Neurofibromatosis type 1 (NF1) is caused by the loss of neurofibromin, a protein shown to regulate metabolism and feeding behavior in Drosophila melanogaster. Using the FlyPad two-choice feeding assay, we compared feeding microstructure between NF1 null mutants, NF1 heterozygotes, and wildtype controls. We find that NF1 mutants took significantly more sips of sucrose and with greater urgency, without changes in sip duration or sucrose taste sensitivity. Overall, these results suggest loss of NF1 increases feeding frequency.
","acknowledgements":"We want to thank Dr. Seth Tomchik and members of the Brown Lab for helpful discussions and constructive feedback, and Dr. Pavel Itskov for technical assistance. We also extend our appreciation to Dr. Barbara Shoplock and the FSU Young Scholars Program for academic and research support.
","authors":[{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","writing_originalDraft","writing_reviewEditing"],"email":"ZP23@fsu.edu","firstName":"Zohre","lastName":"Panahidizjikan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000000187195266"},{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","visualization","validation","writing_reviewEditing"],"email":"Sjb21q@fsu.edu","firstName":"Sydney","lastName":"Born","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0009-0008-7067-8876 "},{"affiliations":["Miami Palmetto Senior High School","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"maiaespinoza11@gmail.com","firstName":"Maia","lastName":"Espinoza","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Magnet Innovation Center, Inlet Beach, FL, USA","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"starcatcherfour@gmail.com","firstName":"Tate","lastName":"Lillian","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Florida State University, Tallahassee, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences","Program in Neuroscience"],"credit":["conceptualization","formalAnalysis","fundingAcquisition","methodology","project","resources","supervision","visualization","writing_originalDraft","writing_reviewEditing"],"email":"EBBrown@fsu.edu","firstName":"Elizabeth","lastName":"Brown","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-0825-8199"}],"awards":[{"awardId":"R00AG071833","funderName":"National Institutes of Health (United States)","awardRecipient":"Elizabeth Brown"},{"awardId":"","funderName":"Lamarr and Edith Trott Endowed Scholarship","awardRecipient":"Zohre Panahidizjikan"}],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for this work was provided by the National Institutes of Health (R00AG071833 to EBB) and FSU Sponsored Research, as well as the Lamarr and Edith Trott Endowed Scholarship to ZP.
","image":{"url":"https://portal.micropublication.org/uploads/3ac5c8ce9b9f8676a3c596ee59c8c4b7.jpg"},"imageCaption":"(A) Schematic of the FlyPad two-choice feeding assay, in which individual flies are offered simultaneous access to 1 mM and 5 mM sucrose. (B) There is a significant effect of genotype on sip number (H test: H2 = 23.54, P<0.0001). Loss of NF1 significantly increases sip number (+, P<0.0001; het, P<0.0085). (C) There is a significant effect of genotype on sip number when binned by sucrose concentration (REML: F2,268 = 11.00, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3706; het, P<0.3915), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0149). (D) Loss of NF1 has no effect on preference for 5mM sucrose (H test: H2 = 4.470, P<0.1070). (E) There is no effect of genotype on sip duration (REML: F2,134 = 2.895, P<0.0588). (F) There is no effect of genotype on intersip interval (REML: F2,132 = 1.253, P<0.2890). (G) There is a significant effect of genotype on feeding intensity (REML: F2,260 = 13.25, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3399; het, P<0.5110), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0001). (H-J) Distribution of the durations of sips of sucrose in (H) wildtype flies, (I) NF1 heterozygous flies, and (J) NF1 mutant flies. Error bars indicate ± SEM. *p<0.05; **p<0.01; ****p<0.0001.
","imageTitle":"Loss of NF1 increases food intake
","methods":"Drosophila care and maintenance
Flies were maintained on standard cornmeal-molasses-yeast food (Bloomington Recipe, Genesee Scientific, San Diego, CA) at 25°C and at ~50% relative humidity under a 12 h light: 12 h dark cycle. Mated ~3–5-day-old male flies were collected under light CO2 anesthesia and allowed to recover for at least 24 h prior to all experiments.
FlyPAD two-choice feeding assay
Feeding behavior was quantified using the flyPAD (fly Proboscis and Activity Detector; Itskov et al., 2014). Prior to testing, flies were starved for 18 h in vials on wet Kimwipe paper. On the day of testing, individual males were aspirated without anesthesia into flyPAD arenas configured for a two-choice assay, with the two food channels containing either 1 mM or 5 mM sucrose in 1% agarose. Flies were allowed to feed for 60 min, after which data were extracted and analyzed using established flyPAD analysis software (Itskov et al., 2014). The Preference Index (PI) was calculated as: (# sips of 5 mM sucrose − # sips of 1 mM sucrose) / (total # of sips), where a value of 1.0 indicates complete preference for 5mM sucrose and a value of -1.0 indicates a complete preference for 1mM sucrose. Feeding Intensity was calculated as: # sips / sip duration (Collins et al., 2026). Flies that did not feed were excluded from downstream analyses. All experiments were performed between ZT 0-2 at 25°C.
Statistical analysis
Feeding metrics are visualized as graphs displaying mean ± standard error. Dots represent individual fly measurements. Four independent runs were performed. The data were not normally distributed and so the non-parametric Kruskal-Wallis test was used to compare genotypes. To compare genotypes and sucrose concentration, a restricted maximum likelihood (REML) estimation was used. All post hoc analyses were performed using Dunn’s multiple comparisons test. Data were analyzed and visualized using InStat software (GraphPad Software 11.0).
","reagents":"Drosophila lines
The control line, w1118 was obtained from the Bloomington Stock Center (#5905). The NF1P1 line was a gift from Seth Tomchik’s lab at the University of Iowa (The et al., 1997). This line was backcrossed to the w1118 laboratory strain for 10 generations prior to testing.
","patternDescription":"Neurofibromatosis type 1 (NF1) is a neurodevelopmental disorder caused by mutations in the NF1 gene. This gene encodes neurofibromin, a GTPase-activating protein that negatively regulates the RAS signaling pathway (Buchberg et al., 1990; Xu et al., 1990). In humans, mutations in NF1 are associated with disruptions in several physiological and behavioral traits, including body size, metabolic function, and behavior (Botero & Tomchik, 2024). Many aspects of this disorder can be phenocopied using the fruit fly, Drosophila melanogaster: NF1-deficient flies have reduced body size, elevated metabolic rate, and reduced triglyceride stores (Botero et al., 2021, 2026; Brown et al., 2023; Sofela et al., 2024; The et al., 1997; Walker et al., 2006). Notably, loss of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024), consistent with a homeostatic response to their elevated metabolic rate and depleted energy stores. In this study, we further characterized the feeding behavior of NF1 flies.
Several approaches have been developed to quantify feeding behavior in Drosophila. These include the capillary feeder (CAFE) assay, which measures cumulative food consumption from a graduated capillary (Ja et al., 2007), manual scoring of proboscis extensions (Mair et al., 2005; Wong et al., 2009), as well as dye-labeling and radioisotope-labeling based approaches that estimate ingestion from colorimetric food additives or radiolabeled food, respectively (Carvalho et al., 2005; King & Wilson, 1955; Shimada et al., 1987; Tanimura et al., 1982). These approaches provide robust measures of total intake but lack the temporal and behavioral resolution to resolve individual feeding events. We therefore used the flyPAD (fly Proboscis and Activity Detector), which can detect each contact between a fly’s proboscis and a food source as a discrete change in capacitance, thereby enabling real-time, high-resolution quantification of feeding microstructure including sip number, duration, and choice between food sources in individual, freely feeding flies (Itskov et al., 2014).
