Post Doctoral Associate May 2012 to August 2015
Mentor: Karen Mesce
Department of Entomology
University of Minnesota, St. Paul, MN
Post Doctoral Scholar September 2009 to April 2012
Mentor: Daniel Wagenaar
Department of Biology
Broad Fellows Program in Brain Circuitry
California Institute of Technology, Pasadena, CA
Visiting Research Scholar Summer 2006
Mentor: Ansgar Büschges
Institute for Zoology at University of Cologne Germany
Mentors: Josef Schmitz, Volker Dürr
Department of Biological Cybernetics at University of Bielefeld.
PhD Biology Awarded August 2009
Mentor: Roy Ritzmann
Thesis title “Negotiation of Barriers by Intact and Brain-Lesioned Cockroaches”
Department of Biology
Case Western Reserve University, Cleveland, OH
B.S. Biology Awarded January 2002
Department of Biology
Case Western Reserve University, Cleveland, OH
Throughout my career I have been interested in how sensory information guides motor decisions. Despite their seeming simplicity, invertebrates are capable of complex behaviors. As with higher vertebrates, behavioral complexity depends on the brain. My thesis work focused specifically on the central complex, a region of the insect brain composed of multiple sub-regions. My work showed that each of these sub-regions had its own function in governing particular complex behaviors. Ultimately, my work has helped to illuminate how encoded sensory information passes from structure to structure eventually becoming transformed into motor commands. At the cellular level, how does sensory information morph into motor commands? This question led me to study the leech-- a system which has been used to identify neural circuits underlying many behaviors at the level of individual cells. Some of the best studied circuits are involved in controlling rhythmic behaviors such as swimming and crawling. Why would a leech choose one behavior over another? In my research, I found that during predatory behavior, leeches are biased toward crawling, and crawling makes them better able to localize pretty. In contrast, another study showed that when presented with UV light, a noxious stimulus, leeches swim away from its source. Armed with these robust behavioral responses and a suite of anatomical and physiological tools, I aim to examine, at the level of the neural network, how sensory information such as this is encoded and used to guide context-dependent locomotor decisions.
Courses designed and instructed:
Human Biology Fall 2017-Spring 2018
Department of Natural Sciences
Metropolitan State University
A human physiology course for undergraduate non-science majors
Introduction to Neurophysiology Spring 2017 to present
Department of Natural Sciences
Metropolitan State University
An undergraduate capstone course that teaches neuroscience from cells to complex behaviors
Comparative Physiology Spring 2016 to present
Department of Natural Sciences
Metropolitan State University
An undergraduate upper division course with lab which examines common ground plans of physiological systems and how they become adapted over evolutionary time to different conditions.
Human Anatomy and Physiology I and II Spring 2016 to present
Metropolitan State University
A two semester anatomy and physiology course (including lab) which serves to teach students desiring health related careers about the anatomy (form) and physiology (function) of the human body.
Sensory Neuroethology Fall 2015
Metropolitan State University
An undergraduate capstone course teaching neuroethology focusing on sensory systems and experimental methodologies.
Scientific Communication and Ethics Fall 2014
Department of Entomology
University of Minnesota
A graduate-level course about ethics and how to effectively present data in a variety of formats.
The Sensory Neuroethology of Courtship Spring 2012
Division of Biology
California Institute of Technology
A primary literature course focusing on how sensory systems are adapted for animal communication and exploited for predatory gains.
