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Refereed publications


Sokolowski, D.J., Vasquez O., Wilson, M.D., Sokolowski, M.BAnreiter, I. 2023.Transcriptomic effects of the foraging gene shed light on pathways of pleiotropy and plasticity. Annals of the New York Academy of Sciences (early access  

Vasquez, O.E.1Allen, A.M.2, So, A-K 3Nguyen, Q.H.4, Krause, H.5, Levine, J.D.6,7Sokolowski M.B.1,7 2023. Characterizing the protein isoforms of foraging (for), the PKGI ortholog in Drosophila melanogaster. International Journal of Molecular Science. 6;24(12):10219. doi: 10.3390/ijms241210219. 

Gorchkova, M., Anreiter, ASokolowski, M.B. 2023. Individual Differences: Gene-Environment Interplay and Epigenetics. In Sex, Gender and Epigenetics. Edited by M.J. Legato, D. Feldberg, M. Glezerman.  Elsevier. ISBN: 987-0-12-823937-7, Chapter 17: pages 207-223.

Gram, S., Sokolowski, D.J., Sokolowski, M.B. 2023. Genes, Environments, Epigenetics. Principles of Gender-Specific Medicine: The Sex-Specific Biology in the Postgenomic Era 4e, Edited by Marianne J. Legato, Elsevier. ISBN: 978-0-323-88534-8, Chapter 12: pages 181-196.


Chen, M. and Sokolowski, M.B. 2022. How social experience and environment impacts behavioural plasticity in Drosophila. Fly. 16(1):68-84. pdf


Allen, A.M. Sokolowski, M.B. 2021. Expression and regulation of the foraging gene in Drosophila melanogaster. J. Neurogenetics 35(3): 192-213.(doi: 10.1080/01677063.2021.1941946).

Alwash, N., Allen, A.M., Sokolowski, M.B., Levine, J.D. 2021. The Drosophila melanogaster foraging gene affects social networks J. Neurogenetics 35(3): 249-262. (

Gray, L.J., Sokolowski, M.B., Simpson, S.J. 2021. Drosophila as a useful model for understanding the evolutionary physiology of obesity resistance and metabolic thrift. Fly 15: 47-51. DOI: 10.1080/19336934.2021.1896960

Gatev, E., Inkster, A.M., Negri, G.L., Konwar, C., Lussier, A.A., Skakkebaek, A., Sokolowski, M.B., Gravholt, C.H., Dunn, E.C., Kobor, M.S. and Aristizabal, M.J. 2021. Autosomal sex-associated co-methylated regions predict biological sex from DNA methylation. Nucleic Acids Research. 49(16):9097-9116. pdf

Dason, J.S. and Sokolowski, M.B. 2021. A cGMP-dependent protein kinase, encoded by the Drosophila foraging gene, regulates neurotransmission through changes in synaptic structure and function. Journal of Neurogenetics. 35(3):213-220. pdf

Anreiter, I., Allen, A.M., Vasquez, O.E., To, L., Douglas, S.J., Alvarez, J.V., Ewer, J. and Sokolowski, M.B. 2021. The Drosophila foraging gene plays a vital role at the start of metamorphosis for subsequent adult emergence. Journal of Neurogenetics. 35(3):179-191. pdf

Edelsparre, A.H., Hefley, T.J., Rodríguez, M.A., Fitzpatrick, M.J. and Sokolowski, M.B., 2021. Scaling up: understanding movement from individual differences to population-level dispersal. bioRxiv. 2021-01. pdf



Boyce, W.T., Sokolowski, M.B., Robinson, G.E. 2020. Genes and environments, development and time. Proceedings of the National Academy of Sciences. 117(38):23235-23241. pdf

Sinha, S., Jones, B.M., Traniello, I.M., Bukhari, S.A., Halfon, M.S., Hofmann, H.A., Huang, S., Katz, P.S., Keagy, J., Lynch, V.J., Sokolowski, M.B., Stubbs, L.J., Tabe-Bordbar, S., Wolfner, M.F., Robinson, G.E. 2020. Behavior-related gene regulatory networks: A new level of organization in the brain. Proceedings of the National Academy of Sciences. 117(38):23270-23279. pdf

Edelsparre, A.H., Fitzpatrick, M.J., Rodríguez, M.A., Sokolowski, M.B. 2020. Tracking dispersal across a patchy landscape reveals a dynamic interaction between genotype and habitat structure. Oikos. 130(1):79-94. pdf

Aristizabal, M.J., Anreiter, I., Halldorsdottir, T., Odgers, C.L., McDade, T.W., Goldenberg, A., Mostafavi, S.,  Kobor, M.S., Binder, E.B., Sokolowski, M.B., O’Donnell, K.J. 2020. Biological embedding of experience: a primer on epigenetics. Proceedings of the National Academy of Sciences.117(38):23261-23269. pdf

Dason, J.S., Cheung, A., Anreiter, I., Montemurri, V.A., Allen, A.M., Sokolowski, M.B. 2020. Drosophila melanogaster foraging regulates a nociceptive-like escape behavior through a developmentally plastic sensory circuit. Proceedings of the National Academy of Sciences. 117(38):23286-23291. pdf

Sokolowski, M.B. 2020. Honey bee colony aggression and indirect genetic effects. Proceedings of the National Academy of Sciences. 117(31):18148-18150. pdf

Sokolowski, M.B. 2020. Functional testing of ASD-associated genes. Proceedings of the National Academy of Sciences. 117(1):26-28. pdf


Anreiter I, Sokolowski MB. 2019. The foraging Gene and Its Behavioral Effects: Pleiotropy and Plasticity. Annual Review of Genetics. 53:373–92. pdf

Dason JS, Cheung A, Anreiter I, Montemurri VA, Allen AM, Sokolowski MB. 2019. Drosophila melanogaster foraging regulates a nociceptive-like escape behavior through a developmentally plastic sensory circuit. Proceedings of the National Academy of Sciences 201820840. pdf

