The Path to Engagement Faculty spotlight

fellows by the numbers

CONTENTS

Introduction Letter

To the B&B Community Welcome to the Spring 2016 issue of The Potential, featuring community education. Georgia State University boasts an especially vibrant neuroscience research and training environment, thanks to the amazing contributions of faculty members, staff, students, and community partners. Not only do we engage in cutting-edge research in a wide variety of basic and applied subfields of neuroscience, but also we support strong professional development opportunities and critical community education endeavors. The B&B Program is essential to all of these, but focuses on community education in this issue of The Potential. Spurred by long-standing science education initiatives such as the BioBus and the entire education arm of the Center for Behavioral Neuroscience, Georgia State neuroscientists offer numerous K-12, college, and public education programs. For example, we reach broad audiences in our K-12 Classroom Visits Program with the Atlanta Chapter of the Society for Neuroscience, as well as through the new Atlanta Science Festival, where our volunteers translate their research findings into lesson plans for novices or non-scientists. Combined, these reach at least 5,000 learners of all ages and stages each year. We also target elite populations to come learn neuroscience and conduct research with us through the Institute On Neuroscience (ION) for Outstanding High School Students and Teachers, as well as Atlanta’s Neuroscience Education and Training (NET/work) Program. All of our programs emphasize diversity by highlighting the relevance and accessibility of neuroscience to individuals from diverse backgrounds. Participant feedback suggests that our education and training programs pique curiosity, advance skills and knowledge, elevate positive attitudes about science at Georgia State, and even launch careers. Moreover, our education research reveals that these science education programs are beneficial for our volunteer instructors as well, e.g. volunteers learn to modify their teaching styles on the fly, responding immediately to learner feedback regarding vocabulary levels, concepts of interest, and opportunities to expand on basic lesson plans. Such experience builds our volunteers’ confidence in teaching, diversifies their styles of scientific communication, and achieves goals in community service. Such healthy integrations of research, professional development, and community service help to make our undergraduate, graduate, post-doctoral, and even faculty and staff training environments very lively and robust.

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Faculty Spotlight on Joe Normandin

5

Fellows by the numbers

7

Path to Engagement

9

Geert de Vries, Ph.D. Neuroscience Institute Director [email protected] Elizabeth Weaver, M.S. Associate Director [email protected] Rob Poh IT Director [email protected] Tara Alexander, M.S. Business Manager [email protected] Chriselle Manabat Creative Assistant

[email protected]

Elliott Albers, Ph.D. Neuroscience Institute [email protected] Deborah Baro, Ph.D. Biology [email protected] Sarah Brosnan, Ph.D. Psychology [email protected] Andrea Scarantino, Ph.D. Philosophy Chair [email protected] Walt Wilczynski, Ph.D. Neuroscience Institute [email protected]

15 B&B AMBASSADORS

Sincerely yours, Kyle Frantz Professor, Neuroscience Institute Senior Faculty Associate for Special Programs

Distinguished Lecture Series

INTERDISCIPLINARY COMMITTEE

Keep up the great work, and don’t hesitate to contact me to volunteer!

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Grad Student Op-Ed: Different Shades of Neuronal Subtype Diversity

ADMINISTRATORS

Hands-on, real-world education and training is integral to the age-old practice of experiential learning, which is a concept of increasing emphasis throughout Georgia State. Faculty and staff members all over campus value active teaching and learning, integrated classroom and professional learning environments, and career preparation. As university graduation rates climb at impressive rates, more and more students are reaching the workforce equipped with the professional skills and dispositions they need in order to succeed. Prime examples of undergraduate involvement in pre-professional and community endeavors exist in every college, such as research assistantships and co-operative education programs in the College of Arts and Sciences, student teaching in the College of Education and Human Development, clinical rotations in the School of Nursing and Health Professions, government internships in the School of Policy Studies, and client-based market research in the College of Business. Some of these opportunities are encapsulated in courses known as “Signature Experiences”, whereas others are facilitated through offices such as Career Services, Study Abroad, and the Office of Civic Engagement. It has been an honor for me in particular to support the enhancement and tracking of these opportunities as the Senior Faculty Associate for Special Programs, in the office of the Vice Provost, Dr. Timothy Renick.

