boost energy levels as temperatures fall. Biol. Lett. (in press).
  • *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.
  • Shain DH (2004) The ice worm’s secret? Alaska Park Science (in press).
  • Duchovnay B and Shain DH (2002) Worms on Ice. The Science Teacher 69, 20-25.
  • 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.
  • 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.


  • 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.
  • Shain DH, Zuber MX and Neuman T (1998) Transcription initiation from a poly d(A) tract. Nucl. Acids Res. 26(4), 1019-1025.
  • 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.