Anreiter, I., Biergans, S., and Sokolowski, M.B. 2018. Epigenetic regulation of behaviour in Drosophila melanogaster. Current Opinion in Behavioral Sciences, 25:44-50. pdf

Struk, A.A., Mugon, J.,  Huston, A., Scholer, A.A., Stadler, G., Higgins, E.T., Sokolowski, M.B., Danckert, J. 2019. Self-regulation and the foraging gene (PRKG1) in humans. PNAS, ahead of print pdf 

Van Wert, M.*, Anreiter, I.*, Fallon, B.A., Sokolowski, M.B. 2019. Intergenerational Transmission of Child Abuse and Neglect: A Cross-Disciplinary Analysis. Gender and the Genome, 3:1-21. pdf


Anreiter, I., Sokolowski, M.B. 2018. Deciphering Pleiotropy: How Complex Genes Regulate Behavior. Communicative and Integrative Biology. pdf

Allen, A., Anreiter, I., Vesterberg, A. Sokolowski, M.B. 2018. Pleiotropy of the Drosophila melanogaster foraging gene on feeding-related traits. J. Neurogenetics. pdf

Hughson, B.A.*, Anreiter, I.*, Murphy K., Jackson-Chornenki, N., Ja, W., Huber, R., Sokolowski, M.B. 2017. The adult foraging assay (AFA) allows detection of strain and food-deprivation effects in feeding-related traits of Drosophila melanogasterJ. Insect Physiol 106(1): 20-29*co first author. pdf

Pan, P., Lawson, D.O., Dudin, A., Vasquez, O.E., Sokolowski, M.B., Fleming A.S., McGowan, P.O. 2018. Both maternal care and genotype influence stress-related phenotype in female rats. Developmental Psychobiology, 60(8):889-902. pdf


Anreiter, I., Kramer, J.M., Sokolowski, M.B. 2017. Epigenetic mechanisms modulate differences in Drosophila foraging behavior. PNAS 114(47): 12518–12523. pdf 10.1073/pnas.1710770114

Anreiter, I., Sokolowski, H.M., Sokolowski, M.B. 2017. Gene-Environment Interplay and Individual differences in behaviour.  Mind, Brain and Education.  pdf 10.1111/mbe.12158.

Malé PG, Turner KM, Doha M, Anreiter I, Allen AM, Sokolowski MB, Frederickson ME. 2017. An ant-plant mutualism through the lens of cGMP-dependent kinase genes. Proc Biol Sci. Sep 284(1862):20170896. pdf   doi: 10.1098/rspb.2017.0896

Wang, S. and Sokolowski, M.B. 2017. Aggressive behaviours, food-deprivation and the foraging gene. Royal Society Open Science 4: 170042. pdf

Allen, A.M., Anreiter, I., Neville M.C., Sokolowski, M.B. 2017. Feeding-related traits are afSfected by dosage of the foraging gene in Drosophila melanogaster. Genetics 205, 761-773.  pdf    doi: 10.1534/genetics.116.197939

Sokolowski, H.M., Vasquez, O.E., Unternaehrer, E., Sokolowski, D.J. Biergans, S.D., Atkinson, L., Gonzalez, A., Silveira, P.P., Levitan, R., O’Donnell, K.J., Steiner, M., Kennedy, J.,  Meaney, M.J., Fleming, A. S., Sokolowski M.B. on behalf of the MAVAN and Toronto Longitudinal Cohort research teams. (2017). The Drosophila foraging gene human orthologue PRKG1 predicts individual differences in the effects of early adversity on maternal sensitivity. Cognitive Development  pdf doi: 10.1016/j.cogdev.2016.11.001

Jonas, W. Fleming, A. Steiner, M. Meaney, M.J. Atkinson, L. Mileva, V. Sokolowski, M. Kennedy, J. 2017. Early adversity symptoms of depression and adversity. European Psychiatry 41:S30 pdf

Patrícia P. Silveira, Irina Pokhvisneva, I., Hélène Gaudreau, Leslie Atkinson, Alison S. Fleming, Marla B. Sokolowski, Meir Steiner, James L. Kennedy, Laurette Dubé, Robert D. Levitan, Michael J. Meaney, on behalf of the MAVAN research team. 2017. Fetal growth interacts with multilocus genetic score reflecting dopamine signaling capacity and predicts spontaneous palatable food intake in children. Appetite. 2017 Oct 14;120:596-601.  pdf doi: 10.1016/j.appet.2017.10.021


Levitan RD, Jansen P, Wendland B, Tiemeier H, Jaddoe VW, Silveira PP, Kennedy JL, Atkinson L, Fleming A, Sokolowski M, Gaudreau H, Steiner M, Dubé L, Hamilton J, Moss E, Wazana A, Meaney M. 2016. A DRD4 gene by maternal sensitivity interaction predicts risk for overweight or obesity in two independent cohorts of preschool children. J Child Psychol Psychiatry. pdf  doi: 10.1111/jcpp.12646

Anreiter I, Vasquez OE, Allen AM, Sokolowski MB. 2016. Foraging Path-length Protocol for Drosophila melanogaster Larvae. J. Vis. Exp. (110), e53980. pdf  doi:10.3791/53980

Riedl CAL, Oster S, Busto M, Mackay TFC, Sokolowski MB. 2016. Natural variability in Drosophila larval and pupal NaCl tolerance. J. Insect Physiol. 88:15-23. pdf  doi: 10.1016/j.jinsphys.2016.02.007.