Recent & Upcoming events

Igor Belykh, Ph.D. Math & Statistics [email protected]

Tricia King, Ph.D. Psychology [email protected]

Richard Briggs, Ph.D. Physics & Astronomy [email protected]

Vincent Rehder, Ph.D. Biology [email protected]

Nancy Forger, Ph.D. Neuroscience Institute [email protected]

Daniel Weiskopf, Ph.D. Philosophy [email protected]

Markus Germann, Ph.D. Chemistry [email protected]

Ying Zhu, Ph.D. Computer Science [email protected]

Editor in Chief: Elizabeth Weaver Design & Layout: Chriselle Manabat

r ec ent & UPC OMING EVENTS See pg. 3

2015 -2016

Walk to End Alzheimers

SEPTEMBER

Professional Development Interview Prep Workshop

NOVEMBER

OCTOBER

Society for Neuroscience Conference

Distinguished Lecture Series

Robert Williams, Ph.D. will talk about “Relational Leadership & Emotional Intelligence” on March 25th at 11:30am EST.

The 2016 B&B Retreat will feature senior Fellow talks, a keynote, and a poster session.

Open to staff, faculty, & students. RSVP only to Liz Weaver at [email protected] Professional Development

RSVP: Chriselle Manabat at mmanabat1@ student.gsu.edu

Workshop

APRIL 22

MARCH 25

JANUARY

MARCH 23

B&B Retreat

APRIL 4

Seed Grant Deadline

Call for Undergraduate Scholars Call for Seed Grant Applications went out in January. The deadline for Seed Grant Applications is Wednesday, March 23rd at 5:00pm EST

Scholars will engage in professional development workshops throughout the summer and will present their research at the end of the summer at a poster symposium.

THE POTENTIAL

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Distinguished Lecture Speakers

Distinguished Lecture Series

1/22

Eve Marder, Ph.D. Brandeis University Title: “Variability, modulation and homeostasis in neurons and networks.” Host: Andrey Shilnikov, Ph.D.

One of the ways the Brains & Behavior Program, otherwise known as B&B, promotes

2/12

research collaborations and dialog is through the Distinguished Lecture Series. These

Ned Block, Ph.D. New York University Title: “Is there a joint in nature between cognition and perception?”

events are held every second Friday of each month and are open to the public.

3/11

Host: Dan Weiskopf, Ph.D. Catherine Carr, Ph.D. University of Maryland Title: “Maps and meters for sound localization.” Host: Paul Katz, Ph.D.

4/8

Alcino Silva, Ph.D. University of California, Los Angeles Title: “Molecular, cellular and systems allocation mechanisms that link memories across time.” Host: Marise Parent, Ph.D

5/13 Greg Wray, Ph.D. Duke University Title: “Evolving a bigger brain: not so easy.” Host: Paul Katz, Ph.D.

Spring 2016

THE POTENTIAL

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faculty Spotlight on joe normandin

Thank you so much Joe for being our faculty spotlight for this issue of the Potential. You received your PhD in the Neurobiology program here. Now having become a teacher here in the last 2 years, you seem to be thriving. What made you decide to come back to GSU as a teacher and how has that transition been for you? I received my Ph.D. in the Neurobiology & Behavior program in the Biology Department in 2010 and went on to a post-doctoral position at the Netherlands Institute for Neuroscience in Amsterdam. While I had fantastic collaborators and loved the opportunity to live abroad, I discovered that an intense research-focused environment really didn’t drive me in the way I thought it would. I returned to the U.S., and I honestly felt a little lost. The one thing that kept coming back to me was how much I enjoyed my teaching experiences as a graduate student. So, I started to look at college-level teaching positions in biology, psychology, and neuroscience. I felt I had the breadth and depth of knowledge to teach almost any class within those disciplines, and I was lucky to find that GSU’s “new” Neuroscience Institute was looking for a Visiting Lecturer. It was very odd interviewing with quite a few people at GSU who knew me well. After my first telephone interview, I thought there was no chance I was going to get the job. I was nervous, and felt like every answer sounded canned. To my surprise, I got a second interview, with a mock lecture, and I think that was what won the committee over. I went into this interview/mock lecture and pretended we were in my classroom (even chastising Dr. Murphy for looking at her cell phone during my lecture). I felt confident, I was

