Jun-Ru Lee and Dian-Han Kuo (2020). Netrin expressed by the ventral ectoderm lineage guides mesoderm migration in epibolic gastrulation of the leech. Developmental Biology 463, 39-52.

Dian-Han Kuo (2019). Comparative embryology as a way to understand evolution. In Old Questions and Young Approaches to Animal Evolution, (edited by José M. Martín-Durán and Bruno C. Vellutini), pp. 57-72. Cham: Spinger.

Dian-Han Kuo and Yi-Te Lai (2019). On the origin of leeches by evolution of development. Development, Growth and Differentiation 61, 43-57.

Dian-Han Kuo and Yu-Hsiang Hsiao (2018). Duplicated FoxA genes in the leech Helobdella: insights into the evolution of direct development in clitellate annelids. Developmental Dynamics 247, 763-778.

Dian-Han Kuo (2017). The polychaete-to-clitellate transition: an EvoDevo perspective. Developmental Biology 427, 230-240.

David A. Weisblat and Dian-Han Kuo (2014). Developmental biology of the leech Helobdella. International Journal of Developmental Biology 58, 429-433.


Ulrich Kutschera, Henning Langguth, Dian-Han Kuo, David A. Weisblat, and Marty Shankland (2013). Description of a new leech species from North America, Helobdella austinensis n. sp. (Hirudinea: Glossiphoniidae), with observations on its feeding behaviour. Zoosystematics and Evolution 89, 239-246.


Oleg Simakov, Ferdinand Marletaz, Sung-Jin Cho, Eric Edsinger-Gonzales, Paul Havlak, Uffe Hellsten, Dian-Han Kuo, Tomas Larsson, Jie Lv, Detlev Arendt, Robert Savage, Kazutoyo Osoegawa, Pieter de Jong, Jane Grimwood, Jarrod A. Chapman, Harris Shapiro, Andrea Aerts, Robert P. Otillar, Astrid Y. Terry, Jeffrey L. Boore, Igor V. Grigoriev, David R. Lindberg, Elaine C. Seaver, David A. Weisblat, Nicholas H. Putnam, and Daniel S. Rokhsar (2013). Insights into bilaterian evolution from three spiralian genomes. Nature 493, 526-531.


Dian-Han Kuo, Marty Shankland and David A. Weisblat (2012). Regional differences in BMP-dependence of dorsoventral patterning in the leech Helobdella. Developmental Biology 368, 86-94.


Dian-Han Kuo and David A. Weisblat (2011). Intermediate filament genes as differentiation markers in the leech Helobdella. Development, Genes and Evolution 221, 225-240. 


Dian-Han Kuo and David A. Weisblat (2011). A new molecular logic for BMP-mediated dorsoventral patterning in the leech Helobdella. Current Biology 21, 1282-1288.


David A. Weisblat and Dian-Han Kuo (2009). Helobdella (Leech): a model for developmental studies. In Emerging Model Organisms: a Laboratory Manual, pp. 245-267. Cold Spring Harbor, New York: Cold Spring Harbor Laboratory Press. (also a special section in Cold Spring Harbor Protocols)

Subsections Handling of Helobdella (leech) embryos. Microinjection of Helobdella (leech) embryos. Devitellinization of living Helobdella (leech) embryos. Arnolds' silver staining of Helbodella (Leech) embryos. Immunostaining Helobdella (leech) embryos. In situ hybridization of Helobdella (leech) embryos. Whole-mount preparation of Helobdella (leech) embryos for microscopy.

Stephanie E. Gline, Dian-Han Kuo, Alberto Stolfi and David A. Weisblat (2009). High resolution cell lineage tracing reveals developmental variability in leech. Developmental Dynamics 238, 3139-3151.


Shaobing O. Zhang, Dian-Han Kuo and David A. Weisblat (2009). Grandparental stem cells in leech segmentation: differences in CDC42 expression are correlated with an alternating pattern of blast cell fates. Developmental Biology 336, 112-121.


Dian-Han Kuo (2009). Annelids in evolutionary developmental biology. In Annelids in Modern Biology, (edited by Daniel H. Shain), pp. 65-87. Hoboken, New Jersey: Wiley-Blackwell.


Dian-Han Kuo and Marty Shankland (2004). Evolutionary diversification of specification mechanisms within the O/P equivalence group of the leech genus Helobdella. Development 131, 5859-5869.

Dian-Han Kuo and Marty Shankland (2004). A distinct patterning mechanism of the O and P cell fates in the development of the rostral segments of the leech Helobdella robusta: implication for the evolutionary dissociation of developmental pathway and morphological outcome. Development 131, 105-115.

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