Silveira PP, Gaudreau H, Atkinson L, Fleming AS, Sokolowski MB, Steiner M, Kennedy JL, Meaney MJ, Levitan RD, Dubé L. 2016. Genetic differential susceptibility to socio-economic status and childhood obesogenic behavior: Why targeted prevention may be the best societal investment. Jama Pediatrics (in press, Sept 2015, 5pgs). Editorial by Jay Belsky The differential susceptibility hypothesis, Sensitivity to the environment for better or for worse. Jama Pediatrics 02/01/20016 2pgs pdf     10.1001/jamapediatrics.2015.4253


Wazana, A, Moss, E, Jolicoeur-Martineau, A., Graffi, J., Tsabari, G., Lecompte, V., Pascuzzo, K., Babineau, V., Gordon-Green, C., Mileva, V., Atkinson, L., Minde, K., Bouvette-Turcot, A.A., Sassi, R., St-André, M., Carrey, N., Matthews, S., Sokolowski, M., Lydon, J., Gaudreau, H., Steiner, M., Kennedy, J.L., Fleming, A., Levitan, R., Meaney, M.J. 2015. The interplay of birth weight, dopamine receptor D4 gene (DRD4), and early maternal care in the prediction of disorganized attachment at 36 months of age. Dev Psychopathol. 2015 Nov;27(4 Pt 1):1145-61. pdf doi: 10.1017/S0954579415000735.

Bouvette-Turcot A.A., Pluess, M.,  Bernier, A., Pennestri, M.H.,  Levitan, R., Sokolowski, M.B., Kennedy, J., Minde, K., Steiner, M.,  Pokhvisneva, I.,  Meaney, M.J.,  Gaudreau, H. 2015. Effects of genotype and sleep on temperament. Pediatrics 136(Oct): 914-921. pdf

Bouvette-Turcot, A.A., Fleming, A.S., Wazana, A., Sokolowski, M.B., Gaudreau, H., Gonzalez, A., Deslauriers, J., Kennedy, J.L., Steiner, M., Meaney, M.J. 2015. Maternal childhood adversity and child temperament: An association moderated by child 5-HTTLPR genotype. Genes Brain Behav. pdf doi: 10.1111/gbb.12205. (in print)

Narasimha S, Kolly S, Sokolowski MB, Kawecki TJ, Vijendravarma RK. (2015) Prepupal Building Behavior in Drosophila melanogaster and Its Evolution under Resource and Time Constraints. PLoS One. 11;10(2):e0117280. doi:10.1371/journal.pone.0117280. pdf


Urquhart-Cronish, M. and Sokolowski, M.B. 2014. Gene-environment interplay in Drosophila melanogaster: chronic nutritional deprivation in larval life affects adult fecal output. Journal of Insect Physiology. 69:95-100. pdf

Hou, N., Armstrong, G.A., Chakraborty-Chatterjee, M., Sokolowski, M.B. and Robertson, R.M., 2014. Na+-K+-ATPase trafficking induced by heat shock pretreatment correlates with increased resistance to anoxia in locusts. Journal of Neurophysiology. 112(4):814-823. pdf

O’Donnell KA, Gaudreau H, Colalillo S, Steiner M, Moss E, Goldberg S, Karama S, Matthews SG, Lydon JE, Silveira PP, Wazana A, Levitan, RD, Sokolowski MB, Kennedy JL, Fleming A, Meaney MJ. 2014. The Maternal Adversity Vulnerability and Neurodevelopment (MAVAN) Project: Theory and methodology. The Canadian Journal of Psychiatry. 59(9):497–508. pdf


Kohn NR, Reaume CJ, Moreno C, Burns JC* Sokolowski MB, Mery F. 2013. Social environment influences performance in a cognitive task in natural variants of the foraging gene. PLoS One. 8(12): e81272.

Akbari EM, Shams S, Belay HT, Kaiguo M, Razak Z, Kent CF, Westwood T, Sokolowski MB, Fleming AS. 2013. The effects of parity and maternal behavior on gene expression in the medial pre optic area and the medial amygdala in postpartum and virgin female rats: A microarray study. Behav Neurosci. 127(6): 913-22.

Mileva-Seitz V., Kennedy J.L., Steiner M., Meaney M.J., Atkinson L., Levitan R.D., Sokolowski M.B., and Fleming A.S. 2013. Interaction between oxytocin genotypes and early experience predicts quality of mothering and postpartum mood. PLoS One 8: e61443.

Lovic, V., Belay, H., Walker, C.D., Burton, C.L., Meaney, M.J., Sokolowski, M., and Fleming, A.S. 2013. Early postnatal experience and DRD2 genotype affect dopamine receptor expression in the rat ventral striatum. Behavioural Brain Research 237: 278-282.

Sokolowski, M.B., Boyce W.T and McEwen B.S. 2013. Scarred for life? New Scientist 26:20-30. 


Eddison, M., Belay, A.T., Sokolowski, M.B., and Heberlein, U. 2012. A genetic screen for olfactory habituation mutations in Drosophila: Analysis of novel foraging alleles and an underlying neural circuit. PLoS One 7: e51684.

Donlea, J., Leahy, A., Thimgan, M.S., Suzuki, Y., Hughson, B.N., Sokolowski, M.B., and Shaw, P.J. 2012. foraging alters resilience/vulnerability to sleep disruption and starvation in Drosophila. PNAS 109: 2613-2618.

Burns, J.G., Svetec, N., Rowe, L., Mery, F., Dolan, M.J., Boyce, W.T., and Sokolowski, M.B. 2012. Gene-environment interplay in Drosophila melanogaster: Chronic food deprivation in early life affects adult exploratory and fitness traits. PNAS 109: 17239-17244.

Boyce, W.T., Sokolowski, M.B., and Robinson, G.E. 2012. Toward a new biology of social adversity. PNAS 109: 17143-17148.

Mileva-Seitz, V., Fleming, A.S., Meaney, M.J., Mastroianni, A., Sinnwell, J.P., Steiner, M., Atkinson, L., Levitan, R.D., Matthews, S.G., Kennedy, J.L., and Sokolowski, M.B. 2012. Dopamine receptors D1 and D2 are related to observed maternal behavior. Genes, Brain and Behavior 11: 684-694.