Spring 2016

You seem to have a passion for community engagement. What led to your enthusiasm for this kind of advocacy? This is going to sound self-aggrandizing and a bit sappy, but it’s all true. I was a smart kid, and I’ve always been rather impatient. My mom had to constantly reign me in and she was always telling me that my intelligence needed to be used to help people and not hurt them by being impatient and belittling. I think that’s where it all began; this idea that I could use my own strengths to help other people. It really aligns with the Catholic faith I grew up with (even though I’m not religious now). I’ve come to the realization that I’m not so special…yeah, intelligent, but weak in other ways. So, I do what I can to inspire and teach others using my strengths, and rely on others to help me where I have challenges. Well… ideally, but perhaps not always perfectly. As I grew up, I used my confidence to help other young gay kids feel comfortable in their own skin by doing a lot of public speaking on what it was like to be a gay person. I did speaking engagements all over Massachusetts, where I grew up. It was basically a crashcourse in public speaking, and has left me feeling generally comfortable in front of a crowd. I also think I may have saved some lives just by telling my story, and it taught me the power of reaching out to people. During grad school at GSU, I met Dr. Kyle Frantz and Dr. Laura Carruth, and volunteered for many of the public science education outreach programs they coordinated. I remember being recruited for a particularly challenging outreach with a class of young kids with behavioral problems. It was a challenge, but one of the most rewarding experiences I’ve had in teaching. The kids were so eager to learn, and so bright, they just needed some extra help to get where they wanted to go. This volunteer work with Kyle and Laura helped me to teach science to so many people from so many walks of life. These experiences not only have tremendous personal value for me, but I also believe that scientific literacy is critical to a future where we can make reasoned policy decisions for our society. What community projects are in the works for you right now?

well-prepared, and I channeled all the best instructors I had ever had. Ever since then, I’ve tried to approach each class the same way and with the same energy. I’ve heard, until that mock lecture, I was not the committee’s first choice, so I guess I did something right. I’ve been teaching in the NI for three years now and this year was hired as a Lecturer within the department. This has given me a bit more job security, along with more responsibility in helping the other faculty to run the NI. In fact, I was entrusted with creating a new course, Introduction to Neuroscience, to meet our student’s need to get their feet wet early, and to expand on what our department offers to any student at GSU (there are no prerequisites to the course). I was honored that my colleagues had faith in my expertise to create such a course. Right from the beginning the other faculty treated me with the same respect they would treat any other faculty member; I never felt like I was still their “student.” Grad school, and the classes I took with some of the NI faculty, seem so very far away. That being said, the lessons I learned years ago from some who are now my colleagues, permeate my classes. One thing that comes to mind is that I owe Dr. Paul Katz gratitude for a tough lesson on the scientific method, hypothesis generation, and the difference between a hypothesis and a prediction. But that’s another story.

I’m currently working on a big project as chair of the Atlanta Science Festival at GSU. We’ve been a big part of the Atlanta Science Festival since the first festival two years ago. This year, we know what has worked and what has not worked, so we are focusing our energy to make what’s been great in the past into something even more fantastic. I really hope everyone comes out for our Discovery Day, March 19th, at the Petit Science Center! There is something for everyone and every age group. It’s a big responsibility and Dr. Frantz and Dr. Amy Horner-Reber (Biology) are instrumental in putting this together with me as the Chair Emerita and CoChair, respectively. I’ve also been doing a few smaller things for the NI and the University. I organized our annual NI Trivia Night last fall, for example. In addition, I had the highest-rated mock college lecture at the Panther Previews sessions (a recruitment event for high school students) last fall (TYVM)! Outside of work, I’m also about to start volunteering with an organization called Lost & Found, a shelter for homeless GLBT youth in Atlanta. They are working on setting up a GED program, and I’d like to help them as a tutor. Depending on demand, there may be some opportunity for any of us here at the NI to help out!