Courses instructed:
General Biology 2 Fall 2015
Department of Natural Sciences
Metropolitan State University
Biorobotics Spring 2008, Spring 2009
Graduate Teaching Assistant
Case Western Reserve University
Mentored and assisted students working within interdisciplinary groups of biologists and engineers on a semester long research project for this project-based biologically inspired robotics course
Organisms and Ecosystems (Physiology) Spring 2004, 2005, 2006, 2007
Graduate Teaching Assistant
Case Western Reserve University
Lectured, monitored, and graded for the undergraduate physiology lab
Neurobiology of Behavior Fall 2004
Graduate Teaching Assistant
Case Western Reserve University
Led discussions about impactful literature within the field of neuroethology and graded for the upper level course ‘neurobiology of behavior’
Invited lectures in other courses:
Insect Structure and Function December 2, 2013
Guest Lecture: “Insect locomotion”
Department of Entomology
University of Minnesota
Microbiology April 28, 2008 and April 27, 2009
Guest Lecture: “Beer: An Introduction”
Department of Biology
Case Western Reserve University
Harley, C. M. (2021) Leech locomotion: Invertebrate Physiology. Apple Press. In review
Nichols S., Kehl C. E., Taylor BK, Harley CM, (2020) "Bioinspired Navigation Based on Distributed Sensing in the Leech" Living Machines 2020
Harley CM, Edgerton, J., Auletta A (2020) "To Vaccinate or Not to Vaccinate: A Classroom Activity illustrating Herd Immunity" CourseSource
Auletta, A, Rue, M.C P, Harley, CM, Mesce, K. A. (2019) The Anatomical Distribution Pattern of Dopaminergic Neurons in the Central Nervous System of the Wolf Spider, Hogna lenta (Araneae: Lycosidae) J. Comp Neurol.
*Harley, CM, Asplen MA (2018) Annelid Vision. Oxford Encyclopedia of Neuroscience
* Thompson KJ, Harley CM, Barthel GM, Sanders MA, and Mesce,KA (2015) Plasmon Resonance and the imaging of metal-impregnanted neurons with the laser scanning confocal microscope. Elife 4:e09388
* Martin JP, Guo PG, Mu L, Harley, CM, Ritzman, RE (2015) Central-complex control of movement in the freely walking cockroach. Current Biology, 25: 2795-2803
Harley CM, Reilly, MG, Stewart C, Schlegel C, Morley E, Puhl JG, Nagel, C, Crisp, KM, Mesce KA (2015) Compensatory plasticity restores locomotion following chronic removal of descending projections. J. Neurophysiology, 113(10): 3610-3622
Harley CM, Wagenaar DA (2013) Scanning behavior in the leech Hirudo Verbana. PLOS ONE 9(1):e86120
Harley CM (2013) A win for science: The benefits of mentoring high school students in the lab. JUNE (Journal for Undergraduate Neuroscience Education) 12(1): E1-5
* Harley CM, Rossi M, Cienfuegos J, Wagenaar DA (2013) Discontinuous locomotion and prey sensing in the leech. Journal of Experimental Biology 216 (10): 1890-1897
Ritzmann RE, Harley CM, Daltorio KA, Tietz BR, Pollack AJ, Bender JA, Guo P, Horomanski AL, Kathman ND, Nieuwoudt C, Brown AE, Quinn RD (2012) Deciding which way to go: How do insects alter movements to negotiate barriers? Frontiers in Neuroscience, 6: 97.
* Harley CM, Cienfuegos J, Wagenaar DA (2011) Developmentally regulated multisensory integration for prey localization in the medicinal leech. Journal of Experimental Biology, 214(22): 3801-3807
Harley CM and Ritzmann RE (2010) Electrolytic lesions within central complex neuropils of the cockroach brain affect negotiation of barriers. Journal of Experimental Biology, 213(16): 2851-2864
* Harley CM, English BA, Ritzmann RE (2009) Characterization of obstacle negotiation behaviors in the cockroach, Blaberus discoidalis. Journal of Experimental Biology, 212(10): 1463-1476
Lewinger WA, Harley CM, Watson MS, Ritzmann RE, Branicky MS, Quinn RD (2008) Insect and animal-inspired sensing to enable autonomous mobile robot obstacle avoidance. Applied Bionics and Biomechanics, 6(1): 43-61
Lewinger WA, Harley CM, Ritzmann RE, Branicky MS, Quinn RD (2005) Insect-like antennal sensing for climbing and tunneling behavior in a biologically-inspired mobile robot. IEEE International Conference on Robotics and Automation (ICRA05), Barcelona.
† Co- first author
* Featured article
How cold weather effects the body an ebook written with the Visible Body team
Laboratory Activities written for Visible Body’s Human Anatomy Atlas
Special senses: vision
Special senses: the ear
Special senses: smell and taste
Laboratory Activities written for Visible Body’s Anatomy and Physiology App
Special senses: system primer
Special senses: taste and smell
Special senses: vision
Special senses: the ear
I also was responsible for the anatomical markers for Broadmann's areas in Visible Body's 3D brain model.
