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Alex Shingleton

Associate Professor of Biology


Integrative developmental biology: developmental biology, physiology and evolution.

Research Interests

Our laboratory studies the molecular-genetic and physiological regulation of body and organ size, and how this regulation evolves. We integrate molecular biology, physiology and evolutionary biology to generate a synthetic understanding of why animals are the size and shape that they are. Our research uses the fruit fly, Drosophila melanogaster, as a model organism for animal development.

See my research webpage for complete details.


Postdoctoral Fellow, Ecology and Evolutionary Biology, Princeton University 
PhD Zoology, Clare College, University of Cambridge 
BA Zoology, St. Peter’s College, University of Oxford 

Positions Held

08.2013 –: Associate Professor, Department of Biology, Lake Forest College 

07.2012 – 08.2013: Associate Professor, Department of Zoology, Michigan State University 

01.2006 – 06.2012: Assistant Professor, Department of Zoology, Michigan State University: 

Selected Courses Taught

Biology 120L: Organismal Biology
Biology 131: Biological Inquiry: The Biology of Dogs
Biology 143: Biological Inquiry: The Biology of Sex 
Biology 389: Evolution

Selected Publications

Research Articles

For complete list see here.

Shingleton, A.W., Masandika, J. R., Thorsen, L.S., Zhu, Y., Mirth, C.K. 2017. The sex-specific effects of diet quality versus quantity on size and shape in Drosophila melanogaster. Royal Society Open Science. 4: 170375.

Saleh Ziabari, O., Shingleton, A.W. 2017. Quantifying abdominal pigmentation in Drosophila melanogasterJournal of Visualized Experiments. 124. 55732

Stillwell, R.C., Shingleton, A.W., Dworkin, I., Frankino, W.A. 2016. Tipping the scales: Evolution of the allometric slope independent of average trait size. Evolution. 7: 433-444.

Dreyer, A.P., Saleh Ziabari, O., Swanson, E.M., Chawla, A., Frankino, W.A., Shingleton, A.W. 2016. Cryptic individual scaling relationships and the evolution of morphological scaling. Evolution. 70: 1703-1706.

Herbosa, L., Oliviera, M.M., Talamillo, Pérez, C., González, M., Martín, D., Sutherland, J.D., Shingleton, A.W., Mirth, C.K., Barrio, R. 2015. Ecdysone promotes growth of imaginal discs through the regulation of Thor in D. melanogaster. Scientific Reports. 5: 12383

Koyama, T., Rodrigues, M.A., Athanasiadas, A., Shingleotn, A.W., Mirth, C.K. 2014. Nutritional control of body size through FoxO-Ultraspiracle mediated ecdysone biosynthesis. eLife. 3:e03091.

Oliveira, M.M., Shingleton, A.W., Mirth, C.K. 2014. Coordination of Wing and Whole-Body Development at Developmental Milestones Ensures Robustness against Environmental and Physiological Perturbations. PLoS Genetics. 10(6): e1004408

Mirth, C.K., Tang, H., Makohon-Moore, S., Salhadar, S., Gokhale, R.H., Warner, R.D., Koyama, T., Riddiford, L.M., Shingleton, A.W., 2014. Juvenile hormone regulates body size and perturbs insulin-signaling in Drosophila. Proceedings of the National Academy of Science, doi: 10.1073/pnas.1313058111

Callier, V., Shingleton, A.W., Brent, C.S., Ghosh, S.M., Kim, J., Harrison, J.F., 2013. The role of reduced oxygen in the developmental physiology of growth and metamorphosis initiation in Drosophila melanogaster. The Journal of Experimental Biology, 216, 4334-4340

Ghosh, S.M., Testa, N.D., Shingleton, A.W. 2013. Tempertature Size Rule is mediated by thermal plasticity of critical size in Drosophila melanogasterProceedings of the Royal Society, London. Series B. 280, no 1760.

Testa, N.D., Ghosh, S.M., Shingleton, A.W. 2013. Sex-specific weight loss mediates sexual size dimorphism in Drosophila melanogaster. PLoS One, 8(3): e58936.