Recently, in a couple of your classes, you have initiated a way for students to earn extra credit by participating in art projects. What kind of art do neuroscience students come up with and how do you believe this is helping them understand the concepts you are teaching? I’ve always been a fan of modern and contemporary art and design, but I have no formal education or training in any of this. I love looking at new things in new ways. I love when my mind is challenged. These are also student experiences that are inherent in the two courses I currently teach (Introduction to Neuroscience and Principles of Neuroscience). Students begin to see that our nervous system is everything we are, in all its complexity, and it’s beautiful. So, this art project begins there, when a student is inspired by something in class. Sometimes it’s something that just seems “cool,” sometimes it’s a more personal revelation. Either way, I wanted to try to capture these moments. The consequence is that students must think deeply about this moment of inspiration, and learn more about the scientific concept. Initially, I help students create a framework so that the work is “significant” and has depth, then they are on their own. Lastly, we work together to explain the neuroscience concept expressed in their work as part of a “gallery card.” I’ve been very impressed with the work thus far, and it’s obvious that each student needed to think deeply about themselves, and their neuroscience concept, to create the work. Come see all their work in my office (PSC 801)! You seem to be thinking and teaching outside the box. What is next on the horizon for Dr. Normandin and his students? I have a number of things bouncing around my head. Poor Dr. Geert De Vries, the NI Director, has to endure hearing these ideas every time I see him. For Principles of Neuroscience, I want to create a more immersive neuroanatomy lesson that makes use of some 3-D scanning technology, a plastinated human head I have access to, and the Curve video wall in Library South. This will augment the sheep brain dissection that we already do. It’s a big project, so I hope I find the time to work on it soon. I’m also thinking into the future, and I wonder if Introduction to Neuroscience might not be a great course for the NI to offer the world as an online course. As part of developing that idea, I’m going to be recording all of my lectures this semester in the new Active Learning Classroom (CS 226) that I’m using for one of my sections. I also want to show my Intro. Neuro. students that science and scientists are accessible, so I hope to skype-in a neuroscientist whose work we are reviewing. As for the NI itself, I want to expand a bit on making connections within our department. For example, post-docs get lost in limbo as they strive for research success, but they need other experiences as well. Maybe post-docs would benefit from a mentorship program for neuro majors that need some extra attention? I haven’t floated that idea to Geert yet, so hopefully he will read this and say “ya” (that’s Dutch for yes)

Biology

5

7

Philosophy

4

4

Chemistry

6

Computer Science

3

Math

DEPARTMENT BREAKDOWN

Psychology

Neuroscience

15

24% of Fellows used their travel award to present their research nationally & internationally

4

Physics

48 TOTAL FELLOWS

Total of 69 Presentations

In 2014, 73% of fellows presented

In 2014, 58% of fellows published

Total of 51 Publications

FELLOWS BY THE NUMBERS

Path to Engagement

285

chastain park

kittredge magnet school The galloway school

Your Brain On Awareness

Ga Aquarium

Decatur high School

78

CENTENNIAL OLYMPIC PARK

mlk historic site

Morehouse School of Medicine

20

chapel hill middle school

Grant park

85 zoo atlanta

west end marta station Turner Field hartsfield Jackson airport westview cemetery

civil war museum

THE POTENTIAL

YOUR BRAIN ON AWARENESS

See below for a couple of the programs that make up the past and present fabric of our unique community: • The DANA Foundation supported Lending Library, where people can check out brain models and educational gear is a staple program that allows our educators to go into K-12 classrooms fully prepared.

by Liz Weaver

LEANING IN

#1

Katie Partrick, second year Neuroscience graduate student, sits across from me with a smile from ear to ear. Enthusiasm might as well be her middle name, because she has a lot of it. Energy and enthusiasm are two really good qualities when you have what Katie has ahead of her. As of a couple months ago, Katie has taken on two big roles: the chair of community engagement for the Neuroscience Graduate Student Association and the Georgia State University representative at the Atlanta Chapter of the Society for Neuroscience (ACSfN). In the following year, Katie will lead the charge, with an army of dedicated volunteers, and organize what our neuroscience department has become well known for: 52 weeks of Brain Awareness. With classroom visits, science fair visits, and STEM nights, in addition to collaborating with other Atlanta institutions to provide neuroscience education to the public (see side box #1), you can begin to sense what a substantial undertaking this will be for Katie. “It definitely takes time and effort. However, the benefit of sharing what you are passionate about and seeing it be received through gratitude and excitement over the brain, the benefits far outweigh the costs. Not to mention, there is constant motivation from a great team of supportive people.” But Katie isn’t the first to hold positions such as the ones above. There have been a stellar list of science educators before her that laid the foundation for what is today one of the nation’s most active neuroscience outreach communities.