Kuntz, S., Poeck, B., Sokolowski, M.B., and Strauss, R. 2012. The visual orientation memory of Drosophila requires Foraging (PKG) upstream of Ignorant (RSK2) in ring neurons of the central complex. Learning & Memory 19: 337-340.


Reaume, C.J. and Sokolowski, M.B. 2011. Conservation of gene function in behaviour. Philosophical Transactions of the Royal Society B: Biological Sciences 366: 2100-2110.

Reaume, C.J., Sokolowski, M.B., and Mery, F. 2011. A natural genetic polymorphism affects retroactive interference in Drosophila melanogaster. Philosophical Transactions of the Royal Society B: Biological Sciences 278: 91-98.

Belay, H., Burton, C.L., Lovic, V., Meaney, M.J., Sokolowski, M., and Fleming, A.S. 2011. Early adversity and serotonin transporter genotype interact with hippocampal glucocorticoid receptor mRNA expression, corticosterone, and behavior in adult male rats. Behavioral Neuroscience 125: 150-160. 

Mileva-Seitz, V., Kennedy, J., Atkinson, L., Steiner, M., Levitan, R., Matthews, S.G., Meaney, M.J., Sokolowski, M.B., and Fleming, A.S. 2011. Serotonin transporter allelic variation in mothers predicts maternal sensitivity, behavior and attitudes toward 6-month-old infants. Genes, Brain and Behavior 10: 325-333.

Kramer, J.M., Kochinke, K., Oortveld, M.A., Marks, H., Kramer, D., de Jong, E.K., Asztalos, Z., Westwood, J.T., Stunnenberg, H.G., Sokolowski, M.B., Keleman, K., Zhou, H., van Bokhoven, H., and Schenck, A. 2011. Epigenetic regulation of learning and memory by Drosophila EHMT/G9a. PLoS Biology 9: e1000569. 


Dawson-Scully, K., Bukvic, D., Chakaborty-Chatterjee, M., Ferreira, R., Milton, S.L., and Sokolowski, M.B. 2010. Controlling Anoxic Tolerance in adult Drosophila via the cGMP-PKG Pathway. The Journal of Experimental Biology 213: 2410-2416.

Lucas, C., Kornfein, R., Chakaborty-Chatterjee, M., Schonfeld, J., Geva, N., Sokolowski, M.B., and Ayali, A. 2010. The locust foraging gene. Archives of Insect Biochemistry and Physiology 74: 52-66.

Reed, L.K., Williams, S., Springston, M., Brown, J., Freeman, K., Desroches, C., Sokolowski, M.B., and Gibson, G. 2010. Genotype-by-diet interactions drive metabolic phenotype variation in Drosophila melanogaster. Genetics 185: 1009-1019.

Desroches, C., Busto, M., Riedl, C.A.L., Mackay, T.F.C., and Sokolowski, M.B. 2010. Quantitative trait locus mapping of geotaxis behaviour in Drosophila melanogaster. Genetics Research 92: 167-174.

Sokolowski, M.B. 2010. Social interactions in “simple” model systems. Neuron 65: 780-794.

Kent, C.F., Daskalchuk, T., Cook, L., Sokolowski, M.B., and Greenspan, R.J. 2009. The Drosophila foraging gene mediates adult plasticity and gene-environment interactions in behaviour, metabolites, and gene expression in response to food deprivation. PLoS Genetics 5: e1000609.


Lucas, C., Hughson, B.N., and Sokolowski, M.B. 2009. Job switching in ants: role of a kinase. Communicative & Integrative Biology 3: 1-3.

Lucas, C. and Sokolowski, M.B. 2009. Molecular basis for changes in behavioral state in ant social behaviors. PNAS 106: 6351-6356.

Reaume, C.J. and Sokolowski, M.B. 2009. cGMP-dependent protein kinase as a modifier of behaviour. Handbook of Experimental Pharmacology 191: 423-443.

Williams, K.D. and Sokolowski, M.B. 2009. Evolution: How Fruit Flies Adapt to Seasonal Stresses. Current Biology 19: R63-64.


Kaun, K.R. and Sokolowski, M.B. 2008. cGMP dependent protein kinase: linking foraging to energy homeostasis. Genome 52: 1-7.

Kaun K.R. Chakaborty-Chatterjee, M., and Sokolowski, M.B. 2008. Natural variation in plasticitiy of glucose homeostasis and food intake. The Journal of Experimental Biology 211: 3160-3133.

Sokolowski, M.B. and Fitzpatrick, M.J. 2008. Behaviour genetics: worms seek that old beetle smell. Current Biology 10: R480-R482.


Belay, A.T., Scheiner, R., So, A.K.-C., Douglas, S.J., Chakaborty-Chatterjee, M., Levine, J.D., and Sokolowski, M.B. 2007. The foraging gene of Drosophila melanogaster: spatial-expression analysis and sucrose responsiveness. Journal of Comparative Neurology 504: 570-582.

Dawson-Scully, K., Armstrong, G.A.B., Kent, C., Robertson, R.M., and Sokolowski, M.B. 2007. Natural variation in the thermotolerance of neural function and behavior due to a cGMP-dependent protein kinase. PLoS One 2: e773.

Kaun, K., Hendel, T., Gerber B., and Sokolowski, M.B. 2007. Natural variation in Drosophila larval reward learning and memory due to a cGMP-dependent protein kinase. Learning & Memory 14: 342-349.

Kaun, K.R., Riedl, C.A.L., Chakaborty-Chatterjee, M., Belay, A.T., Douglas, S., Gibbs, A.G., and Sokolowski, M.B. 2007. Natural variation in food acquisition mediated via a Drosophila cGMP-dependent protein kinase. The Journal of Experimental Biology 210: 3547-3558.