Shingleton, A.W., Tang, H.Y. 2012. Plastic flies: The regulation and evolution of trait variability in Drosophila. Fly, 6, 1-3

Tang, H., Smith-Caldas, M.S.R., Driscoll, M.V., Salhadar, S. &Shingleton, A.W.  2011. FOXO regulates organ-specific phenotypic plasticity in Drosophila. PLoS Genetics, 7, e1002373

Parker, N.F. & Shingleton, A.W.  2011. The coordination of growth among Drosophila organs in response to localized growth perturbation. Developmental Biology, 357, 318-325

Shingleton, A.W., Estep, C.M., Driscoll, M.V., Dworkin, I. 2009.Many ways to be small: Different environmental regulators of size generate different scaling relationships in Drosophila melanogasterProceedings of the Royal Society, London. Series B. 276, 2625-2633

Stieper, B.C., Kupershtok, M., Driscoll, M.V. & Shingleton, A.W. 2008. Imaginal disc growth regulates the timing of metamorphosis in Drosophila melanogaster. Developmental Biology. 321, 18-26

Shingleton, A.W., Mirth, C.K. & Bates, P.W. 2008. Developmental model of static allometry in holometabolous insects. Proceedings of the Royal Society, London. Series B. 275, 1875-1885

Shingleton, A.W., Das, J., Vinicius, L. & Stern, D.L. 2005. The temporal requirements for insulin signaling during development in Drosophila, PLoS Biology, 3 (9)


Shingleton, A.W. 2015. Physiology: Female flies have the guts for reproduction. Current Biology. 25: R716-718.

Gokhale, R.H., Shingleton, A.W. 2015. Size Control. WIRE: Developmental Biology. 4: 335-356.

Shingleton, A.W., Frankino, W.A. 2013. New perspectives on the evolution of exaggerated traits. BioEssays. 35, 100-107

Kültz, D., Clayton, D.F., Robinson, G.E., Albertson, C., Carey, H.V., Cummings, M.E., Dewar, K., Edwards, S.V., Hofmann, H.A., Gross, L.J., Kingsolver, J.G., Meaney, M.J., Schlinger, B.A., Shingleton, A.W., Sokolowski, M.B., Somero, G.N., Stanzione, D.C., Todgham, A.E. 2013. New frontiers for organismal biology. BioScience. 63, 464-471.

Mirth, C.K., Shingleton, A.W. 2012. Intergrating body and organ size in Drosophila: recent advances and outstanding problems. Frontiers in Experimental Endocrinology, 3:49

Shingleton, A.W. 2010. Allometry. Nature Education Knowledge. 1(9):2

Shingleton, A.W., Frankino, W.A., Flatt, T. Nijhout, H.F. & Emlen, D.J. 2007. Size and Shape: The regulation of static allometry in insects, BioEssays, 29 (6), 536-548

Shingleton, A.W. 2005, Body-Size Regulation: Combining Genetics and Physiology, Current Biology, 15 (20), R825-R827

Book Chapters

Shingleton, A.W. 2011. The regulation and evolution of growth and body size. in “Mechanisms of Life History Evolution” (T. Flatt & A. Heyland, Eds), OUP

Frankino, W.A., Emlen, D.J. & Shingleton, A.W.2009.Experimental approaches to studying the evolution of morphological allometries: The shape of things to come. in “Experimental Evolution” (T. Garland & M.R. Rose, Eds), University of California Press, Berkley.


2016 – 2019: NSF (IOS-1557638): The proximate basis of individual variation in phenotypic plasticity. ($436,772). PI. Shingleton

2013 – 2017:  NSF (IOS-1256565): Is hypoxia a critical cue for molting in Drosophila? ($326,000). PI: Shingleton

2009 – 2014: NSF (IOS-0919855): CAREER: Size Matters – The control of allometry in Drosophila. ($591,517). PI: Shingleton

2009 – 2013: NSF (IOS-0845847): Tipping the Scales - A selection approach to the developmental regulation of morphological scaling ($218,626). PI: Shingleton; Co-PI: I. Dworkin