A HISTORY OF DEDICATION In 2000, prior to the formation of the Neuroscience Institute, the Center for Behavioral Neuroscience (CBN, a state wide consortium of neuroscientists from Emory University, Georgia Tech, Morehouse College & School of Medicine, Spelman and Georgia State University) was hiring faculty in a multitude of areas. Two new hires here at Georgia State University (the lead institution), Drs. Kyle Frantz and Laura Carruth, were hired both as researchers in their specific areas as well science educators. After all, one of the core missions of the CBN then, and still today, is to be an advocate for and leader in science education. To that mission, Drs. Frantz and Carruth vigorously wrote science education grants. With the invaluable help from organizations such as the Dana Foundation (see side box #2) and hundreds of volunteers of faculty and students, GSU built a science education empire of sorts. From undergraduate & high school research

programs to Spring 2016

• Neuroscience grad student and B&B Fellow, Kat Sturhman worked with Fernbank science museum for their “Science at Hand Day”.

• The Institute on Neuroscience (ION), a summer research experience for high schoolers, has grown and now has a teacher-training component. Teachers have the opportunity to do experiments and write lesson plans under the guidance of university level faculty.

• In addition, recent neuroscience graduate Dr. Elizabeth Jeffress, joined forces with Fernbank’s Science Open Labs to present one of GSU’s most popular interactive neuroscience booths, “Touch a Brain”.

• The Brain Bee, a Society for Neuroscience program brought to Atlanta by CBN, has its roots in good ol’ fashion competition. High school students compete to convey their knowledge of the brain (and it’s a lot!) in a spelling bee format. Winners go on to a national competition.

• This past year, GSU grad students reached in to reach out!

#2 Founded in 1950, The DANA Foundation is a private philanthropic organization committed to advancing brain research and to educating the public in a responsible manner about research’s potential. They promote dialogue between researchers and lay audiences, provide validated information about the latest advances in research through free publications and websites and engage people worldwide through the Alliances and International Brain Awareness Week (BAW). See more at: www.dana.org

#3 From undergraduate & high school research programs, to teaching teachers, to classroom visits, the science education arm of the CBN set the stage for, what has now become, a culture of community engagement within the Neuroscience Institute here at Georgia State University. In addition, and something Katie looks forward to taking an active role in, The ACSfN plays a critical role in the partnership of science education efforts between three particularly active institutions: Georgia Tech, Emory and GSU.

A BRIGHT FUTURE In 2007, through tireless efforts of the CBN and ACSfN, the Atlanta neuroscience community knew they needed more than just one Brain Awareness Week to spread the brain love, if you will. The DANA initiative Brain Awareness Week in March quickly became Brain Awareness Month here in Atlanta. Even the governor’s office year after year acknowledges this month. A month was enough, right? Wrong. Thanks to the dedication of CBN, ACSfN and people like Katie, we have gone from one month focused on brain awareness to, what Drs. Frantz and Carruth coined as, ‘52 weeks of brain awareness.’

Today, while Drs. Carruth and Frantz are still both heavily involved in neuroscience advocacy and community enrichment activities, a lot of day to day work has become institutionalized within the Neuroscience Graduate Student Association (NGSA), CBN, and the ACSfN. Some initiatives from the early days are no longer active. Others have morphed into alternate undertakings. Still others are continuing to operate and thrive.

Founded by Emory University, the Georgia Institute of Technology, and the Metro Atlanta Chamber, the Atlanta Science Festival is a collaboration among diverse community partners planning a collection of events for young people, families, and adults. It is an annual public celebration of local science and technology. Curious people of all ages explore the science and technology in our region and see how science is connected to all parts of our lives. Scientists and educators from museums, local schools, universities, and companies uncover mysteries and explain discoveries in a variety of hands-on activities. Facility tours, stimulating presentations, and riveting performances expand our community of science enthusiasts and inspire a new generation of curious thinkers. With more than 100 individual events each year since the inaugural Festival in 2014, they have reached over 75,000 people. Collaborations with 100-plus community partners, including school districts, post-secondary institutions, museums, businesses, civic and community groups facilitated the festival’s success. Atlanta Science Festival is the proud winner of the 2015 Technology Association of Georgia STEM Education Award for Post-Secondary Outreach. (Excerpt reworded from http://atlantasciencefestival.org/about-asf)

• One of the signature programs originally adapted from an Emory program, BRAIN (Behavioral Research Advancements In Neuroscience), was an intensive summer research experience for undergrads. Although it has since sunsetted, its offspring of sorts, NET/work (Neuroscience Education & Training program) still carries on BRAIN’s charge to provide research training for undergrads. • The Neuroscience Expo was an exciting annual event of 2 days worth interactive activities. Running for 5 years, this far-reaching public awareness project made its way from the Civic Center to Zoo Atlanta, engaging tens of thousands of people and volunteers. • Founded by Emory, Georgia Tech, and the Metro Atlanta Chamber in 2014, The Atlanta Science Festival incorporates several brain-related aspects including interactive booths such as build-a-brain, touch-a-brain, robo-roach, zombie brains, and the smile booth (see side box #3). • Finally, there are undergraduate organizations such as the CNS, (Collegiate Neuroscience Society) and NuRhoPsi (Neuroscience national honors society) who put together noteworthy lectures and expert seminars open to the public as well.