Mery, F., Belay A., Sokolowski, M.B., and Kawecki, T.J. 2007. Natural polymorphism affecting learning and memory in Drosophila. PNAS 104: 13051-13055.

Riedl, C.A.L., Riedl, M., Mackay, T.F.C., and Sokolowski, M.B. 2007. Genetic and behavioural analysis of natural variation in Drosophila melanogaster pupation position. Fly 1: 23-32.

Fitzpatrick, M.J., Feder, E., Rowe, L., and Sokolowski, M.B. 2007. Maintaining a behaviour polymorphism by frequency-dependent selection on a single gene. Nature 447: 210-212.


Williams, K.D., Busto, M., Suster, M.L., So, A.K.-C., Ben-Shahar, Y., Leevers, S.J., and Sokolowski, M.B. 2006. Natural variation in Drosophila melanogaster diapause due to the insulin-regulated PI3-kinase. PNAS 103: 15911-15915.

Reaume, C.J. and Sokolowski, M.B. 2006. The nature of Drosophila melanogaster. Current Biology 16: R623-R628.


Douglas, S.J., Dawson-Scully, K., and Sokolowski, M.B. 2005. The neurogenetics and evolution of food-related behaviour. Trends in Neuroscience 28: 664-652.

Zakharkin, S.O., Belay, A., Fernandez, J.R., De Luca, V., Kennedy, J.L., Sokolowski, M.B., and Allison, D.B. 2005. Lack of association between polymorphism of the human cyclic GMP-dependent protein kinase gene and obesity. International Journal of Obesity 29: 872-874.

Riedl, C.A.L, Neal, S.J., Robichon, A., Westwood, J.T., and Sokolowski, M.B. 2005. Drosophila soluble guanylyl cyclase mutants exhibit increased foraging locomotion: behavioral and genomic investigations. Behavior Genetics 35: 231-244.

Hendel, T., Michels, B., Neuser, K., Schipanski, A., Kaun, K.R., Sokolowski, M.B., Marohn, F., Michel, R., Heisenberg, M., and Gerber, B. 2005. The carrot, not the stick: appetitive rather than aversive gustatory stimulation support associative olfactory learning in individually assayed Drosophila larvae. Journal of Comparative Physiology A 191: 265-279.

Fitzpatrick, M.J., Ben-Shahar, Y., Smid, H.M., Vet, L.E.M., Robinson, G.E., and Sokolowski M.B. 2005. Candidate genes for behavioural ecology. Trends in Ecology and Evolution 20: 96-104


Riedl, C.A.L. and Sokolowski, M.B. 2004. Behavioral genetics: guanylyl cyclase prompts worms to party. Current Biology 14: R657-R658.

Fitzpatrick, M.J. and Sokolowski, M.B. 2004. In search of food: exploring the evolutionary link between cGMP-dependent protein kinase (PKG) and behaviour. Integrative and Comparative Biology 44: 28-36.

Scheiner, R., Sokolowski, M.B., and Erber, J. 2004. Activity of cGMP-dependent protein kinase (PKG) affects sucrose responsiveness and habituation in Drosophila melanogaster. Learning & Memory 11: 303-311.

Suster, M.L., Karunanithi, S., Atwood, H.L., and Sokolowski, M.B. 2004. Turning behaviour in Drosophila larvae: a role for the small scribbler transcript. Genes, Brain and Behavior 3: 273-286.

Suster, M.L., Seugnet, L., Bate, M., and Sokolowski, M.B. 2004. Refining GAL4-driven transgene expression in Drosophila with a GAL80 enhancer-trap. Genesis 39: 240-245.


Sokolowski, M.B. 2003. NPY and the regulation of behavioral development. Neuron 39: 6-8. 

Ben-Shahar, Y., Leung, H.-T., Pak, W.L., Sokolowski, M.B., and Robinson, G.E. 2003. cGMP-dependent changes in phototaxis: a possible role for the foraging gene in honey bee division of labor. The Journal of Experimental Biology 206: 2507-2515. 


Sokolowski, M.B. 2002. Social eating for stress. Nature 419: 893-894. 

De Luca, V., Muglia, P., Jain, U., Basile, V.S., Sokolowski, M.B., and Kennedy, J.L. 2002. A Drosophila model for ADHD: no evidence of association with PRKGI gene. Neuromolecular Medicine 2: 281-287. 

Ben-Shahar, Y., Robichon, A., Sokolowski, M.B., and Robinson, G.E. 2002. Influence of gene action across different time scales on behavior. Science 296: 741-744. 


Sokolowski, M.B. 2001. Drosophila: Genetics meets behaviour. Nature Reviews Genetics 2: 879-890. 

Osborne, K, de Belle J.S., and Sokolowski, M.B. 2001. Foraging behaviour in Drosophila larvae: mushroom body ablation. Chemical Senses 26: 223-230. 


Yang, P., Shaver, S. A. Hilliker, A.J., and Sokolowski, M.B.. 2000. Abnormal turning behavior in Drosophila larvae: identification and molecular analysis of scribbler (sbb). Genetics 155: 1161-1174. 

Shaver, S.A., Riedl, C.A.L., Parkes, T.L., Sokolowski, M.B., and Hilliker, A.J. 2000. Isolation of larval behavioral mutants in Drosophila melanogaster. Journal of Neurogenetics 14: 193-205. 

Tsuji, L.J., DeIuliis, G., Hansell, R.I.C., Kozlovic, D.R., and Sokolowski, M.B. 2000. Non-random mating in classical lekking grouse species: seasonal and diurnal trends. International Journal of Biometeorology 44: 134-140. 

Engel, J.E., Xie, X.-J., Sokolowski, M.B., and Wu, C.-F. 2000. A cGMP-dependent protein kinase gene, foraging, modifies habituation-like response decrement of the giant fiber escape circuit in Drosophila. Learning & Memory 7: 341-352. 


