Publications

Peer-reviewed papers, open hardware, and classroom neuroscience.

A curated bibliography of Backyard Brains publications focused on neuroscience education, open-source lab tools, classroom electrophysiology, plant and animal behavior, and the occasional piece of scientific mischief. This public list emphasizes BYB education, open hardware, classroom electrophysiology, plant and animal behavior, and playful science communication.

Curated list

Included works22
Since 202011
SpikerBox Scholar results~279
Newest publication2025
    • 2025
    • Human

    Study while you sleep: using targeted memory reactivation as an independent research project for undergraduates

    J Mar'i, R Zhang, S Mircic, E Serbe-Kamp, M Meier, A Leonhardt, JW Antony, KA Norman, TC Marzullo, et al.

    Advances in Physiology Education 49 (1), 1-10

    Abstract: Why study when you could just sleep? We demonstrate how students can perform scalable research investigations to manipulate memory processing during sleep. It is a hands-on way to advance students' understanding of sleep-based memory consolidation and the corresponding neural mechanisms using open-source software and do-it-yourself EEG tools.

    View publication
    • 2024
    • Plant

    A library of electrophysiological responses in plants - a model of transversal education and open science

    Danae Madariaga, Derek Arro, Catalina Irarrázaval, Alejandro Soto, Felipe Guerra, Angélica Romero, Fabián Ovalle, Elsa Fedrigolli, Thomas DesRosiers, Étienne Serbe-Kamp, Timothy Marzullo

    Plant Signaling & Behavior 19 (1), 2310977

    Abstract: Electrophysiology in plants is understudied, and, moreover, an ideal model for student inclusion at all levels of education. Here, we report on an investigation in open science, whereby scientists worked with high school students, faculty, and undergraduates from Chile, Germany, Serbia, South Korea, and the USA. The students recorded the electrophysiological signals of >15 plant species in response to a flame or tactile stimulus applied to the leaves. We observed that approximately 60% of the plants studied showed an electrophysiological response, with a delay of ~ 3-6 s after stimulus presentation. In preliminary conduction velocity experiments, we verified that observed signals are indeed biological in origin, with information transmission speeds of ~ 2–9 mm/s. Such easily replicable experiments can serve to include more investigators and students in contributing to our understanding of plant electrophysiology. KEYWORDS: Plant electrical signals, electrophysiology, environmental stress, plant physiology, comparative physiology, DIY science, open science

    View publication
    • 2024
    • Neuroengineering
    • Preprint

    Building brains for robots: a hands-on approach to learning neuroscience in the classroom

    R Kannan, M Gendreau, A Hatch, SK Free, K Muriungi, YA Garje, et al.

    bioRxiv, 2024.05.15.594177

    Abstract: As the relevance of neuroscience in education grows, effective methods for teaching this complex subject in high school classrooms remain elusive. Integrating classroom experiments with brain-based robots offers a promising solution. This paper presents a structured curriculum designed around the use of camera-equipped mobile robots which enables students to construct and explore artificial neural networks.

    View publication
    • 2024
    • Human

    Low-cost classroom and laboratory exercises for investigating both wave and event-related electroencephalogram potentials

    K Smith, A Pilger, MLM Amorim, S Mircic, Z Reining, N Ristow, D Miller, JC Donovan, M Meier, TC Marzullo, et al.

    Journal of Undergraduate Neuroscience Education 22 (3), A197

    Abstract: Electroencephalography (EEG) has given rise to a myriad of new discoveries over the last 90 years. EEG is a non-invasive technique that has revealed insights into the spatial and temporal processing of brain activity over many neuroscience disciplines, including sensory, motor, sleep, and memory formation. However, most undergraduate students lack laboratory access to EEG recording equipment or the skills to perform an experiment independently. Here, we provide easy-to-follow instructions to measure both wave and event-related EEG potentials using a portable, low-cost amplifier (Backyard Brains, Ann Arbor, MI) that connects to smartphones and PCs, independent of their operating system.

    View publication
    • 2022
    • Invertebrate
    • Animal behavior

    Gills just want to have fun: can fish play games, just like us?

    S Eisenbeiser, E Serbe-Kamp, GJ Gage, TC Marzullo

    Animals 12 (13), 1684

    Abstract: It is common to observe play in dogs, cats, and birds, but have we been ignoring play in one of the most common house pets of all... fish? Aquarium fish are often used as meditative decoration in family households, but it could be that fish have similarly diverse behavioral repertoires as mammals and birds. To examine this theory, we conducted field tests at local pet stores where a range of aquarium fish species was tested for responsiveness to laser pointer stimuli.