C.M. Harley “Active Learning in 3D” Visible Body, Newton Mass. May 2021
C.M. Harley “The Neuroscience of Resilience” Metropolitan State University April 2021
C.M. Harley “The Separable Roles of Motivation and Recovery” UNC/Duke November 2021
C.M. Harley “When it comes to the nervous system, leeches don’t make sucky teachers” Institute de Fisica, Federal University of Rio October 2021
C.M. Harley “Circulatory and Immune systems of Invertebrates” Minnesota Virtual Academy Feb 2021
C.M. Harley “The History of Studying Viruses” Minnesota Virtual Academy Feb 2021.
Post hoc recording here: https://youtu.be/sJ3kxnvzpE0
C.M. Harley "Visible Body Office Hours" Public webinars for Visible Body. July 2020, August 2020
C.M. Harley "Putting pieces together: How breaking something can teach you about it" University of Oklahoma June 2020
C.M. Harley "At the heart of Adaptation" Visible Body Webinar March 2020
C.M. Harley “How Hirudo got its groove back: examining the role of physical therapy in recovery following nerve cord injury" Society for Experimental Biology 2019
C.M. Harley “Learning: A Neural Perspective” Brain Awareness Week, Metropolitan State University 2018
C.M. Harley “Textbooks are not the only option—teaching Anatomy and Physiology using an App” STAR Symposium February 2018
C.M. Harley “The eyes have it: The visual guidance of host localization and escape in the medicinal leech” JB Johnston Club Annual Meeting. Washington, D.C., November 2014
C.M. Harley “Small minds, Big World: Learning from invertebrate navigation” Brain awareness week. St. Olaf College. Northfield MN October 2013
C.M. Harley “Trick or Treat: How leeches localize their prey” Smith College. Northampton, MA October 2013
C.M. Harley “Closing multiple loops: How sensory processing influences locomotor behavior and vice versa” Gordon Research Seminar: Closing the Loop. West Dover, VT August 2013
C.M. Harley “Crawl vs Swim: How leeches decide between two competing motor programs” University of Minnesota. St. Paul, MN April 2013
C.M. Harley and K.A. Mesce "You can’t always get what you want, but if you crawl sometimes, you might just find, you get what you need". Wright State University. Dayton, OH March 2013
C.M. Harley "Beyond the pest: what cockroaches can tell us about neural control of complex behavior". University of Minnesota. St. Paul, MN November 2012
C.M. Harley M. Rossi, J. Cienfuegos, K.A. Mesce and D. A. Wagenaar "To swim or to crawl: the effect of different locomotor states on prey Localization". JB Johnston Club Annual Meeting. New Orleans, LA October 2012
C.M. Harley and D.A. Wagenaar "Finding the right food: Developmental regulation of prey sensing". Flow Sensing in Air and Water. Bonn, Germany July 2011
C.M. Harley and R.E. Ritzmann “Why do cockroaches have brains?” Case Western Reserve University. Cleveland, OH August 2009
C.M. Harley and R.E. Ritzmann “Using your head: the involvement of the cockroach brain and head sensors in obstacle negotiation behaviors” California Institute of Technology. Pasadena, CA February 2009
C.M. Harley and R.E. Ritzmann “If I only had a brain: the involvement of the cockroach brain and head sensors in obstacle negotiation behaviors” University of Maryland Baltimore County. Baltimore, MD November 2008
C.M. Harley and R.E. Ritzmann “Using behavioral analysis to diagnose deficits in insects with discrete lesions of the central body complex” Insect Behavior: Small Brains, Big Functions, Janelia Farm. Washington, D.C. March 2007