Here, we utilized the flyPAD to investigate the effects of NF1 on food intake and feeding microstructure (Fig 1A). Starved male flies were provided the opportunity to feed from 1 mM and 5 mM sucrose, a paradigm that reports both feeding motivation and the ability to discriminate caloric value (Collins et al., 2026). We used the NF1P1 mutant line, which has a deletion that removes all the NF1 gene except for the first exon (The et al., 1997). We found that NF1P1 mutant flies take significantly more sips than both control and NF1P1 heterozygote flies (Fig 1B), consistent with previous findings that disruption of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024). We next broke down the number of sips by sucrose concentration and found that all genotypes took more sips of 5mM sucrose than of 1mM sucrose. However, NF1P1 mutant flies consistently took more sips of 5mM sucrose relative to control and NF1P1 heterozygote flies (Fig 1C). To evaluate the possibility that NF1P1 mutant flies are more sensitive to 5mM sucrose, we calculated a Preference Index for 5mM sucrose, where a positive value indicates stronger preference for 5mM sucrose and a negative value indicate stronger preference for 1mM sucrose. We found that preference for 5mM sucrose was consistent across all genotypes (Fig 1D), suggesting taste sensitivity to sucrose does not mediate the increased sips taken by NF1P1 flies.
In wild-type flies, food deprivation alters the microstructure of feeding by increasing the total number and duration of sips, and by decreasing the length between sips (Itskov et al., 2014; Banu et al., 2023). We next investigated whether NF1 functions to regulate these additional metrics of feeding microstructure. We found that the duration of each sip was not significantly different between genotypes (Fig 1E), nor was the interval between sips (Fig 1F). Therefore, NF1P1 flies consume more 5mM sucrose by increasing the frequency of sipping, not by taking longer sips or reducing time between sips, suggesting that loss of NF1 engages a more restricted feeding-drive mechanism than food deprivation alone. This is further reflected in measurements of feeding urgency (Collins et al., 2026), where we find that NF1P1 flies have significantly higher feeding intensity on 5mM sucrose relative to control and NF1P1 heterozygotes (Fig 1G). This may be driven primarily by the observation that NF1P1 flies have a greater frequency of very short duration sips relative to control and heterozygotes (Fig 1H-J). Overall, our results suggest that loss of NF1 further increases feeding frequency without altering the duration of individual sipping episodes or the ability to discriminate between sucrose concentrations, consistent with an increase in feeding drive.
","references":[{"reference":"Banu A, Gowda SBM, Salim S, Mohammad F. 2023. Serotonergic control of feeding microstructure in Drosophila. Frontiers in Behavioral Neuroscience 16: 10.3389/fnbeh.2022.1105579.
","pubmedId":"","doi":"10.3389/fnbeh.2022.1105579"},{"reference":"Botero V, Barrios J, Knauss A, Dahlen G, Rosendahl E, Colodner KJ, Tomchik SM. 2026. Metabolic and behavioral effects of neurofibromin result from differential recruitment of MAPK and mTOR signaling. PLOS Genetics 22: e1012061.
","pubmedId":"","doi":"10.1371/journal.pgen.1012061"},{"reference":"Botero V, Stanhope BA, Brown EB, Grenci EC, Boto T, Park SJ, et al., Tomchik. 2021. Neurofibromin regulates metabolic rate via neuronal mechanisms in Drosophila. Nature Communications 12: 10.1038/s41467-021-24505-x.
","pubmedId":"","doi":"10.1038/s41467-021-24505-x"},{"reference":"Botero V, Tomchik SM. 2024. Unraveling neuronal and metabolic alterations in neurofibromatosis type 1. Journal of Neurodevelopmental Disorders 16: 10.1186/s11689-024-09565-6.
","pubmedId":"","doi":"10.1186/s11689-024-09565-6"},{"reference":"Brown EB, Zhang J, Lloyd E, Lanzon E, Botero V, Tomchik S, Keene AC. 2023. Neurofibromin 1 mediates sleep depth in Drosophila. PLOS Genetics 19: e1011049.
","pubmedId":"","doi":"10.1371/journal.pgen.1011049"},{"reference":"Buchberg AM, Cleveland LS, Jenkins NA, Copeland NG. 1990. Sequence homology shared by neurofibromatosis type-1 gene and IRA-1 and IRA-2 negative regulators of the RAS cyclic AMP pathway. Nature 347: 291-294.
","pubmedId":"","doi":"10.1038/347291a0"},{"reference":"Carvalho GB, Kapahi P, Benzer S. 2005. Compensatory ingestion upon dietary restriction in Drosophila melanogaster. Nature Methods 2: 813-815.
","pubmedId":"","doi":"10.1038/nmeth798"},{"reference":"Collins NJ, Endres MN, Sinakevitch IT, Shao L. 2026. Protocol for quantifying Drosophila feeding behavior using flyPAD and optoPAD. STAR Protocols 7: 104711.
","pubmedId":"","doi":"10.1016/j.xpro.2026.104711"},{"reference":"Itskov PM, Moreira JM, Vinnik E, Lopes Ga, Safarik S, Dickinson MH, Ribeiro C. 2014. Automated monitoring and quantitative analysis of feeding behaviour in Drosophila. Nature Communications 5: 10.1038/ncomms5560.
","pubmedId":"","doi":"10.1038/ncomms5560"},{"reference":"Ja WW, Carvalho GB, Mak EM, de la Rosa NN, Fang AY, Liong JC, Brummel T, Benzer S. 2007. Prandiology of\n Drosophila\n and the CAFE assay. Proceedings of the National Academy of Sciences 104: 8253-8256.
","pubmedId":"","doi":"10.1073/pnas.0702726104"},{"reference":"King, R. C., & Wilson, L. P. (1955). Studies with radiophosphorus in Drosophila. V. The phosphorus balance of adult females. Journal of Experimental Zoology, 130(1), 71–82.
","pubmedId":"","doi":""},{"reference":"Mair W, Piper MDW, Partridge L. 2005. Calories Do Not Explain Extension of Life Span by Dietary Restriction in Drosophila. PLoS Biology 3: e223.
","pubmedId":"","doi":"10.1371/journal.pbio.0030223"},{"reference":"Shimada I, Nakao M, Kawazoe Y. 1987. Acute differential sensitivity and role of the central nervous system in the feeding behavior of Drosophila melanogaster. Chemical Senses 12: 481-490.
","pubmedId":"","doi":"10.1093/chemse/12.3.481"},{"reference":"Sofela FA, Lopez Valencia M, Jongens TA, Sehgal A. 2024. Effects of\n Nf1\n on sleep behavior are mediated through starvation caused by deficits in SARM1 dependent NAD+ metabolism. : 10.1101/2024.09.14.612058.
","pubmedId":"","doi":"10.1101/2024.09.14.612058"},{"reference":"Tanimura T, Isono K, Takamura T, Shimada I. 1982. Genetic dimorphism in the taste sensitivity to trehalose inDrosophila melanogaster. Journal of Comparative Physiology ? A 147: 433-437.
","pubmedId":"","doi":"10.1007/BF00612007"},{"reference":"The I, Hannigan GE, Cowley GS, Reginald S, Zhong Y, Gusella JF, Hariharan IK, Bernards A. 1997. Rescue of a Drosophila NF1 mutant phenotype by protein kinase A. Science 276(5313): 791-4.
","pubmedId":"9115203","doi":""},{"reference":"Walker JA, Tchoudakova AV, McKenney PT, Brill S, Wu D, Cowley GS, Hariharan IK, Bernards A. 2006. Reduced growth of Drosophila neurofibromatosis 1 mutants reflects a non-cell-autonomous requirement for GTPase-Activating Protein activity in larval neurons. Genes Dev 20(23): 3311-23.
","pubmedId":"17114577","doi":""},{"reference":"Wong R, Piper MDW, Wertheim B, Partridge L. 2009. Quantification of Food Intake in Drosophila. PLoS ONE 4: e6063.
","pubmedId":"","doi":"10.1371/journal.pone.0006063"},{"reference":"Xu G, O'Connell P, Viskochil D, Cawthon R, Robertson M, Culver M, et al., Weiss. 1990. The neurofibromatosis type 1 gene encodes a protein related to GAP. Cell 62: 599-608.
","pubmedId":"","doi":"10.1016/0092-8674(90)90024-9 "}],"title":"Neurofibromin 1 alters feeding microstructure in Drosophila
","reviews":[{"reviewer":{"displayName":"Amy Poe"},"openAcknowledgement":false,"status":{"submitted":true}},{"reviewer":{"displayName":"Woo Jae Kim"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Steven Marygold"},"openAcknowledgement":false,"submitted":null}]},{"id":"6142577e-f316-4a69-aae3-9803c17e5ac3","decision":"accept","abstract":"Neurofibromatosis type 1 (NF1) is caused by the loss of neurofibromin, a protein shown to regulate metabolism and feeding behavior in Drosophila melanogaster. Using the FlyPad two-choice feeding assay, we compared feeding microstructure between NF1 null mutants, NF1 heterozygotes, and wildtype controls. We find that NF1 mutants took significantly more sips of sucrose and with greater urgency, without changes in sip duration or sucrose taste sensitivity. Overall, these results suggest loss of NF1 increases feeding frequency.