    View publication
    • 2022
    • Human
    • Book

    How your brain works: neuroscience experiments for everyone

    G Gage, T Marzullo

    MIT Press, 2022

    Publisher description: Armed with some DIY electrodes, readers will get to see what brain activity really looks like through simple neuroscience experiments. Written by two neuroscience researchers who invented open-source techniques to record signals from neurons, muscles, hearts, eyes, and brains, How Your Brain Works includes more than forty-five experiments to gain a deeper understanding of your brain.

    View publication
    • 2021
    • Plant

    Authentic research investigations of a controversial question: can plants learn?

    J Shin, E Serbe, GJ Gage

    The American Biology Teacher 83 (4), 222-228

    Abstract: Can plants learn? This question stirs up controversy and speculation in the classroom, as it is currently doing in the scientific community at large. We leverage the controversy to ask students to contribute to the greater body of knowledge by using scientific principles in creative research projects.

    View publication
    • 2021
    • Neuroengineering

    The construction of high-magnification homemade lenses for a simple microscope: an easy DIY tool for biological and interdisciplinary education

    DP Flores, TC Marzullo

    Advances in Physiology Education

    Abstract: The rise of microscopy in the seventeenth century allowed scientists to discover a new world of microorganisms and achieve great physiological advances. Since Leeuwenhoek's design uses a single ball lens, it is possible to fabricate variations for educational activities in physics and biology university and high school classrooms.

    View publication
    • 2020
    • Human

    Developing and implementing low-cost remote laboratories for undergraduate biology and neuroscience courses

    C Hanzlick-Burton, J Ciric, M Diaz-Rios, W Colgan III, GJ Gage

    Journal of Undergraduate Neuroscience Education 19 (1), A118

    Abstract: The global pandemic caused by the novel coronavirus (SARS-COV-2) has forced many universities to abruptly change the delivery of courses from in-person to online. This change to remote learning requires creating new ways to deliver lectures, exams, and discussion groups through online meeting platforms. An often-overlooked challenge is performing lab courses that require access to specialized equipment and resources typically found in the undergraduate laboratory classrooms. Here we discuss some strategies for developing and implementing a full semester neuroscience laboratory course that allows students to fully participate in laboratory exercises at home or in their dorm rooms. Performing lab exercises remotely and independently was shown to significantly improve participant’s self-efficacy and confidence that they can learn complex neuroscience material, when compared to participants who passively watch experiments online. We review best practices to ensure that lessons can be successfully demonstrated by the instructor and carried out by all students. Finally, we discuss the need to provide a level playing field such that all students may succeed, regardless of their current technology resources at home. Keywords: distance learning, remote labs, neuroscience labs, remote teaching, anatomy & physiology (A&P)

    View publication
    • 2020
    • Neuroengineering

    Neurorobotics workshop for high school students promotes competence and confidence in computational neuroscience

    CA Harris, L Guerri, S Mircic, Z Reining, M Amorim, D Jovic, W Wallace, et al.

    Frontiers in Neurorobotics 14, 6

    Abstract: Understanding the brain is a fascinating challenge, captivating the scientific community and the public alike. The lack of effective treatment for most brain disorders makes the training of the next generation of neuroscientists, engineers and physicians a key concern. Here we introduce the use of neurorobots - robots controlled by computer models of biological neural networks - to teach computational neuroscience.

    View publication
    • 2019
    • Invertebrate

    An electrophysiological investigation of power-amplification in the ballistic mantis shrimp punch

    DJ Pollak, KD Feller, E Serbe, S Mircic, GJ Gage

    Journal of Undergraduate Neuroscience Education 17 (2), T12

    Abstract: Mantis shrimp are aggressive, burrowing crustaceans that hunt using one the fastest movements in the natural world. These stomatopods can crack the calcified shells of prey or spear down unsuspecting fish with lighting speed. Their strike makes use of power-amplification mechanisms to move their limbs much faster than is possible by muscles alone. Other arthropods such as crickets and grasshoppers also use power-amplified kicks that allow these animals to rapidly jump away from predator threats. Here we present a template laboratory exercise for studying the electrophysiology of power-amplified limb movement in arthropods, with a specific focus on mantis shrimp strikes. The exercise is designed in such a way that it can be applied to other species that perform power-amplified limb movements (e.g., house crickets, Acheta domesticus ) and species that do not (e.g., cockroaches, Blaberus discoidalis ). Students learn to handle the animals, make and implant electromyogram (EMG) probes, and finally perform experiments. This integrative approach introduces the concept of power-amplified neuromuscular control; allows students to develop scientific methods, and conveys high-level insights into behavior, and convergent evolution, the process by which different species evolve similar traits. Our power-amplification laboratory exercise involves a non-terminal preparation which allows electrophysiological recordings across multiple days from arthropods using a low-cost EMG amplifier. Students learn the principles of electrophysiology by fabricating their own electrode system and performing implant surgeries. Students then present behaviorally-relevant stimuli that generate attack strikes in the animals during the electrophysiology experiments to get insight into the underlying mechanisms of power amplification. Analyses of the EMG data (spike train burst duration, firing rate, and spike amplitude) allow students to compare mantis shrimp with other power-amplifying species, as well as a non-power-amplifying one. The major learning goal of this exercise is to empower students by providing an experience to develop their own setup to examine a complex biological principle. By contrasting power-amplifiers with non-power-amplifiers, these analyses highlight the peculiarity of power amplification at multiple levels of analysis, from behavior to physiology. Our comparative design requires students to consider the behavioral function of the movement in different species alongside the neuromuscular underpinnings of each movement. This laboratory exercise allows students to develop methodology, problem-solving and inquisitive skills crucial for pursuing science.