C.M. Harley and R.E. Ritzmann “The role of head sensors in the avoidance of complex obstacles” Universität zu Köln. Cologne, Germany June 2006
C.M. Harley and R.E. Ritzmann “The role of head sensors in the avoidance of complex obstacles” University of Bielefeld. Bielefeld, Germany July 2006
C.M. Harley and R.E. Ritzmann “Complex decisions about simple obstacles: cockroach climbing and tunneling” East Coast Nerve Net. Woods Hole, MA April 2006
C.M. Harley and R.E. Ritzmann “How cockroaches use head sensors to navigate complex environments” Case Western Reserve University. Cleveland, OH March 2006
C.M. Harley and R.E. Ritzmann “Investigating the role of motion senstitive cells in mantis strike” Case Western Reserve University. Cleveland, OH April 2005. (as part of qualifying exam)
C.M. Harley "The Impact of Visible Body use on Student Performance and Engagement in Anatomy and Physiology Classes" (2021) SABER
C.M. Harley "The Effectiveness of Visible Body for Teaching A&P" (2021) HAPS
McDuffee Altekreus A*; Harley, CM; Li. J Edon JA Zemke, D More is not necessarily better: Physical therapy differentially influences crawl quality and quantity in Hirudo verbana SICB 2020 Austin TX
Kelly Sheehy, Emiliya Bayko, Elizabeth Baseka-Nemmeh, Andrew Clayburn, Brandon Young, C.M. Harley (2018) Crawling Back to you: A Study of Leeches and Light. Midbrains 2018
C.M. Harley (2018) “Homeostasis: The Game”— Using a game to teach the relationship between the endocrine system and homeostasis. SABER Minneapolis MN July 2018
C. Yoder, B. Chi, C.M.Harley (2017) “In search of the leech optimotor response” Student Scholars Syposium St Paul MN, April 2018
C. Yoder, B. Chi, C.M.Harley (2017) “In search of the leech optimotor response” Midbrains St Paul MN, Nov 2017
A. Auletta, C.M.Harley , M. CP Rue, K.A. Mesce (2016) The Organization of the Spider Nervous System as Revealed by Dopamine Immunolabeling and Confocal Microscopy. International Society of Arachnology. Colorado.
A. Auletta, C.M.Harley , M. CP Rue, K.A. Mesce (2016) The Organization of the Spider Nervous System as Revealed by Dopamine Immunolabeling and Confocal Microscopy. Entomological Society of America Minneapolis MN
A. Auletta, C.M. Harley, K.A. Mesce (2015) Journey to the Center of the Spider: The First Immunocytochemical Analysis of Dopamine in the CNS of a Spider & Possible Behavioral Correlates Gordon Research Seminar. Lucca Italy
C. M. Harley, M. Reilly, C. Stewart, E. Morley, M. Songpitak, S. Hadir, K. Crisp, K.A. Mesce (2014) Homeostatic plasticity directed by physical therapy facilitates locomotor recovery after removal of cephalic inputs. Soc. Neurosci. Abstr. Washington, D.C.
K.J. Thompson, C. M. Harley, M. A. Sanders, K.A. Mesce (2014) Imaging silver-impregnated histological preparations with confocal microscopy. Soc. Neurosci. Abstr. Washington, D.C.
C. M. Harley, M. Reilly, C. Stewart, E. Morley, M. Songpitak, S. Hadir, K. Crisp, K.A. Mesce (2014) Compensatory plasticity directed by physical therapy facilitates locomotor recovery after removal of cephalic inputs. Intl. Soc. Neuroethology. Abstr. Sapporo, Japan
C. M. Harley, K.J. Thompson, M. A. Sanders, K.A. Mesce (2014) Seeing a new silver lining: Imaging silver-impregnated histological preparations with confocal microscopy. Intl. Soc. Neuroethology. Abstr. Sapporo, Japan
C. M. Harley, M. Reilly, C. Schlegel, S.M.A.J. Haidir, K.A. Mesce (2013) Physical therapy in an invertebrate model facilitates locomotor recovery after nerve cord injury. Soc Neurosci Abstr. San Diego, CA