","acknowledgements":"We want to thank Dr. Seth Tomchik and members of the Brown Lab for helpful discussions and constructive feedback, and Dr. Pavel Itskov for technical assistance. We also extend our appreciation to Dr. Barbara Shoplock and the FSU Young Scholars Program for academic and research support.
","authors":[{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","writing_originalDraft","writing_reviewEditing"],"email":"ZP23@fsu.edu","firstName":"Zohre","lastName":"Panahidizjikan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000000187195266"},{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","visualization","validation","writing_reviewEditing"],"email":"Sjb21q@fsu.edu","firstName":"Sydney","lastName":"Born","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0009-0008-7067-8876 "},{"affiliations":["Miami Palmetto Senior High School","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"maiaespinoza11@gmail.com","firstName":"Maia","lastName":"Espinoza","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Magnet Innovation Center, Inlet Beach, FL, USA","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"starcatcherfour@gmail.com","firstName":"Tate","lastName":"Lillian","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Florida State University, Tallahassee, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences","Program in Neuroscience"],"credit":["conceptualization","formalAnalysis","fundingAcquisition","methodology","project","resources","supervision","visualization","writing_originalDraft","writing_reviewEditing"],"email":"EBBrown@fsu.edu","firstName":"Elizabeth","lastName":"Brown","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-0825-8199"}],"awards":[{"awardId":"R00AG071833","funderName":"National Institutes of Health (United States)","awardRecipient":"Elizabeth Brown"},{"awardId":"","funderName":"Lamarr and Edith Trott Endowed Scholarship","awardRecipient":"Zohre Panahidizjikan"}],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for this work was provided by the National Institutes of Health (R00AG071833 to EBB) and FSU Sponsored Research, as well as the Lamarr and Edith Trott Endowed Scholarship to ZP.
","image":{"url":"https://portal.micropublication.org/uploads/6dbb1b22553139c36cdc94a56fdfb0e7.png"},"imageCaption":"(A) Schematic of the FlyPad two-choice feeding assay, in which individual flies are offered simultaneous access to 1 mM and 5 mM sucrose. (B) There is a significant effect of genotype on sip number (H test: H2 = 23.54, P<0.0001). Loss of NF1 significantly increases sip number (+, P<0.0001; het, P<0.0085). (C) There is a significant effect of genotype on sip number when binned by sucrose concentration (REML: F2,268 = 11.00, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3706; het, P<0.3915), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0149). (D) Loss of NF1 has no effect on preference for 5mM sucrose (H test: H2 = 4.470, P<0.1070). (E) There is no effect of genotype on sip duration (REML: F2,134 = 2.895, P<0.0588). (F) There is no effect of genotype on intersip interval (REML: F2,132 = 1.253, P<0.2890). (G) There is a significant effect of genotype on feeding intensity (REML: F2,260 = 13.25, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3399; het, P<0.5110), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0001). (H-J) Distribution of the durations of sips of sucrose in (H) wildtype flies, (I) NF1 heterozygous flies, and (J) NF1 mutant flies. Error bars indicate ± SEM. *p<0.05; **p<0.01; ****p<0.0001.
","imageTitle":"Loss of NF1 increases food intake
","methods":"Drosophila care and maintenance
Flies were maintained on standard cornmeal-molasses-yeast food (Bloomington Recipe, Genesee Scientific, San Diego, CA) at 25°C and at ~50% relative humidity under a 12 h light: 12 h dark cycle. Mated ~3–5-day-old male flies were collected under light CO2 anesthesia and allowed to recover for at least 24 h prior to all experiments.
FlyPAD two-choice feeding assay
Feeding behavior was quantified using the flyPAD (fly Proboscis and Activity Detector; Itskov et al., 2014). Prior to testing, flies were starved for 18 h in vials on wet Kimwipe paper. On the day of testing, individual males were aspirated without anesthesia into flyPAD arenas configured for a two-choice assay, with the two food channels containing either 1 mM or 5 mM sucrose in 1% agarose. Flies were allowed to feed for 60 min, after which data were extracted and analyzed using established flyPAD analysis software (Itskov et al., 2014). The Preference Index (PI) was calculated as: (# sips of 5 mM sucrose − # sips of 1 mM sucrose) / (total # of sips), where a value of 1.0 indicates complete preference for 5mM sucrose and a value of -1.0 indicates a complete preference for 1mM sucrose. Feeding Intensity was calculated as: # sips / sip duration (Collins et al., 2026). Flies that did not feed were excluded from downstream analyses. The resulting sample size for this experiment were as follows: controls (N = 43), NF1P1 heterozygotes (N = 39), and NF1P1 mutants (N = 55). All experiments were performed between ZT 0-2 at 25°C.
Statistical analysis
Feeding metrics are visualized as graphs displaying mean ± standard error. Dots represent individual fly measurements. Four independent runs were performed. The data were not normally distributed and so the non-parametric Kruskal-Wallis test was used to compare genotypes. To compare genotypes and sucrose concentration, a restricted maximum likelihood (REML) estimation was used. All post hoc analyses were performed using Dunn’s multiple comparisons test. Data were analyzed and visualized using InStat software (GraphPad Software 11.0).
","reagents":"Drosophila lines
The control line, w1118 was obtained from the Bloomington Stock Center (#5905). The NF1P1 line was a gift from Seth Tomchik’s lab at the University of Iowa (The et al., 1997). This line was backcrossed to the w1118 laboratory strain for 10 generations prior to testing.
","patternDescription":"Neurofibromatosis type 1 (NF1) is a neurodevelopmental disorder caused by mutations in the NF1 gene. This gene encodes neurofibromin, a GTPase-activating protein that negatively regulates the RAS signaling pathway (Buchberg et al., 1990; Xu et al., 1990). In humans, mutations in NF1 are associated with disruptions in several physiological and behavioral traits, including body size, metabolic function, and behavior (Botero & Tomchik, 2024). Many aspects of this disorder can be phenocopied using the fruit fly, Drosophila melanogaster: NF1-deficient flies have reduced body size, elevated metabolic rate, and reduced triglyceride stores (Botero et al., 2021, 2026; Brown et al., 2023; Sofela et al., 2024; The et al., 1997; Walker et al., 2006). Notably, loss of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024), consistent with a homeostatic response to their elevated metabolic rate and depleted energy stores. In this study, we further characterized the feeding behavior of NF1 flies.
Several approaches have been developed to quantify feeding behavior in Drosophila. These include the capillary feeder (CAFE) assay, which measures cumulative food consumption from a graduated capillary (Ja et al., 2007), manual scoring of proboscis extensions (Mair et al., 2005; Wong et al., 2009), as well as dye-labeling and radioisotope-labeling based approaches that estimate ingestion from colorimetric food additives or radiolabeled food, respectively (Carvalho et al., 2005; King & Wilson, 1955; Shimada et al., 1987; Tanimura et al., 1982). These approaches provide robust measures of total intake but lack the temporal and behavioral resolution to resolve individual feeding events. We therefore used the flyPAD (fly Proboscis and Activity Detector), which can detect each contact between a fly’s proboscis and a food source as a discrete change in capacitance, thereby enabling real-time, high-resolution quantification of feeding microstructure including sip number, duration, and choice between food sources in individual, freely feeding flies (Itskov et al., 2014).
