Publications from the lab and its collaborators.

Spiny mice (Acomys) regenerate wounded whisker pad skin with whisker follicles, muscles, and targeted innervation.

Varholick, J.A., *Kondapaneni, R., Maden, M. (2025). npj Regenerative Medicine, 10:28

The first study to demonstrate that spiny mice can regenerate their whisker follicles and the associated structures after removal, establishing a new model system for studying cutaneous nerve regeneration.

Integrating regenerative biology with developmental psychobiology to understand behavioral recovery.

Varholick, J.A. (2025). Developmental Dynamics, Early View

A theory paper establishing the intimate connection of behavior and neuroplasticity with tissue regeneration.

Stroke-induced neuroplasticity in the spiny mouse, Acomys cahirinus.

*Kidd, B.M., Varholick, J.A., *Tuyn, D.M., Kamat, P.K., Simon, Z.D., Liu, L., Meckler, M.P., Pompilus, M., Bubenik, J.L., Davenport, M.L., Carter, H.A., Grudny, M.M., Barbazuk, W.B., Doré, S., Febo, M., Candelario-Jalil, E., Maden, M., Swanson, M.S. (2024). npj Regenerative Medicine, 9, 41.

After a severe stroke, spiny mice could not regenerate their brain, but they fully recovered behaviors within 48 hours. We collaborated on behavioral ethograms and assisted with MRI scans.

Older 6-9-month-old spiny mice (Acomys cahirinus) have delayed and spatially heterogeneous ear wound regeneration.

Varholick, J.A., *Thermolice, J., *Godinez, G., *Dos Santos, V., *Kondapaneni, R., Maden, M. (2024). Biology Open, 13 (10): bio060565.

Regeneration speed declines quickly within the first year of life, associated with a reduction in regeneration quality.

Bite wounds and dominance structures in male and female African spiny mice (Acomys cahirinus): Implications for animal welfare and the generalizability of results.

Varholick, J.A., *Godinez, G., *Jenkins, A., *Mobin, S., Maden, M. (2023). Animals, 14 (1), 64.

Weak skin and frequent fighting can lead to worrisome bite wounds in spiny mice. Most animals will have a bite wound in their lifetime despite stable dominance structures.

Spiny Mouse (Acomys): As an emerging research organism for regenerative medicine with applications beyond the skin.

Gaire, J., Varholick, J.A., Rana, S., Sunshine, M.D., Ahmad, A.S., Chan, K.M., Allen, K., Doré, S., Fuller, D.D., Maden, M., Simmons, C. (2021). npj Regenerative Medicine, 6(1), 1-6.

An overview of regeneration studies in spiny mice to acquaint translational researchers with this model organism.

Model systems for regeneration: the spiny mouse, Acomys cahirinus.

Maden, M., Varholick, J.A. (2020). Development, 147(dev167718).

A primer for regenerative biologists to understand and begin studying regeneration in spiny mice.

A bespoke water T-maze apparatus and protocol: An optimized, reliable, and repeatable method for screening learning, memory, and executive functioning in laboratory mice.

Bailoo, J.D., Bergeson, S.E., Ponomarev, I., Wilms, J.O., Kisby, B.R., Cornwall, G.A., MacDonald, C.C., Lawrence, J.J., Ganapathy, V., Sivaprakasam, S., Panthagani, P., Trasti, S., Varholick, J.A., Findlater, M., Deonarine, A. (2024). Frontiers in Behavioral Neuroscience, Learning and Memory, 18.

A novel water T-maze that is less stressful, more efficient, and more reliable than the traditional Morris water maze.

A systematic review and meta-analysis of the relationship between social dominance status and common behavioral phenotypes in male laboratory mice.

Varholick, J.A., Bailoo, J.D., *Jenkins, A., Voelkl, B., Würbel, H. (2021). Frontiers in Behavioral Neuroscience, Learning and Memory, 14(624036).

Methods of measuring dominance and stability are too heterogeneous to conclude whether dominance affects common behavioral measures in neuroscience.

Effects of weaning age and housing conditions on phenotypic differences in mice.

Bailoo, J.D., Voelkl, B., Varholick, J.A., Novak, J., Eimear, M., Russo, M., Palme, R., Würbel, H. (2020). Scientific Reports, 10(11684).

Altering weaning time and housing type had minimal effects on a broad range of phenotypic measures and likely has little effect on reproducibility in mouse experiments.

Social dominance hierarchy type and rank contribute to phenotypic variation within cages of laboratory mice.

Varholick, J.A., *Pontiggia, A., Murphy, E., *Daniele, V., Palme, R., Voelkl, B., Würbel, H., Bailoo, J.D. (2019). Scientific Reports, 9(13650).

The stability of dominance relationships, and position within them, consistently accounts for random variation on a range of phenotypic measures.

Effects of cage enrichment on behavior, welfare and outcome variability in female mice.

Bailoo, J.D., Murphy, E., Boada-Sana, M., Varholick, J.A., Hintze, S., Baussiere, C., Hahn, K.C., Gopfert, C., Palme, R., Voelkl, B., Würbel, H. (2018). Frontiers in Behavioral Neuroscience, 12, p 232.

Variability in enriched animals was similar to those housed in barren conditions, and enriched conditions reduced stereotypic behavior.

Phenotypic variability between social dominance ranks in laboratory mice.

Varholick, J.A., Bailoo, J.D., Palme, R., Würbel, H. (2018). Scientific Reports, 8(6593).

Dominance in the home cage was more dynamic than expected and had little effect on a range of phenotypic measures.

Evaluation of space allowance on measures of animal welfare and outcome variation in laboratory mice.

Bailoo, J.D., Murphy, E., Varholick, J.A., Novak, J., Palme, R., Voelkl, B., Würbel, H. (2018). Scientific Reports, 8(713).

Larger cages and varying group size did not affect welfare or outcome variation, so studies should consider group-housing mice in larger cages.

Maternal separation followed by isolation-housing differentially affects prepulse inhibition of the acoustic startle response in C57BL/6 mice.

Bailoo, J.D., Varholick, J.A., Garza, X.J., Jordan, R.L., & Hintze, S. (2016). Developmental Psychobiology, 58(8), 937-944.

Integrating regenerative biology with developmental psychobiology to understand behavioral recovery.

Varholick, J.A. (2024). Preprint on OSF Preprints.

An earlier and extended version of the theory paper published in Developmental Dynamics in 2025.

Social dominance status and social stability in spiny mice (Acomys cahirinus) and its relation to ear-hole regeneration and glucocorticoids.

Varholick, J.A., *Godinez, G., *Jenkins, A., *Mobin, S., Romeo, R.D., Corll, J., Barbazuk, W., Maden, M. (2022). Preprint on bioRxiv.

An earlier version of our dominance research on spiny mice, limited by low animal numbers; it serves as preliminary data for follow-up studies.