    View publication
    • 2019
    • Human

    The case for neuroscience research in the classroom

    GJ Gage

    Neuron 102 (5), 914-917

    Abstract: Neuroscience courses, largely relegated to advanced undergraduate or graduate universities, are now being offered in high schools and middle schools. Low-tech versions of advanced neuroscience research tools are being used in hands-on labs. In this NeuroView, I will argue the need for and provide an overview of neuroscience research beyond academia.

    View publication
    • 2017
    • Invertebrate

    Grasshopper DCMD: an undergraduate electrophysiology lab for investigating single-unit responses to behaviorally-relevant stimuli

    DMT Nguyen, M Roper, S Mircic, RM Olberg, GJ Gage

    Journal of Undergraduate Neuroscience Education 15 (2), A162

    Abstract: Avoiding capture from a fast-approaching predator is an important survival skill shared by many animals. Investigating the neural circuits that give rise to this escape behavior can provide a tractable demonstration of systems-level neuroscience research for undergraduate laboratories. In this paper, we describe three related hands-on exercises using the grasshopper and affordable technology to bring neurophysiology, neuroethology, and neural computation to life and enhance student understanding and interest. We simplified a looming stimuli procedure using the Backyard Brains SpikerBox bioamplifier, an open-source and low-cost electrophysiology rig, to extracellularly record activity of the descending contralateral movement detector (DCMD) neuron from the grasshopper’s neck. The DCMD activity underlies the grasshopper’s motor responses to looming monocular visual cues and can easily be recorded and analyzed on an open-source iOS oscilloscope app, Spike Recorder. Visual stimuli are presented to the grasshopper by this same mobile application allowing for synchronized recording of stimuli and neural activity. An in-app spike-sorting algorithm is described that allows a quick way for students to record, sort, and analyze their data at the bench. We also describe a way for students to export these data to other analysis tools. With the protocol described, students will be able to prepare the grasshopper, find and record from the DCMD neuron, and visualize the DCMD responses to quantitatively investigate the escape system by adjusting the speed and size of simulated approaching objects. We describe the results from 22 grasshoppers, where 50 of the 57 recording sessions (87.7%) had a reliable DCMD response. Finally, we field-tested our experiment in an undergraduate neuroscience laboratory and found that a majority of students (67%) could perform this exercise in one two-hour lab setting, and had an increase in interest for studying the neural systems that drive behavior.

    View publication
    • 2017
    • Neuroengineering
    • History

    The missing manuscript of Dr. Jose Delgado's radio controlled bulls

    TC Marzullo

    Journal of Undergraduate Neuroscience Education

    Abstract: Neuroscience systems level courses teach: 1) the role of neuroanatomical structures of the brain for perception, movement, and cognition; 2) methods to manipulate and study the brain including lesions, electrophysiological recordings, microstimulation, optogenetics, and pharmacology; 3) proper interpretation of behavioral data to deduce brain circuit operation; and 4) the similarities, differences, and ethics of animal models and their relation to human physiology. These four topics come together quite dramatically in Dr. Jose Delgado’s 1960s famous experiments on the neural correlates of aggression in which he stopped bulls in mid-charge by electrically stimulating basal ganglia and thalamic structures. Technical documentation on these experiments is famously difficult to find. Here I translate and discuss a Spanish language article written by Dr. Delgado in 1981 for an encyclopedia on bull fighting published in Madrid. Here Dr. Delgado appears to give the most complete explanation of his experiments on microstimulation of bovine brains. Dr. Delgado’s motivations, methods, and his interpretation of the bull experiments are summarized, as well as some accompanying information from his 1970 English language book: “Physical Control of the Mind.” This review of Dr. Delgado’s written work on the bull experiments can provide a resource to educators and students who desire to learn more about and interpret the attention-calling experiments that Dr. Delgado did on a ranch in Andalucía over 50 years ago.