Here, we utilized the flyPAD to investigate the effects of NF1 on food intake and feeding microstructure (Fig 1A). Starved male flies were provided the opportunity to feed from 1 mM and 5 mM sucrose, a paradigm that reports both feeding motivation and the ability to discriminate caloric value (Collins et al., 2026). We used the NF1P1 mutant line, which has a deletion that removes all the NF1 gene except for the first exon (The et al., 1997). We found that when we summed the total number of sips taken from both wells over the duration of the experiment NF1P1 mutant flies take significantly more sips than both control and NF1P1 heterozygote flies (Fig 1B), consistent with previous findings that disruption of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024). We next broke down the number of sips by sucrose concentration and found that all genotypes took more sips of 5mM sucrose than of 1mM sucrose. However, NF1P1 mutant flies consistently took more sips of 5mM sucrose relative to control and NF1P1 heterozygote flies (Fig 1C). To evaluate the possibility that NF1P1 mutant flies are more sensitive to 5mM sucrose, we calculated a Preference Index for 5mM sucrose, where a positive value indicates stronger preference for 5mM sucrose and a negative value indicate stronger preference for 1mM sucrose. We found that preference for 5mM sucrose was consistent across all genotypes (Fig 1D), suggesting taste sensitivity to sucrose does not mediate the increased sips taken by NF1P1 flies.
In wild-type flies, food deprivation alters the microstructure of feeding by increasing the total number and duration of sips, and by decreasing the length between sips (Itskov et al., 2014; Banu et al., 2023). We next investigated whether NF1 functions to regulate these additional metrics of feeding microstructure. We found that the duration of each sip was not significantly different between genotypes (Fig 1E), nor was the interval between sips (Fig 1F). Therefore, NF1P1 flies consume more 5mM sucrose by increasing the frequency of sipping, not by taking longer sips or reducing time between sips, suggesting that loss of NF1 engages a more restricted feeding-drive mechanism than food deprivation alone. This is further reflected in measurements of feeding urgency (Collins et al., 2026), where we find that NF1P1 flies have significantly higher feeding intensity on 5mM sucrose relative to control and NF1P1 heterozygotes (Fig 1G). This may be driven primarily by the observation that NF1P1 flies have a greater frequency of very short duration sips relative to control and heterozygotes (Fig 1H-J). Overall, our results suggest that loss of NF1 further increases feeding frequency without altering the duration of individual sipping episodes or the ability to discriminate between sucrose concentrations, consistent with an increase in feeding drive.
","references":[{"reference":"Banu A, Gowda SBM, Salim S, Mohammad F. 2023. Serotonergic control of feeding microstructure in Drosophila. Frontiers in Behavioral Neuroscience 16: 10.3389/fnbeh.2022.1105579.
","pubmedId":"","doi":"10.3389/fnbeh.2022.1105579"},{"reference":"Botero V, Barrios J, Knauss A, Dahlen G, Rosendahl E, Colodner KJ, Tomchik SM. 2026. Metabolic and behavioral effects of neurofibromin result from differential recruitment of MAPK and mTOR signaling. PLOS Genetics 22: e1012061.
","pubmedId":"","doi":"10.1371/journal.pgen.1012061"},{"reference":"Botero V, Stanhope BA, Brown EB, Grenci EC, Boto T, Park SJ, et al., Tomchik. 2021. Neurofibromin regulates metabolic rate via neuronal mechanisms in Drosophila. Nature Communications 12: 10.1038/s41467-021-24505-x.
","pubmedId":"","doi":"10.1038/s41467-021-24505-x"},{"reference":"Botero V, Tomchik SM. 2024. Unraveling neuronal and metabolic alterations in neurofibromatosis type 1. Journal of Neurodevelopmental Disorders 16: 10.1186/s11689-024-09565-6.
","pubmedId":"","doi":"10.1186/s11689-024-09565-6"},{"reference":"Brown EB, Zhang J, Lloyd E, Lanzon E, Botero V, Tomchik S, Keene AC. 2023. Neurofibromin 1 mediates sleep depth in Drosophila. PLOS Genetics 19: e1011049.
","pubmedId":"","doi":"10.1371/journal.pgen.1011049"},{"reference":"Buchberg AM, Cleveland LS, Jenkins NA, Copeland NG. 1990. Sequence homology shared by neurofibromatosis type-1 gene and IRA-1 and IRA-2 negative regulators of the RAS cyclic AMP pathway. Nature 347: 291-294.
","pubmedId":"","doi":"10.1038/347291a0"},{"reference":"Carvalho GB, Kapahi P, Benzer S. 2005. Compensatory ingestion upon dietary restriction in Drosophila melanogaster. Nature Methods 2: 813-815.
","pubmedId":"","doi":"10.1038/nmeth798"},{"reference":"Collins NJ, Endres MN, Sinakevitch IT, Shao L. 2026. Protocol for quantifying Drosophila feeding behavior using flyPAD and optoPAD. STAR Protocols 7: 104711.
","pubmedId":"","doi":"10.1016/j.xpro.2026.104711"},{"reference":"Itskov PM, Moreira JM, Vinnik E, Lopes Ga, Safarik S, Dickinson MH, Ribeiro C. 2014. Automated monitoring and quantitative analysis of feeding behaviour in Drosophila. Nature Communications 5: 10.1038/ncomms5560.
","pubmedId":"","doi":"10.1038/ncomms5560"},{"reference":"Ja WW, Carvalho GB, Mak EM, de la Rosa NN, Fang AY, Liong JC, Brummel T, Benzer S. 2007. Prandiology of\n Drosophila\n and the CAFE assay. Proceedings of the National Academy of Sciences 104: 8253-8256.
","pubmedId":"","doi":"10.1073/pnas.0702726104"},{"reference":"King, R. C., & Wilson, L. P. (1955). Studies with radiophosphorus in Drosophila. V. The phosphorus balance of adult females. Journal of Experimental Zoology, 130(1), 71–82.
","pubmedId":"","doi":""},{"reference":"Mair W, Piper MDW, Partridge L. 2005. Calories Do Not Explain Extension of Life Span by Dietary Restriction in Drosophila. PLoS Biology 3: e223.
","pubmedId":"","doi":"10.1371/journal.pbio.0030223"},{"reference":"Shimada I, Nakao M, Kawazoe Y. 1987. Acute differential sensitivity and role of the central nervous system in the feeding behavior of Drosophila melanogaster. Chemical Senses 12: 481-490.
","pubmedId":"","doi":"10.1093/chemse/12.3.481"},{"reference":"Sofela FA, Lopez Valencia M, Jongens TA, Sehgal A. 2024. Effects of\n Nf1\n on sleep behavior are mediated through starvation caused by deficits in SARM1 dependent NAD+ metabolism. : 10.1101/2024.09.14.612058.
","pubmedId":"","doi":"10.1101/2024.09.14.612058"},{"reference":"Tanimura T, Isono K, Takamura T, Shimada I. 1982. Genetic dimorphism in the taste sensitivity to trehalose inDrosophila melanogaster. Journal of Comparative Physiology ? A 147: 433-437.
","pubmedId":"","doi":"10.1007/BF00612007"},{"reference":"The I, Hannigan GE, Cowley GS, Reginald S, Zhong Y, Gusella JF, Hariharan IK, Bernards A. 1997. Rescue of a Drosophila NF1 mutant phenotype by protein kinase A. Science 276(5313): 791-4.
","pubmedId":"9115203","doi":""},{"reference":"Walker JA, Tchoudakova AV, McKenney PT, Brill S, Wu D, Cowley GS, Hariharan IK, Bernards A. 2006. Reduced growth of Drosophila neurofibromatosis 1 mutants reflects a non-cell-autonomous requirement for GTPase-Activating Protein activity in larval neurons. Genes Dev 20(23): 3311-23.
","pubmedId":"17114577","doi":""},{"reference":"Wong R, Piper MDW, Wertheim B, Partridge L. 2009. Quantification of Food Intake in Drosophila. PLoS ONE 4: e6063.
","pubmedId":"","doi":"10.1371/journal.pone.0006063"},{"reference":"Xu G, O'Connell P, Viskochil D, Cawthon R, Robertson M, Culver M, et al., Weiss. 1990. The neurofibromatosis type 1 gene encodes a protein related to GAP. Cell 62: 599-608.