Renger, J.J., Yao, W.-D., Sokolowski, M.B., and Wu, C.-F. 1999. Neuronal polymorphism among natural alleles of a cGMP-dependent kinase gene, foraging, in Drosophila. The Journal of Neuroscience 19: 1-8. 


Sokolowski, M.B. 1998. Genes for normal behavioral variation: recent clues from flies and worms. Neuron 21: 463-466. 

Shaver, S.A., Varnam, C., Hilliker, A.J., and Sokolowski, M.B. 1998. The foraging gene affects adult but not larval olfactory-related behavior in Drosophila. Behavioural Brain Research 95: 23-29. 


Osborne, K.A., Robichon, A., Burgess, E., Butland, S., Shaw, R.A., Coulthard, A. Pereira, H.S., Greenspan, R.J., and Sokolowski, M.B. 1997. Natural behavior polymorphism due to a cGMP-dependent protein kinase of Drosophila. Science 277: 834-836. 

Sokolowski, M.B., Pereira, H.S., and Hughes, K. 1997. Evolution of foraging behavior in Drosophila by density dependent selection. PNAS 94: 7373-7377. 

Rodd, F.H., Reznick, D.N., and Sokolowski, M.B. 1997. Phenotypic plasticity in the life history traits of guppies: response to social environment. Ecology 78: 419-433. 


Varnam, C., Strauss, R., De Belle, J.S., and Sokolowski, M.B. 1996. Larval behavior of Drosophila central complex mutants: interactions between no bridge, foraging, and chaser. Journal of Neurogenetics 11: 99-115. 

Tsuji, L.J.S., Karagatzides, and Sokolowski, M.B. 1996. Diet and internal anatomy of male sharp-tailed grouse Tympanuchus phasianellus as related to age and position on the lek. Canadian Field Naturalist 109: 433-436.

Hughes, K. and Sokolowski, M.B. 1996. Natural selection in the laboratory for a change in resistance by Drosophila melanogaster to the parasitoid wasp Asobara tabida. Journal of Insect Behavior 9: 477-491. 


Sawin-McCormack, E.P., Sokolowski, M.B., and Campos, A.R. 1995. Characterization and genetic analysis of Drosophila melanogaster photobehavior during larval development. Journal of Neurogenetics 10: 119-135. 

Pereira, H.S., MacDonald, D.E., Hilliker, A.J., and Sokolowski, M.B. 1995. Chaser (Csr), a new gene affecting larval foraging behavior in Drosophila melanogaster. Genetics 140: 263-270. 

Blows, M. and Sokolowski, M.B. 1995. The expression of additive and nonadditive genetic variation under stress. Genetics 140: 1149-1159. 

Rodd, F.H. and Sokolowski, M.B. 1995. Complex origins of variation in the sexual behaviour of male Trinidadian guppies, Poecilia reticulata: interactions among social environment, heredity, body size and age. Animal Behaviour 49: 1139-1159. 


Tsuji, L.J.S., Kozlovic, D.R., Sokolowski, M.B., and Hansell, R.I.C. 1994. Relationship of body size of male Sharp-tailed Grouse to location of individual territories on leks. Wilson Bulletin 106: 329-337. 

Sawin, E.P., Harris, L.R., Campos, A.R., and Sokolowski, M.B. 1994. Sensorimotor transformation from light reception to phototactic behavior in Drosophila larvae (Diptera: Drosophilidae). Journal of Insect Behavior 7: 553-567. 

Sawin, E.P., Dowse, H.B. Hamblen-Coyle, M.J., Hall, J.C., and Sokolowski, M.B. 1994. A lack of locomotor activity rhythms in Drosophila melanogaster larvae (Diptera: Drosophilidae). Journal of Insect Behavior 7: 249-262. 

Welbergen, P. and Sokolowski, M.B. 1994. Development time and pupation behavior in the Drosophila melanogaster subgroup (Diptera: Drosophiladae). Journal of Insect Behavior 7: 263-277. 

Carton, Y. and Sokolowski, M.B. 1994. Parasitization of embedded and non-embedded Drosophila melanogaster (Diptera: Drosophiladae) pupae by the Hymenopteran parasitoid Pachyerepoideus vindemniae (Hymenoptera: Pteromalidae). Journal of Insect Behavior 7: 129-131. 1


Vet, L.E.M., Sokolowski, M.B., MacDonald, D.E., and Snellen, H. 1993. Responses of a generalist and a specialist parasitoid (Hymenoptera: Eucoilidae) to Drosophilid larval kairomones. Journal of Insect Behavior 6: 615-624. 

Williams, K. and Sokolowski, M.B. 1993. Diapause in Drosophila melanogaster females: a genetic analysis. Heredity 71: 312-317. 

Pereira, H.S. and Sokolowski, M.B. 1993. Mutations in the larval foraging gene affect adult locomotory behavior after feeding in Drosophila melanogaster. PNAS 90: 5044-5046. 

De Belle, J.S., Sokolowski, M.B., and Hilliker, A.J. 1993. Genetic analysis of the foraging microregion of Drosophila melanogaster. Genome 36: 94-101. 


Tsuji, L.J.S., Kozlovic, D.R., and Sokolowski, M.B. 1992. Territorial position in Sharp-tailed Grouse leks and the probability of fertilization. Condor 94: 1030-1031. 

Sokolowski, M.B. and Hansell, K.P. 1992. The foraging locus: Behavioral tests for normal muscle movement in rover and sitter Drosophila melanogaster larvae. Genetica 85: 205-209. 

Carton, Y. and Sokolowski, M.B. 1992. Interactions between searching strategies of Drosophila parasitoids and the polymorphic behavior of their hosts. Journal of Insect Behavior 5: 161-175. 