    View publication
    • 2016
    • Invertebrate

    Leg Regrowth in Blaberus discoidalis following limb autotomy versus limb severance and relevance to neurophysiology experiments

    TC Marzullo

    PLOS ONE

    Abstract: Many insects can regenerate limbs, but less is known about the regrowth process with regard to limb injury type. As part of our neurophysiology education experiments involving the removal of a cockroach leg, the ability of Blaberus discoidalis cockroaches to regenerate a metathoracic leg was examined following autotomy at the femur/trochanter joint versus severance via a transverse coxa-cut.

    View publication
    • 2015
    • Neuroengineering
    • Open hardware

    Open Labware: 3-D printing your own lab equipment

    T Baden, AM Chagas, G Gage, T Marzullo, LL Prieto-Godino, T Euler

    PLOS Biology 13 (3), e1002086

    Abstract: The introduction of affordable, consumer-oriented 3-D printers is a milestone in the current maker movement. Combined with free and open sharing of detailed design blueprints and accessible development tools, rapid prototypes of complex products can now be assembled in one's own garage. Free-and-open online designs for self-built, sophisticated lab equipment provide access to teaching and research tools for an increasingly broad audience worldwide.

    View publication
    • 2014
    • Invertebrate

    Portable conduction velocity experiments using earthworms for the college and high school neuroscience teaching laboratory

    KM Shannon, GJ Gage, A Jankovic, WJ Wilson, TC Marzullo

    Advances in Physiology Education 38 (1), 62-70

    Abstract: The earthworm is ideal for studying action potential conduction velocity in a classroom setting, as its simple linear anatomy allows easy axon length measurements and the worm's sparse coding allows single action potentials to be easily identified. Here, we present a portable and robust experimental setup that allows students to perform conduction velocity measurements within a 30-min to 1-h laboratory session.

    View publication
    • 2013
    • Invertebrate

    Using crickets to introduce neurophysiology to early undergraduate students

    RK Dagda, RM Thalhauser, R Dagda, TC Marzullo, GJ Gage

    Journal of Undergraduate Neuroscience Education 12 (1), A66

    Abstract: Anatomy and physiology instructors often face the daunting task of teaching the principles of neurophysiology as part of a laboratory course with very limited resources. Teaching neurophysiology can be a difficult undertaking as sophisticated electrophysiology and data acquisition equipment is often financially out-of-reach for two-year institutions, and for many preparations, instructors need to be highly skilled in electrophysiology techniques when teaching hands-on laboratories. In the absence of appropriate laboratory tools, many undergraduate students have difficulty understanding concepts related to neurophysiology. The cricket can serve as a reliable invertebrate model to teach the basic concepts of neurophysiology in the educational laboratory. In this manuscript, we describe a series of hands-on, demonstrative, technologically simple, and affordable laboratory activities that will help undergraduate students gain an understanding of the principles of neurophysiology. By using the cerci ganglion and leg preparation, students can quantify extracellular neural activity in response to sensory stimulation, understand the principles of rate coding and somatotopy, perform electrical microstimulation to understand the threshold of sensory stimulation, and do pharmacological manipulation of neuronal activity. We describe the utility of these laboratory activities, provide a convenient protocol for quantifying extracellular recordings, and discuss feedback provided by undergraduate students with regards to the quality of the educational experience after performing the lab activities.

    View publication
    • 2012
    • Invertebrate
    • Open hardware

    The SpikerBox: a low cost, open-source bioamplifier for increasing public participation in neuroscience inquiry

    TC Marzullo, GJ Gage

    PLOS ONE 7 (3), e30837

    Abstract: Although people are generally interested in how the brain functions, neuroscience education for the public is hampered by a lack of low cost and engaging teaching materials. To address this, we developed an open-source tool, the SpikerBox, which is appropriate for use in middle/high school educational programs and by amateurs.

    View publication
    • 2008
    • Improbable research

    The cingulate cortex does everything

    GJ Gage, H Parikh, TC Marzullo

    Annals of Improbable Research 14 (3), 55

    Abstract: Here we explain most of the mysteries concerning the brain. We report the Cingular Theory of Unification, which postulates that one brain region, the cingulate cortex, is the alpha and omega, responsible for all of humankind's functions.

    View publication
    • 2006
    • Improbable research

    Stock market behavior predicted by rat neurons

    Timothy C. Marzullo, Edward G. Rantze, Gregory J. Gage

    Annals of Improbable Research 12 (4), 22-23

    Abstract: We here report for the first time, to the best of our knowledge, rat motor cortex neurons predicting the behavior of the American stock market. We implanted the motor cortex of the brains of rats with silicon electrodes. Using the correlation technique, we monitored the activity of neurons in our rats while simultaneously tracking the activity of stocks in the U.S. stock market.

    View publication