","pubmedId":"","doi":"10.1016/0092-8674(90)90024-9 "}],"title":"Neurofibromin 1 alters feeding microstructure in Drosophila
","reviews":[{"reviewer":{"displayName":"Woo Jae Kim"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Steven Marygold"},"openAcknowledgement":false,"submitted":"1791874644215"}]},{"id":"aabefbbd-c973-406e-8ff8-3ae9fdc5b542","decision":"publish","abstract":"Neurofibromatosis type 1 (NF1) is caused by the loss of neurofibromin, a protein shown to regulate metabolism and feeding behavior in Drosophila melanogaster. Using the FlyPad two-choice feeding assay, we compared feeding microstructure between NF1 null mutants, NF1 heterozygotes, and wildtype controls. We find that NF1 mutants took significantly more sips of sucrose and with greater urgency, without changes in sip duration or sucrose taste sensitivity. Overall, these results suggest loss of NF1 increases feeding frequency.
","acknowledgements":"We want to thank Dr. Seth Tomchik and members of the Brown Lab for helpful discussions and constructive feedback, and Dr. Pavel Itskov for technical assistance. We also extend our appreciation to Dr. Barbara Shoplock and the FSU Young Scholars Program for academic and research support.
","authors":[{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","writing_originalDraft","writing_reviewEditing"],"email":"ZP23@fsu.edu","firstName":"Zohre","lastName":"Panahidizjikan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000000187195266"},{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","visualization","validation","writing_reviewEditing"],"email":"Sjb21q@fsu.edu","firstName":"Sydney","lastName":"Born","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0009-0008-7067-8876 "},{"affiliations":["Miami Palmetto Senior High School, Pinecrest, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"maiaespinoza11@gmail.com","firstName":"Maia","lastName":"Espinoza","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Magnet Innovation Center, Inlet Beach, FL, USA","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"starcatcherfour@gmail.com","firstName":"Tate","lastName":"Lillian","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Florida State University, Tallahassee, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences","Program in Neuroscience"],"credit":["conceptualization","formalAnalysis","fundingAcquisition","methodology","project","resources","supervision","visualization","writing_originalDraft","writing_reviewEditing"],"email":"EBBrown@fsu.edu","firstName":"Elizabeth B","lastName":"Brown","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-0825-8199"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for this work was provided by the National Institutes of Health (R00AG071833 to EBB) and FSU Office of Research, as well as the FSU Department of Biological Science Lamar and Edith Trott Endowed Scholarship to ZP.
","image":{"url":"https://portal.micropublication.org/uploads/6dbb1b22553139c36cdc94a56fdfb0e7.png"},"imageCaption":"(A) Schematic of the FlyPad two-choice feeding assay, in which individual flies are offered simultaneous access to 1mM and 5mM sucrose. (B) There is a significant effect of genotype on sip number (H test: H2 = 23.54, P<0.0001). Loss of NF1 significantly increases sip number (+, P<0.0001; het, P<0.0085). (C) There is a significant effect of genotype on sip number when binned by sucrose concentration (REML: F2,268 = 11.00, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3706; het, P<0.3915), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0149). (D) Loss of NF1 has no effect on preference for 5mM sucrose (H test: H2 = 4.470, P<0.1070). (E) There is no effect of genotype on sip duration (REML: F2,134 = 2.895, P<0.0588). (F) There is no effect of genotype on intersip interval (REML: F2,132 = 1.253, P<0.2890). (G) There is a significant effect of genotype on feeding intensity (REML: F2,260 = 13.25, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3399; het, P<0.5110), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0001). (H-J) Distribution of the durations of sips of sucrose in (H) wildtype flies, (I) NF1 heterozygous flies, and (J) NF1 mutant flies. Error bars indicate ± SEM. *p<0.05; **p<0.01; ****p<0.0001.
","imageTitle":"Loss of NF1 increases food intake
","methods":"Drosophila care and maintenance
Flies were maintained on standard cornmeal-molasses-yeast food (Bloomington Recipe, Genesee Scientific, San Diego, CA) at 25°C and at ~50% relative humidity under a 12 h light: 12 h dark cycle. Mated ~3–5-day-old male flies were collected under light CO2 anesthesia and allowed to recover for at least 24 h prior to all experiments.
FlyPAD two-choice feeding assay
Feeding behavior was quantified using the flyPAD (fly Proboscis and Activity Detector; Itskov et al., 2014). Prior to testing, flies were starved for 18 h in vials on wet Kimwipe paper. On the day of testing, individual males were aspirated without anesthesia into flyPAD arenas configured for a two-choice assay, with the two food channels containing either 1 mM or 5 mM sucrose in 1% agarose. Flies were allowed to feed for 60 min, after which data were extracted and analyzed using established flyPAD analysis software (Itskov et al., 2014). The Preference Index (PI) was calculated as: (# sips of 5 mM sucrose − # sips of 1 mM sucrose) / (total # of sips), where a value of 1.0 indicates complete preference for 5mM sucrose and a value of -1.0 indicates a complete preference for 1mM sucrose. Feeding Intensity was calculated as: # sips / sip duration (Collins et al., 2026). Flies that did not feed were excluded from downstream analyses. The resulting sample size for this experiment were as follows: controls (N = 43), NF1P1 heterozygotes (N = 39), and NF1P1 mutants (N = 55). All experiments were performed between ZT 0-2 at 25°C.
Statistical analysis
Feeding metrics are visualized as graphs displaying mean ± standard error. Dots represent individual fly measurements. Four independent runs were performed. The data were not normally distributed and so the non-parametric Kruskal-Wallis test was used to compare genotypes. To compare genotypes and sucrose concentration, a restricted maximum likelihood (REML) estimation was used. All post hoc analyses were performed using Dunn’s multiple comparisons test. Data were analyzed and visualized using InStat software (GraphPad Software 11.0).
","reagents":"Drosophila lines
The control line, w1118 was obtained from the Bloomington Stock Center (#5905). The NF1P1 line was a gift from Seth Tomchik’s lab at the University of Iowa (The et al., 1997). This line was backcrossed to the w1118 laboratory strain for 10 generations prior to testing.
","patternDescription":"Neurofibromatosis type 1 (NF1) is a neurodevelopmental disorder caused by mutations in the NF1 gene. This gene encodes neurofibromin, a GTPase-activating protein that negatively regulates the RAS signaling pathway (Buchberg et al., 1990; Xu et al., 1990). In humans, mutations in NF1 are associated with disruptions in several physiological and behavioral traits, including body size, metabolic function, and behavior (Botero & Tomchik, 2024). Many aspects of this disorder can be phenocopied using the fruit fly, Drosophila melanogaster: NF1-deficient flies have reduced body size, elevated metabolic rate, and reduced triglyceride stores (Botero et al., 2021, 2026; Brown et al., 2023; Sofela et al., 2024; The et al., 1997; Walker et al., 2006). Notably, loss of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024), consistent with a homeostatic response to their elevated metabolic rate and depleted energy stores. In this study, we further characterized the feeding behavior of NF1 flies.
Several approaches have been developed to quantify feeding behavior in Drosophila. These include the capillary feeder (CAFE) assay, which measures cumulative food consumption from a graduated capillary (Ja et al., 2007), manual scoring of proboscis extensions (Mair et al., 2005; Wong et al., 2009), as well as dye-labeling and radioisotope-labeling based approaches that estimate ingestion from colorimetric food additives or radiolabeled food, respectively (Carvalho et al., 2005; King & Wilson, 1955; Shimada et al., 1987; Tanimura et al., 1982). These approaches provide robust measures of total intake but lack the temporal and behavioral resolution to resolve individual feeding events. We therefore used the flyPAD (fly Proboscis and Activity Detector), which can detect each contact between a fly’s proboscis and a food source as a discrete change in capacitance, thereby enabling real-time, high-resolution quantification of feeding microstructure including sip number, duration, and choice between food sources in individual, freely feeding flies (Itskov et al., 2014).