C.M. Harley, K.A. Mesce (2013) Visual Processing in the medicinal leech: More than meets eye. Soc Neurosci Abstr. San Diego, CA
C.M. Harley Closing multiple loops: How sensory processing influences locomotor behavior and vice versa. Gordon Research Conference: Closing the Loop. West Dover, VT August 2013
K.A. Mesce, C. M. Harley, A. J. Shafqat, K. M. Crisp, M. Reilly, E. Morley, C. Schlegel, C. W. Nagel, J.G. Puhl (2012) Organization of dopamine regulated locomotor behavior in the medicinal leech. Soc Neurosci Abstr. New Orleans, LA
C. M. Harley, M. Rossi, J. Cienfuegos, D. A.Wagenaar, K. A. Mesce (2012) Sensory guidance of locomotor behavior in the medicinal leech. Soc Neurosci Abstr. New Orleans, LA
K. A. Mesce, C. M. Harley, K. M. Crisp, A. J.Shafqat, C. W. Nagel, J. G. Puhl (2012) Steps and strides in understanding how a dopamine regulated locomotor behavior is organized in the medicinal leech. Intl. Soc. Neuroethology. Abstr. College Park, MD
C.M. Harley, M. Rossi, J. Cienfuegos, K. A. Mesce, D. A.Wagenaar (2012) To see or not to see: integration of sensory information in the medicinal leech is dependent on mode of locomotion. Intl. Soc. Neuroethology. Abstr. College Park, MD
J. P. Martin, L. Mu, C. M. Harley, M. Kesavan and R. E. Ritzmann(2012) Origin of descending inputs controlling reflex reversals in the cockroach (Blaberus discoidalis). Intl. Soc. Neuroethology. Abstr.College Park, MD
C.M. Harley, J. Nagarah, J. Cienfuegos, M. Rossi, D.A. Wagenaar (2011) Multisensory integration underlying host detection in the medicinal leech. Soc Neurosci Abstr. Washington, D.C.
C.M. Harley, J. Cienfuegos, D.A. Wagenaar (2010) Frequency dependence of responses to prey in the leech Hirudo medicinalis during different life stages. Soc Neurosci Abstr. San Diego, CA
C.M. Harley, J. Cienfuegos, D.A. Wagenaar (2010) Frequency dependence of prey localization responses in the leech Hirudo medicinalis during different life stages. Intl. Soc. Neuroethology Abstr. Salamanca, Spain
C.M. Harley, L. Mu, R.E. Ritzmann (2009) Lesions within specific cockroach brain regions reveal control for obstacle avoidance. Soc. Neurosci. Abstr. Chicago, IL
C.M.Harley, S.J. Awe, L. Mu, J.A. Bender, R.E. Ritzmann (2008) Discrete lesions within the cockroach brain affect obstacle negotiation behaviors. Soc. Neurosci. Abstr. Washington, D.C.
C.M. Harley, B. A. English, R.E. Ritzmann (2008) Using your head: the involvement of the cockroach brain and head sensors in obstacle negotiation behaviors. Adaptive Motion of Animals and Machines Cleveland, OH.
C.M. Harley, B. A. English, R.E. Ritzmann (2007) Discrete lesions within the cockroach brain and their effect on negotiation of barriers. Kleinztiern Konferenz Tutzing, Germany
C.M. Harley, B. A. English, A. Pollack, R.E. Ritzmann (2007) Discrete lesions within the cockroach brain and their effect on negotiation of barriers. Intl. Soc. Neuroethology Abstr. Vancouver, Canada
C.M. Harley, J. Predina, R.E. Ritzmann (2006) Responses to incomplete sensory information in cockroach climbing behavior. Soc. Neurosci. Abstr. Atlanta, GA
C.M. Harley, W.A. Lewinger, R.E. Ritzmann, R.D. Quinn (2006) Characterization of obstacle avoidance behaviors in the cockroach Blaberus discoidalis and implementation in a semi-autonomous robot. Research ShowCase. Cleveland, OH
C.M. Harley, W.A. Lewinger, R.E. Ritzmann, R.D. Quinn (2005) Characterization of obstacle avoidance behaviors in the cockroach Blaberus discoidalis and implementation in a semi-autonomous robot. Soc. Neurosci. Abstr. Washington, D.C.