Rodriguez, L., Sokolowski, M.B., and Shore, J.S. 1992. Habitat selection by Drosophila melanogaster larvae. Journal of Evolutionary Biology 5: 61-70.


Rodriguez, L., Sokolowski, M.B., and Carton, Y. 1991. Intra- and inter-specific variation in pupation behaviours of Drosophila from different habitats. Canadian Journal of Zoology 68: 2616-2619. 


Sokolowski, M.B. 1990. Functions of genetic analysis. European Bulletin of Cognitive Psychology 10: 675-680.

Sokolowski, M.B. and Carton, Y. 1990. Microgeographic variation in Drosophila melanogaster larval behavior. Journal of Insect Behavior 2: 829-833. 


De Belle, J.S., Hilliker, A.J., and Sokolowski, M.B. 1989. Genetic localization of foraging (for): A major gene for larval behaviour in Drosophila melanogaster. Genetics 123: 157-164. 

Sokolowski, M.B. and Bauer, S.J. 1989. Genetic analysis of pupation distance in Drosophila melanogaster. Heredity 62: 177-183. 

De Belle, J.S. and Sokolowski, M.B. 1989. Rover/sitter foraging behaviour in Drosophila melanogaster: genetic localization to chromosome-2L using compound autosomes. Journal of Insect Behavior 2: 291-299. 

Graf, S.A. and Sokolowski, M.B. 1989. The rover/sitter Drosophila foraging polymorphism as a function of larval development, food patch quality and starvation. Journal of Insect Behavior 2: 301-313. 

Bauer, S.J. and Sokolowski, M.B. 1988. Autosomal and maternal effects on pupation behaviour in Drosophila melanogaster. Behavior Genetics 18: 81-97. 

De Belle, J.S. and Sokolowski, M.B. 1987. Heredity of rover/sitter: alternative foraging strategies of Drosophila melanogaster. Heredity 59: 73-83. 

Sokolowski, M.B. and Turlings, T. 1987. Drosophila parasitoid-host interactions: vibrotaxis and ovipositor searching from the host’s perspective. Canadian Journal of Zoology 65: 461-464. 

Sokolowski, M.B., Bauer, S.J., Wai-Ping, V., Rodriguez, L.E., Wong, J., and Kent, C. 1986. Ecological genetics and behaviour of Drosophila melanogaster larvae in nature. Animal Behaviour 34: 403-408. 

Sokolowski, M.B. 1985. Genetic aspects to differences in foraging behavior. Behavioral Brain Sciences 8: 348-349.

Bauer, S.J. and Sokolowski, M.B. 1985. A genetic analysis of path length and pupation height in a natural population of Drosophila melanogaster. Canadian Journal of Genetics and Cytology 27: 334-340. 

Wong, J.L., Sokolowski, M.B., and Kent, C.F. 1985. Prepupation behavior in Drosophila: embedding. Behavior Genetics 15: 155-164. 

Sokolowski, M.B. 1985. Ecology, genetics and behavior of Drosophila larval foraging and pupation behavior. Journal of Insect Physiology 31: 857-864. 


Bauer, S.J. and Sokolowski, M.B. 1984. Larval foraging in isofemale lines of Drosophila melanogaster and D. pseudoobscura. Journal of Heredity 75: 131-134. 

Sokolowski, M.B., Kent, C., and Wong, J. 1984. Drosophila larval foraging behaviour: developmental stages. Animal Behaviour 32: 645-651. 

Sokolowski, M.B., Hansell, R.I.C., and Rotin, D. 1983. Larval foraging behavior. II.  Selection in the sibling species Drosophila melanogaster and D. simulans. Behavior Genetics 13: 169-177. 

Sokolowski, M.B. and Hansell, R.I.C. 1983. Larval foraging behavior. I. The sibling species Drosophila melanogaster and D. simulans. Behavior Genetics 13: 159-168. 

Sokolowski, M.B. and Hansell, R.I.C. 1983. Elucidating the behavioral phenotype of D. melanogaster larvae: correlations between foraging strategies and pupation heights. Behavior Genetics 13: 267-280. 

Sokolowski, M.B. 1982. Drosophila larval foraging behaviour: digging. Animal Behaviour 30: 1252-1253. 

Sokolowski, M.B. 1980. Foraging strategies of Drosophila melanogaster: a chromosomal analysis. Behavior Genetics 10: 291-302. 


Books/Book Chapters, Commentaries and non-Refereed Reports

Sajedinejad, S., Sansone, G., Fallon, B., Miller, S., Birken, C., Denburg, A., Jenkins, J., Levine, J., Mishna, F., Sokolowski, M.B., Stewart, S. 2021. Utilizing Information and Communication Technologies for the Provision of Child Welfare Services During the COVID-19 Pandemic. Toronto, Ontario: Policy
Bench, Fraser Mustard Institute of Human Development, University of Toronto. pdf

Sajedinejad, S., Sansone, G., Fallon, B., Miller, S., Gitterman, A., Cohen, E., Sokolowski, M.B. 2021. From Research to Impact: A Toolkit for Developing Effective Policy Briefs. Toronto, Ontario: Policy Bench, Fraser Mustard Institute of Human Development, University of Toronto. pdf

Alwash, N., Sokolowski, M.B., Levine, J.D. 2021. Exploring the effect of Drosophila melanogaster foraging gene on social networks. The role of the foraging gene (for) on Social Interaction Networks of Drosophila
melanogaster. 91. pdf

Sansone, G., Fallon, B., Miller, S., Birken, C., Denburg, A., Jenkins, J., Levine, J., Mishna, F., Sokolowski, M.B., Stewart, S.  2020. Children Aging Out of Care. Toronto, Ontario: Policy Bench, Fraser
Mustard Institute of Human Development, University of Toronto. pdf

Illes, J., Weiss, S., Bains, J., Chandler, J.A., Conrod, P., De Koninck, Y., Fellows, L.K., Groetzinger, D., Racine, E., Robillard, J.M., Sokolowski, M.B. 2019. A Neuroethics Backbone for the Evolving Canadian Brain Research Strategy. Neuron, 101(3):370-374.