Here, we utilized the flyPAD to investigate the effects of NF1 on food intake and feeding microstructure (Fig 1A). Starved male flies were provided the opportunity to feed from 1 mM and 5 mM sucrose, a paradigm that reports both feeding motivation and the ability to discriminate caloric value (Collins et al., 2026). We used the NF1P1 mutant line, which has a deletion that removes all of the NF1 gene except for the first exon (The et al., 1997). We found that when we summed the total number of sips taken from both wells over the duration of the experiment NF1P1 mutant flies take significantly more sips than both control and NF1P1 heterozygote flies (Fig 1B), consistent with previous findings that disruption of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024). We next broke down the number of sips by sucrose concentration and found that all genotypes took more sips of 5mM sucrose than of 1mM sucrose. However, NF1P1 mutant flies consistently took more sips of 5mM sucrose relative to control and NF1P1 heterozygote flies (Fig 1C). To evaluate the possibility that NF1P1 mutant flies are more sensitive to 5mM sucrose, we calculated a Preference Index for 5mM sucrose, where a positive value indicates stronger preference for 5mM sucrose and a negative value indicate stronger preference for 1mM sucrose. We found that preference for 5mM sucrose was consistent across all genotypes (Fig 1D), suggesting taste sensitivity to sucrose does not mediate the increased sips taken by NF1P1 flies.
In wild-type flies, food deprivation alters the microstructure of feeding by increasing the total number and duration of sips, and by decreasing the length between sips (Itskov et al., 2014; Banu et al., 2023). We next investigated whether NF1 functions to regulate these additional metrics of feeding microstructure. We found that the duration of each sip was not significantly different between genotypes (Fig 1E), nor was the interval between sips (Fig 1F). Therefore, NF1P1 flies consume more 5mM sucrose by increasing the frequency of sipping, not by taking longer sips or reducing time between sips, suggesting that loss of NF1 engages a more restricted feeding-drive mechanism than food deprivation alone. This is further reflected in measurements of feeding urgency (Collins et al., 2026), where we find that NF1P1 flies have significantly higher feeding intensity on 5mM sucrose relative to control and NF1P1 heterozygotes (Fig 1G). This may be driven primarily by the observation that NF1P1 flies have a greater frequency of very short duration sips relative to control and heterozygotes (Fig 1H-J). Overall, our results suggest that loss of NF1 further increases feeding frequency without altering the duration of individual sipping episodes or the ability to discriminate between sucrose concentrations, consistent with an increase in feeding drive.
","references":[{"reference":"Banu A, Gowda SBM, Salim S, Mohammad F. 2023. Serotonergic control of feeding microstructure in Drosophila. Frontiers in Behavioral Neuroscience 16: 10.3389/fnbeh.2022.1105579.
","pubmedId":"","doi":"10.3389/fnbeh.2022.1105579"},{"reference":"Botero V, Barrios J, Knauss A, Dahlen G, Rosendahl E, Colodner KJ, Tomchik SM. 2026. Metabolic and behavioral effects of neurofibromin result from differential recruitment of MAPK and mTOR signaling. PLOS Genetics 22: e1012061.
","pubmedId":"","doi":"10.1371/journal.pgen.1012061"},{"reference":"Botero V, Stanhope BA, Brown EB, Grenci EC, Boto T, Park SJ, et al., Tomchik. 2021. Neurofibromin regulates metabolic rate via neuronal mechanisms in Drosophila. Nature Communications 12: 10.1038/s41467-021-24505-x.
","pubmedId":"","doi":"10.1038/s41467-021-24505-x"},{"reference":"Botero V, Tomchik SM. 2024. Unraveling neuronal and metabolic alterations in neurofibromatosis type 1. Journal of Neurodevelopmental Disorders 16: 10.1186/s11689-024-09565-6.
","pubmedId":"","doi":"10.1186/s11689-024-09565-6"},{"reference":"Brown EB, Zhang J, Lloyd E, Lanzon E, Botero V, Tomchik S, Keene AC. 2023. Neurofibromin 1 mediates sleep depth in Drosophila. PLOS Genetics 19: e1011049.
","pubmedId":"","doi":"10.1371/journal.pgen.1011049"},{"reference":"Buchberg AM, Cleveland LS, Jenkins NA, Copeland NG. 1990. Sequence homology shared by neurofibromatosis type-1 gene and IRA-1 and IRA-2 negative regulators of the RAS cyclic AMP pathway. Nature 347: 291-294.
","pubmedId":"","doi":"10.1038/347291a0"},{"reference":"Carvalho GB, Kapahi P, Benzer S. 2005. Compensatory ingestion upon dietary restriction in Drosophila melanogaster. Nature Methods 2: 813-815.
","pubmedId":"","doi":"10.1038/nmeth798"},{"reference":"Collins NJ, Endres MN, Sinakevitch IT, Shao L. 2026. Protocol for quantifying Drosophila feeding behavior using flyPAD and optoPAD. STAR Protocols 7: 104711.
","pubmedId":"","doi":"10.1016/j.xpro.2026.104711"},{"reference":"Itskov PM, Moreira JM, Vinnik E, Lopes Ga, Safarik S, Dickinson MH, Ribeiro C. 2014. Automated monitoring and quantitative analysis of feeding behaviour in Drosophila. Nature Communications 5: 10.1038/ncomms5560.
","pubmedId":"","doi":"10.1038/ncomms5560"},{"reference":"Ja WW, Carvalho GB, Mak EM, de la Rosa NN, Fang AY, Liong JC, Brummel T, Benzer S. 2007. Prandiology of\n Drosophila\n and the CAFE assay. Proceedings of the National Academy of Sciences 104: 8253-8256.
","pubmedId":"","doi":"10.1073/pnas.0702726104"},{"reference":"King, R. C., & Wilson, L. P. (1955). Studies with radiophosphorus in Drosophila. V. The phosphorus balance of adult females. Journal of Experimental Zoology, 130(1), 71–82.
","pubmedId":"","doi":""},{"reference":"Mair W, Piper MDW, Partridge L. 2005. Calories Do Not Explain Extension of Life Span by Dietary Restriction in Drosophila. PLoS Biology 3: e223.
","pubmedId":"","doi":"10.1371/journal.pbio.0030223"},{"reference":"Shimada I, Nakao M, Kawazoe Y. 1987. Acute differential sensitivity and role of the central nervous system in the feeding behavior of Drosophila melanogaster. Chemical Senses 12: 481-490.
","pubmedId":"","doi":"10.1093/chemse/12.3.481"},{"reference":"Sofela FA, Lopez Valencia M, Jongens TA, Sehgal A. 2024. Effects of\n Nf1\n on sleep behavior are mediated through starvation caused by deficits in SARM1 dependent NAD+ metabolism. : 10.1101/2024.09.14.612058.
","pubmedId":"","doi":"10.1101/2024.09.14.612058"},{"reference":"Tanimura T, Isono K, Takamura T, Shimada I. 1982. Genetic dimorphism in the taste sensitivity to trehalose inDrosophila melanogaster. Journal of Comparative Physiology ? A 147: 433-437.
","pubmedId":"","doi":"10.1007/BF00612007"},{"reference":"The I, Hannigan GE, Cowley GS, Reginald S, Zhong Y, Gusella JF, Hariharan IK, Bernards A. 1997. Rescue of a Drosophila NF1 mutant phenotype by protein kinase A. Science 276(5313): 791-4.
","pubmedId":"9115203","doi":""},{"reference":"Walker JA, Tchoudakova AV, McKenney PT, Brill S, Wu D, Cowley GS, Hariharan IK, Bernards A. 2006. Reduced growth of Drosophila neurofibromatosis 1 mutants reflects a non-cell-autonomous requirement for GTPase-Activating Protein activity in larval neurons. Genes Dev 20(23): 3311-23.
","pubmedId":"17114577","doi":""},{"reference":"Wong R, Piper MDW, Wertheim B, Partridge L. 2009. Quantification of Food Intake in Drosophila. PLoS ONE 4: e6063.
","pubmedId":"","doi":"10.1371/journal.pone.0006063"},{"reference":"Xu G, O'Connell P, Viskochil D, Cawthon R, Robertson M, Culver M, et al., Weiss. 1990. The neurofibromatosis type 1 gene encodes a protein related to GAP. Cell 62: 599-608.
","pubmedId":"","doi":"10.1016/0092-8674(90)90024-9 "}],"title":"Neurofibromin 1 alters feeding microstructure in Drosophila
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Steven Marygold"},"openAcknowledgement":false,"submitted":null}]},{"id":"915b4c70-f0df-4ba5-86d7-23a3fba9f27e","decision":"publish","abstract":"Neurofibromatosis type 1 (NF1) is caused by the loss of neurofibromin, a protein shown to regulate metabolism and feeding behavior in Drosophila melanogaster. Using the FlyPad two-choice feeding assay, we compared feeding microstructure between NF1 null mutants, NF1 heterozygotes, and wildtype controls. We find that NF1 mutants took significantly more sips of sucrose and with greater urgency, without changes in sip duration or sucrose taste sensitivity. Overall, these results suggest loss of NF1 increases feeding frequency.