C.M. Harley, W.A. Lewinger, R.E. Ritzmann, R.D. Quinn (2005) High-speed video and kinematic analysis of transitional behaviors in the cockroach. Research ShowCase Cleveland, OH
R.E. Ritzmann, C.M. Rice*, A.J. Pollack, A.L. Ridgel, D.A. Kingsley, and R.D. Quinn (2001) Roles of descending control in locomotion through complex terrain. Intl. Soc. Neuroethology Abstr. Bonn, Germany, *maiden name
Carol C. Ryan Excellence in Advising Award Spring 2020
CourseSource Scholarship July 2018
Scholarship to attend course source workshop
ADInstruments Crawfly Scholarship July 2017
Scholarship to attend the crawfly workshop in Ithaca, NY
Professional Development Grant Multiple occasions
Metropolitan State University
International Congress of Neuroethology
Elected to general council August 2013
Company of Biologists
Direct Travel Award August 2010
Travel Award to attend the Society for Neuroethology meeting in Salamanca Spain
Society for Neuroscience: Women in Neuroscience
Graduate Travel Fellowship November 2009
Travel Award to attend the Society for Neuroscience meeting in Chicago, I.L.
Company of Biologists
Direct Travel Award November 2008
Travel Award to attend the Society for Neuroscience meeting in Washington, D.C.
Case Western Reserve University
Endowment Sponsored Mentorship Program October 2007
Travel Award to attend the Klienztiern Konferenz in Tutzing, Germany
International Congress of Neuroethology
Congress Award July 2007
Travel award to attend the Society for Neuroethology meeting in Vancouver, Canada
IEEE
Best Video at International Conference on Robotics and Automation (ICRA ’05) Won for the video submission of:
Lewinger, W.A., Harley, C.M., Ritzmann, R.E., Branicky, M.S., Quinn, R.D., (2005) Insect-like Antennal Sensing for Climbing and Tunneling Behavior in a
Biologically-inspired Mobile Robot. IEEE International Conference on Robotics and Automation (ICRA05), Barcelona, Spain.
National Science Foundation
IGERT Fellowship June 2003—May 2006
Received an interdisciplinary fellowship to engage in graduate studies at CWRU
Howard Hughes Medical Institute
Spur Fellowship May 2001—August 2001
Received a grant to continue undergraduate sensory systems research in Ritzmann lab
Funding acquired for additional coursework
University of Minnesota Spring 2014
Regents scholarship received to take Advanced methods in microscopy
Case Western Reserve University Spring 2008
Fellowship course funds received to take German language
Khari Thury Spring 2020
Annie Foster Spring 2020
Robin Johnson Fall 2019-Spring 2020
Niera Belgic Fall 2019
Alyssa McDuffee Altekreuse Summer 2019-Spring 2020
Jing Li Summer 2019
Adefunke Edon Spring 2019
Darlene Zemke Spring 2019
Heather Harrington Spring 2019
Andra Sonnek Spring 2019
Arabella Jones Spring 2019
Kelly Sheehy Fall 2018 to present
Elizabeth Baseka Nemmeh Fall 2018
Andrew Clayburn Summer 2018 to Spring 2019
Emiliya Bayko Summer 2018 to present
Alisha Anderson Spring 2018
Brandon Young Spring 2018 to Spring 2019
Marissa Peterson Fall 2017 to present
Alison Aguilar Fall 2017
Hannah Lee Fall 2017
Brian Chi Fall 2017
Cynthia Yoder Summer 2017 to Spring 2017
Sundus Yusef Summer 2017
Benjamin Kuter Spring 2017
Chee Xiong Summer 2016
Jasmine Engler Summer 2016
Tyler Pinkston Summer 2016
Linna Ahmado Summer 2016
Colleen Smith Summer 2014 to Summer 2016
Melissa Songpitak Summer 2013
Syed “Jabbar” Haidir Fall 2012- Spring 2014
Melissa Reilly Summer 2012-Spring 2013
Chantel Schlegel Summer 2012-Fall 2013
Emma Morely Summer 2012, Summer 2013
* Matt Rossi Spring 2011- Spring 2012
* Javier Cienfuegos Spring 2010- Fall 2011
Amy Brown Spring 2009-Summer 2009
Solomon Awe Spring 2008-Summer 2009
Nicholas Courtney Spring 2008
Sharanya Sivikumar Spring 2008
Brittany English Fall 2006-December 2007
Jarrod Predina Summer 2006
* Denotes high school students