Dason, J.S., Sokolowski, M.B., Wu, C-F. 2018. A reductionist approach to understanding the nervous system: the Harold Atwood legacy. J. Neurogenetics, 32(3):127-130.

Dason, J.S., Hegström-Wojtowicz, M., Sokolowski, M.B. 2018.
A look inside the Atwood lab. J. Neurogenetics, 32(3):279-293.

Williams K.D. and Sokolowski, M.B. Phenotypic plasticity and insect behaviour. Chapter 5.  In Insect Behavior from mechanisms to ecological and evolutionary consequences Alex Córdoba-Aguilar, Daniel González-Tokman & Isaac González-Santoyo (Editors). Cambridge University Press. (in press 2018).

Epigenetics: Synthesis. In: Tremblay RE, Boivin M, Peters RDeV, eds. Sokolowski MB, Boyce WT, topic eds. Encyclopedia on Early Childhood Development[online]. Updated November 2017. Accessed November 13, 2017.

Sokolowski MB, Boyce WT. Biology of the Epigenome. In: Tremblay RE, Boivin M, Peters RDeV, eds. Sokolowski MB, Boyce WT, topic eds. Encyclopedia on Early Childhood Development[online]. Published November 2017. Accessed November 13, 2017.

Sokolowski MB, Boyce WT. Gene-Environment Interplay and Epigenetic Processes. In: Tremblay RE, Boivin M, Peters RDeV, eds. Sokolowski MB, Boyce WT, topic eds. Encyclopedia on Early Childhood Development[online]. Published November 2017. Accessed November 13, 2017.

Sokolowski MB, Boyce WT. Epigenetic Embedding of Early Adversity and Developmental Risk. In: Tremblay RE, Boivin M, Peters RDeV, eds. Sokolowski MB, Boyce WT, topic eds. Encyclopedia on Early Childhood Development[online]. Published November 2017. Accessed November 13, 2017.

Sokolowski MB, Boyce WT. Epigenetics and the Role of Developmental Time. In: Tremblay RE, Boivin M, Peters RDeV, eds. Sokolowski MB, Boyce WT, topic eds. Encyclopedia on Early Childhood Development[online]. Published November 2017. Accessed November 13, 2017.

Sokolowski, M.B., Boyce W.T and McEwen B.S. 2013. Scarred for life? New Scientist 26:20-30.

Hughson, B., Pham, V. and Sokolowski, M.B. 2014. Behaviour genetic analysis of natural variation in foraging behaviour in Drosophila. Handbook of Drosophila Behaviour Genetics. Cambridge University Press.

Sokolowski, M.B. and Goodwin, S.F., eds. Gene-Environment Interplay. Advances in Genetics. Volume 77., Academic Press, 2012. Print.

Sokolowski, M.B. and Levine, J.D. “Nature-Nurture Interactions.” Social Behaviour: Genes, Ecology and Evolution. Ed. Székely, Tamás, Allen J. Moore, and Jan Komdeur. Cambridge University Press, 2010. Print.

Williams, K.D, Schmidt, P.S. and Sokolowski, M.B. “Photoperiodism in Insects: Molecular Basis and Consequences of Diapause.” Photoperiodism: The Biological Calendar. Ed. Nelson, Randy J., David L. Denlinger, and David E. Somers. Oxford University Press, 2009. Print.

Sokolowski, M.B., ed. Socio-Genetics. Academic Press, Advances in Genetics Volume 68.  2009. Print.

Ewer, J. and Sokolowski, M.B. “Drosophila.” The Encyclopaedia of Reproduction. Ed. Knobil, Ernst and Jimmy D. Neill. Academic Press, 2008. Print.

de Bono, M. and Sokolowski, M.B. “Foraging in Flies and Worms.” Invertebrate Neurobiology. Ed. North, Geoffrey and Ralph J. Greenspan. Cold Spring Harbor Lab Press, 2007. Print.

Belay, A.T. and Sokolowski, M.B. “Genetic Analysis of Food Search Behavior in the Fruit Fly (Drosophila melanogaster).” Neurobehavioral Genetics: Methods and Applications, Second Edition. Ed. Mormède, Pierre and Bryon C. Jones. CRC Press, 2006. Print.

Sokolowski, M.B. and Wahlsten, D. 2001. “Gene-environment interaction and complex behavior.” Methods in Genomic Neuroscience. Ed. Chin, Hemin R. and Steven O. Moldin. CRC Press, 2001. Print. PDF

Sokolowski, M.B. and Riedl, C.A.L. “Behavior-genetic and molecular analysis of naturally occurring variation in Drosophila larval foraging behavior.” Handbook of Molecular-Genetic Techniques for Brain and Behavior Research. Ed. Gerlai, R.T. and W.E. Crusio. Elsevier Science, 1999. Print.

Sokolowski, M.B. “Genetic Analysis of Food Search Behavior in the Fruit Fly Drosophila melanogaster.” Neurobehavioral Genetics: Methods and Applications. Ed. Mormède, Pierre and Bryon C. Jones. CRC Press, 1999. Print.

Sokolowski, M.B. “Genetic analysis of behaviour in the fruit fly Drosophila melanogaster.” Techniques for the Genetic Analysis of Brain and Behavior: Focus on the Mouse. Ed. Goldowitz, D., D. Wahlsten, and R.E. Wimer. Elsevier Science Ltd., 1992. Print.

Sokolowski, M.B. “The Relationship between Third Instar Larval Behavior and Age.” Evolutionary Genetics of Invertebrate Behavior. Ed. Huettel, Milton Davis. Plenum Press, 1986. Print.