","acknowledgements":"We want to thank Dr. Seth Tomchik and members of the Brown Lab for helpful discussions and constructive feedback, and Dr. Pavel Itskov for technical assistance. We also extend our appreciation to Dr. Barbara Shoplock and the FSU Young Scholars Program for academic and research support.
","authors":[{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","writing_originalDraft","writing_reviewEditing"],"email":"ZP23@fsu.edu","firstName":"Zohre","lastName":"Panahidizjikan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000-0001-8719-5266"},{"affiliations":["Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences"],"credit":["formalAnalysis","investigation","project","visualization","validation","writing_reviewEditing"],"email":"Sjb21q@fsu.edu","firstName":"Sydney","lastName":"Born","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0009-0008-7067-8876 "},{"affiliations":["Miami Palmetto Senior High School, Pinecrest, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"maiaespinoza11@gmail.com","firstName":"Maia","lastName":"Espinoza","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Magnet Innovation Center, Inlet Beach, FL, USA","Florida State University, Tallahassee, FL, United States"],"departments":["","Young Scholars Program"],"credit":["formalAnalysis","investigation","visualization","writing_reviewEditing"],"email":"starcatcherfour@gmail.com","firstName":"Tate","lastName":"Lillian","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":true,"WBId":null,"orcid":null},{"affiliations":["Florida State University, Tallahassee, FL, United States","Florida State University, Tallahassee, FL, United States"],"departments":["Biological Sciences","Program in Neuroscience"],"credit":["conceptualization","formalAnalysis","fundingAcquisition","methodology","project","resources","supervision","visualization","writing_originalDraft","writing_reviewEditing"],"email":"EBBrown@fsu.edu","firstName":"Elizabeth B","lastName":"Brown","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0000-0003-0825-8199"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for this work was provided by the National Institutes of Health (R00AG071833 to EBB) and FSU Office of Research, as well as the FSU Department of Biological Science Lamar and Edith Trott Endowed Scholarship to ZP.
","image":{"url":"https://portal.micropublication.org/uploads/6dbb1b22553139c36cdc94a56fdfb0e7.png"},"imageCaption":"(A) Schematic of the FlyPad two-choice feeding assay, in which individual flies are offered simultaneous access to 1mM and 5mM sucrose. (B) There is a significant effect of genotype on sip number (H test: H2 = 23.54, P<0.0001). Loss of NF1 significantly increases sip number (+, P<0.0001; het, P<0.0085). (C) There is a significant effect of genotype on sip number when binned by sucrose concentration (REML: F2,268 = 11.00, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3706; het, P<0.3915), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0149). (D) Loss of NF1 has no effect on preference for 5mM sucrose (H test: H2 = 4.470, P<0.1070). (E) There is no effect of genotype on sip duration (REML: F2,134 = 2.895, P<0.0588). (F) There is no effect of genotype on intersip interval (REML: F2,132 = 1.253, P<0.2890). (G) There is a significant effect of genotype on feeding intensity (REML: F2,260 = 13.25, P<0.0001). Loss of NF1 has no effect on sips of 1mM sucrose (+, P<0.3399; het, P<0.5110), but significantly increases sips of 5mM sucrose (+, P<0.0001; het, P<0.0001). (H-J) Distribution of the durations of sips of sucrose in (H) wildtype flies, (I) NF1 heterozygous flies, and (J) NF1 mutant flies. Error bars indicate ± SEM. *p<0.05; **p<0.01; ****p<0.0001.
","imageTitle":"Loss of NF1 increases food intake
","methods":"Drosophila care and maintenance
Flies were maintained on standard cornmeal-molasses-yeast food (Bloomington Recipe, Genesee Scientific, San Diego, CA) at 25°C and at ~50% relative humidity under a 12 h light: 12 h dark cycle. Mated ~3–5-day-old male flies were collected under light CO2 anesthesia and allowed to recover for at least 24 h prior to all experiments.
FlyPAD two-choice feeding assay
Feeding behavior was quantified using the flyPAD (fly Proboscis and Activity Detector; Itskov et al., 2014). Prior to testing, flies were starved for 18 h in vials on wet Kimwipe paper. On the day of testing, individual males were aspirated without anesthesia into flyPAD arenas configured for a two-choice assay, with the two food channels containing either 1 mM or 5 mM sucrose in 1% agarose. Flies were allowed to feed for 60 min, after which data were extracted and analyzed using established flyPAD analysis software (Itskov et al., 2014). The Preference Index (PI) was calculated as: (# sips of 5 mM sucrose − # sips of 1 mM sucrose) / (total # of sips), where a value of 1.0 indicates complete preference for 5mM sucrose and a value of -1.0 indicates a complete preference for 1mM sucrose. Feeding Intensity was calculated as: # sips / sip duration (Collins et al., 2026). Flies that did not feed were excluded from downstream analyses. The resulting sample size for this experiment were as follows: controls (N = 43), NF1P1 heterozygotes (N = 39), and NF1P1 mutants (N = 55). All experiments were performed between ZT 0-2 at 25°C.
Statistical analysis
Feeding metrics are visualized as graphs displaying mean ± standard error. Dots represent individual fly measurements. Four independent runs were performed. The data were not normally distributed and so the non-parametric Kruskal-Wallis test was used to compare genotypes. To compare genotypes and sucrose concentration, a restricted maximum likelihood (REML) estimation was used. All post hoc analyses were performed using Dunn’s multiple comparisons test. Data were analyzed and visualized using InStat software (GraphPad Software 11.0).
","reagents":"Drosophila lines
The control line, w1118 was obtained from the Bloomington Stock Center (#5905). The NF1P1 line was a gift from Seth Tomchik’s lab at the University of Iowa (The et al., 1997). This line was backcrossed to the w1118 laboratory strain for 10 generations prior to testing.
","patternDescription":"Neurofibromatosis type 1 (NF1) is a neurodevelopmental disorder caused by mutations in the NF1 gene. This gene encodes neurofibromin, a GTPase-activating protein that negatively regulates the RAS signaling pathway (Buchberg et al., 1990; Xu et al., 1990). In humans, mutations in NF1 are associated with disruptions in several physiological and behavioral traits, including body size, metabolic function, and behavior (Botero & Tomchik, 2024). Many aspects of this disorder can be phenocopied using the fruit fly, Drosophila melanogaster: NF1-deficient flies have reduced body size, elevated metabolic rate, and reduced triglyceride stores (Botero et al., 2021, 2026; Brown et al., 2023; Sofela et al., 2024; The et al., 1997; Walker et al., 2006). Notably, loss of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024), consistent with a homeostatic response to their elevated metabolic rate and depleted energy stores. In this study, we further characterized the feeding behavior of NF1 flies.
Several approaches have been developed to quantify feeding behavior in Drosophila. These include the capillary feeder (CAFE) assay, which measures cumulative food consumption from a graduated capillary (Ja et al., 2007), manual scoring of proboscis extensions (Mair et al., 2005; Wong et al., 2009), as well as dye-labeling and radioisotope-labeling based approaches that estimate ingestion from colorimetric food additives or radiolabeled food, respectively (Carvalho et al., 2005; King & Wilson, 1955; Shimada et al., 1987; Tanimura et al., 1982). These approaches provide robust measures of total intake but lack the temporal and behavioral resolution to resolve individual feeding events. We therefore used the flyPAD (fly Proboscis and Activity Detector), which can detect each contact between a fly’s proboscis and a food source as a discrete change in capacitance, thereby enabling real-time, high-resolution quantification of feeding microstructure including sip number, duration, and choice between food sources in individual, freely feeding flies (Itskov et al., 2014).