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“Participant feedback suggests that our education and training programs pique curiosity, advance skills and knowledge, elevate positive attitudes about science at Georgia State, and even launch careers.” -Kyle Frantz ...Katie is still in training, but it only takes a few minutes of talking with her to note that her skills are honing in quickly. Most importantly, however, she knows she can reach out to a supportive and highly vibrant neuroscience education community if she needs any guidance or encouragement. In this community and others like it, a lot of work goes in. But the investment gives everyone involved a dividend. Everybody wins.

#5 Brains & Behavior raises awareness too with its annual Alzheimer’s Walk, raising almost $10,000 to date.

EVERYONE WINS #4 According to a report on supplies used from the lending library, the numbers from June 2014 –May 2015 are one reflection of the vitality of our neuroscience outreach community here in Atlanta. • Atlanta Science Fair – 2800 people reached • K-12 Classroom visits – 4,775 students reached • Teaching Undergraduates – 1,000 undergrads reached • Fernbank Natural History Museum Activities – 2,700 guests reached • Some 11,000 people were reached in 2015!

Spring 2016

Every year, it seems like brain awareness activities are growing exponentially, as if Atlanta were the perfect petri dish. Why not do it? It’s a feel good activity, right? Children smiling, people learning, some even going on to spread awareness themselves. In addition, the interactive nature of community engagement provides an avenue towards experiential learning skills across the ranks. However, these rewarding effects aren’t one-sided. Community engagement is mutually beneficial to the people receiving it and the institutions from which it came. Having volunteers in the classroom and graduate students behind the booths afford universities feet on the ground. In other words, it gives institutions direct contact with community partners and school age children. That direct contact and subsequent feedback helps inform brain related pedagogy. In addition, it offers on-site teacher training for graduate students. GSU Lecturer, and long time science educator, Dr. Joe Normadin remembers being a graduate student at GSU and claims that without the numerous opportunities to get engaged through outreach, he “wouldn’t have had the confidence to become an instructor at Georgia State”. Through our programs, Dr. Normandin became proficient in “techniques in teaching active learning curriculum.” Skills, Dr. Normandin admits, he often still uses today while teaching classes such as Intro to Neuroscience...

For more info on how to get involved in community engagement, contact: Katie Patrick: [email protected]

THE POTENTIAL

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Different Shades of Neuronal Subtype Diversity by Richard Campbell

D

uring the late 19th and early 20th century, there was a debate as to whether the nervous system was physically connected (continuous) or independent cells that communicated with one another (contiguous). Around that time, a famous neuroscientist, Santiago Ramón y Cajal, put forth evidence that the brain was made of contiguous units of cells (neurons) based upon their dendritic and axonal architecture. This became the basis of something called the neuron doctrine. The neuron doctrine, which has been essential to modern neuroscience, applies cellular theory to the nervous system (there are many more components to the neuron doctrine). From Ramón y Cajal’s experiments, it was apparent that not only was the nervous system composed of many different neurons, but that there are many subtypes of neurons that share similar morphological features. Many of these distinguishing features of different neuronal types are represented in the beautiful drawings that are still admired today for the paradigm shift they represented in neuroscience [1].