Here, we utilized the flyPAD to investigate the effects of NF1 on food intake and feeding microstructure (Fig 1A). Starved male flies were provided the opportunity to feed from 1 mM and 5 mM sucrose, a paradigm that reports both feeding motivation and the ability to discriminate caloric value (Collins et al., 2026). We used the NF1P1 mutant line, which has a deletion that removes all of the NF1 gene except for the first exon (The et al., 1997). We found that when we summed the total number of sips taken from both wells over the duration of the experiment NF1P1 mutant flies take significantly more sips than both control and NF1P1 heterozygote flies (Fig 1B), consistent with previous findings that disruption of NF1 increases food intake (Botero et al., 2021; Sofela et al., 2024). We next broke down the number of sips by sucrose concentration and found that all genotypes took more sips of 5mM sucrose than of 1mM sucrose. However, NF1P1 mutant flies consistently took more sips of 5mM sucrose relative to control and NF1P1 heterozygote flies (Fig 1C). To evaluate the possibility that NF1P1 mutant flies are more sensitive to 5mM sucrose, we calculated a Preference Index for 5mM sucrose, where a positive value indicates stronger preference for 5mM sucrose and a negative value indicate stronger preference for 1mM sucrose. We found that preference for 5mM sucrose was consistent across all genotypes (Fig 1D), suggesting taste sensitivity to sucrose does not mediate the increased sips taken by NF1P1 flies.
In wild-type flies, food deprivation alters the microstructure of feeding by increasing the total number and duration of sips, and by decreasing the length between sips (Itskov et al., 2014; Banu et al., 2023). We next investigated whether NF1 functions to regulate these additional metrics of feeding microstructure. We found that the duration of each sip was not significantly different between genotypes (Fig 1E), nor was the interval between sips (Fig 1F). Therefore, NF1P1 flies consume more 5mM sucrose by increasing the frequency of sipping, not by taking longer sips or reducing time between sips, suggesting that loss of NF1 engages a more restricted feeding-drive mechanism than food deprivation alone. This is further reflected in measurements of feeding urgency (Collins et al., 2026), where we find that NF1P1 flies have significantly higher feeding intensity on 5mM sucrose relative to control and NF1P1 heterozygotes (Fig 1G). This may be driven primarily by the observation that NF1P1 flies have a greater frequency of very short duration sips relative to control and heterozygotes (Fig 1H-J). Overall, our results suggest that loss of NF1 further increases feeding frequency without altering the duration of individual sipping episodes or the ability to discriminate between sucrose concentrations, consistent with an increase in feeding drive.
","references":[{"reference":"Banu A, Gowda SBM, Salim S, Mohammad F. 2023. Serotonergic control of feeding microstructure in Drosophila. Frontiers in Behavioral Neuroscience 16: 10.3389/fnbeh.2022.1105579.
","pubmedId":"","doi":"10.3389/fnbeh.2022.1105579"},{"reference":"Botero V, Barrios J, Knauss A, Dahlen G, Rosendahl E, Colodner KJ, Tomchik SM. 2026. Metabolic and behavioral effects of neurofibromin result from differential recruitment of MAPK and mTOR signaling. PLOS Genetics 22: e1012061.
","pubmedId":"","doi":"10.1371/journal.pgen.1012061"},{"reference":"Botero V, Stanhope BA, Brown EB, Grenci EC, Boto T, Park SJ, et al., Tomchik. 2021. Neurofibromin regulates metabolic rate via neuronal mechanisms in Drosophila. Nature Communications 12: 10.1038/s41467-021-24505-x.
","pubmedId":"","doi":"10.1038/s41467-021-24505-x"},{"reference":"Botero V, Tomchik SM. 2024. Unraveling neuronal and metabolic alterations in neurofibromatosis type 1. Journal of Neurodevelopmental Disorders 16: 10.1186/s11689-024-09565-6.
","pubmedId":"","doi":"10.1186/s11689-024-09565-6"},{"reference":"Brown EB, Zhang J, Lloyd E, Lanzon E, Botero V, Tomchik S, Keene AC. 2023. Neurofibromin 1 mediates sleep depth in Drosophila. PLOS Genetics 19: e1011049.
","pubmedId":"","doi":"10.1371/journal.pgen.1011049"},{"reference":"Buchberg AM, Cleveland LS, Jenkins NA, Copeland NG. 1990. Sequence homology shared by neurofibromatosis type-1 gene and IRA-1 and IRA-2 negative regulators of the RAS cyclic AMP pathway. Nature 347: 291-294.
","pubmedId":"","doi":"10.1038/347291a0"},{"reference":"Carvalho GB, Kapahi P, Benzer S. 2005. Compensatory ingestion upon dietary restriction in Drosophila melanogaster. Nature Methods 2: 813-815.
","pubmedId":"","doi":"10.1038/nmeth798"},{"reference":"Collins NJ, Endres MN, Sinakevitch IT, Shao L. 2026. Protocol for quantifying Drosophila feeding behavior using flyPAD and optoPAD. STAR Protocols 7: 104711.
","pubmedId":"","doi":"10.1016/j.xpro.2026.104711"},{"reference":"Itskov PM, Moreira JM, Vinnik E, Lopes Ga, Safarik S, Dickinson MH, Ribeiro C. 2014. Automated monitoring and quantitative analysis of feeding behaviour in Drosophila. Nature Communications 5: 10.1038/ncomms5560.
","pubmedId":"","doi":"10.1038/ncomms5560"},{"reference":"Ja WW, Carvalho GB, Mak EM, de la Rosa NN, Fang AY, Liong JC, Brummel T, Benzer S. 2007. Prandiology of\n Drosophila\n and the CAFE assay. Proceedings of the National Academy of Sciences 104: 8253-8256.
","pubmedId":"","doi":"10.1073/pnas.0702726104"},{"reference":"King, R. C., & Wilson, L. P. (1955). Studies with radiophosphorus in Drosophila. V. The phosphorus balance of adult females. Journal of Experimental Zoology, 130(1), 71–82.
","pubmedId":"","doi":""},{"reference":"Mair W, Piper MDW, Partridge L. 2005. Calories Do Not Explain Extension of Life Span by Dietary Restriction in Drosophila. PLoS Biology 3: e223.
","pubmedId":"","doi":"10.1371/journal.pbio.0030223"},{"reference":"Shimada I, Nakao M, Kawazoe Y. 1987. Acute differential sensitivity and role of the central nervous system in the feeding behavior of Drosophila melanogaster. Chemical Senses 12: 481-490.
","pubmedId":"","doi":"10.1093/chemse/12.3.481"},{"reference":"Sofela FA, Lopez Valencia M, Jongens TA, Sehgal A. 2024. Effects of\n Nf1\n on sleep behavior are mediated through starvation caused by deficits in SARM1 dependent NAD+ metabolism. : 10.1101/2024.09.14.612058.
","pubmedId":"","doi":"10.1101/2024.09.14.612058"},{"reference":"Tanimura T, Isono K, Takamura T, Shimada I. 1982. Genetic dimorphism in the taste sensitivity to trehalose inDrosophila melanogaster. Journal of Comparative Physiology ? A 147: 433-437.
","pubmedId":"","doi":"10.1007/BF00612007"},{"reference":"The I, Hannigan GE, Cowley GS, Reginald S, Zhong Y, Gusella JF, Hariharan IK, Bernards A. 1997. Rescue of a Drosophila NF1 mutant phenotype by protein kinase A. Science 276(5313): 791-4.
","pubmedId":"9115203","doi":""},{"reference":"Walker JA, Tchoudakova AV, McKenney PT, Brill S, Wu D, Cowley GS, Hariharan IK, Bernards A. 2006. Reduced growth of Drosophila neurofibromatosis 1 mutants reflects a non-cell-autonomous requirement for GTPase-Activating Protein activity in larval neurons. Genes Dev 20(23): 3311-23.
","pubmedId":"17114577","doi":""},{"reference":"Wong R, Piper MDW, Wertheim B, Partridge L. 2009. Quantification of Food Intake in Drosophila. PLoS ONE 4: e6063.
","pubmedId":"","doi":"10.1371/journal.pone.0006063"},{"reference":"Xu G, O'Connell P, Viskochil D, Cawthon R, Robertson M, Culver M, et al., Weiss. 1990. The neurofibromatosis type 1 gene encodes a protein related to GAP. Cell 62: 599-608.
","pubmedId":"","doi":"10.1016/0092-8674(90)90024-9 "}],"title":"Neurofibromin 1 alters feeding microstructure in Drosophila
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