Spring 2016

During the last 100 years, an explosion of knowledge and advances have been made in studying the nervous system. As with any discipline, as new tools became available (Ex: electron microscopy, GFP, etc.) an understanding of the degree of heterogeneity of neuronal subtypes has increased. With the current tools available, we can identify neuron subtypes based upon a variety of factors. These include and are not limited to: location, morphology, neurotransmitter type, electrophysiological properties and more recently gene expression[2]. These properties have furthered scientific understanding as many different neuronal types and subtypes have been identified. However, many in the field have raised some unanswered questions. Why is there variation in the above mentioned properties from one animal, tissue or even cell to the next? Why is it evolutionarily beneficial to have so many neurons of the same general type in a given region? Like any complex problem, there is probably more than one answer. However, one of those answers may include a solution that is within reach. Recently, advances in sequencing and cell sorting have allowed for the identification of distinct populations of neurons within a given region of the nervous system. Several studies utilizing next generation single-cell sequencing, have revealed that there are many more neuronal subtypes than previously thought[3]. This is also true within populations of neurons, such as the CA1 pyramidal neurons found in the hippocampus. When this technique was applied to a population of CA1 neurons, clustering analysis of gene expression profiles revealed multiple heterogeneous groups [4]. However, this finding is not limited to the hippocampus. Further exploration via RNA-Seq of the cortex has revealed a multitude of subclasses based upon the expression of multiple genes [4, 5]. This is indicative that a single marker (such as single gene or morphology) may not be sufficient to accurately identify a specific subpopulation of neurons.

In various aspects, this principle of many subtypes in a general class of neurons has been intuitive and in some model systems already known. Invertebrate models have long been known to have many different subtypes of neurons that otherwise would appear to be the same. This is further highlighted by studies in which populations of similar neurons express different profiles of genes to create subtypes among the population. Developmental work in C. elegans and D. melanogaster have highlighted that this can be achieved by the use of common gene regulatory networks[6]. These networks are then modified by the expression of subtype specific transcription factors. In some cases, gene regulatory networks control a common identity among the population (Ex: neurotransmitter type) and are conserved from nematodes to mammals [6, 7]. In C. elegans, the cell lineages are well mapped, invariant and the differences between the subtypes of neurons is well described. Can this ability to describe variation between neurons of a similar group be scaled from C. elegans to other animals? It seems very reasonable that in the near future we will be able to identify and deduce the functional differences of subtypes in more complex systems than nematodes. I would predict that some, but not all, variation observed in terms of neuron function or properties may be due to multiple unidentified subtypes within the given region/population of interest. Indeed, differences between two neurons that appear to be the same may be the result of subtleties in their gene expression profile, which could distinguishes them as separate subtypes. It will be exciting to see this tested in the near future. Does this mean that an already incredibly complex system is far more complicated than we originally thought? In my opinion, and in the opinion of many others, it most certainly is. However, this reality provides many exciting opportunities for the future. Although, identify-

ing subtypes within [insert your model/ brain region of interest] is interesting in its own right, this endeavor can be further explored by asking how many different subtypes there are in a given population and delving deeper into the function of each subtype. Furthermore, we can begin to ask which subtypes are related to different behaviors or connections within the nervous system. These studies can be facilitated by the advances in sequencing and genome editing. From my own perspective, this observation of a multitude of subtypes also raises the question of how these different subtypes are determined. Thus, we can further explore the developmental events that lead to their creation. With the current set of tools available, we can begin to address some of the questions raised about neuronal subtypes in the nervous system. Just as the advancements in the late 19th century and 20th century provided an explosion of new information about the different types and functions of neurons, I believe there is still a wealth of unchartered territory to explore. This will provide exciting opportunities for both established and budding scientists alike. REFERENCES 1. De Carlos, J.A. and J. Borrell, A historical reflection of the contributions of Cajal and Golgi to the foundations of neuroscience. Brain Res Rev, 2007. 55(1): p. 8-16. 2. Lodato, S. and P. Arlotta, Generating Neuronal Diversity in the Mammalian Cerebral Cortex. Annu Rev Cell Dev Biol, 2015. 31: p. 699-720. 3. Tasic, B., et al., Adult mouse cortical cell taxonomy revealed by single cell transcriptomics. Nat Neurosci, 2016. 4. Cembrowski, M.S., et al., Spatial Gene-Expression Gradients Underlie Prominent Heterogeneity of CA1 Pyramidal Neurons. Neuron, 2016. 5. Zeisel, A., et al., Brain structure. Cell types in the mouse cortex and hippocampus revealed by single-cell RNA-seq. Science, 2015. 347(6226): p. 1138-42. 6. Hobert, O., Regulation of terminal differentiation programs in the nervous system. Annu Rev Cell Dev Biol, 2011. 27: p. 681-96. 7. Hobert, O., I. Carrera, and N. Stefanakis, The molecular and gene regulatory signature of a neuron. Trends Neurosci, 2010. 33(10): p. 435-45.