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Book Chapters

      1. Bhandari, R.K. (2023). Epigenetic inheritance in aquatic organisms (Chapter 4: 95-115). Epigenetics in Aquaculture. Edited by Drs. Francesc Piferrer and Hanping Wang. 
      2. vom Saal F.S., Parmigiani S., Palanza P., and Bhandari R.K. (2019). Transgenerational effects on fertility of estrogenic chemicals in fish. In: F. Ruggiu, Editor, Nuclear War, and Planetary Emergencies, World Scientific Publishers, Singapore. pp. in press.
      3. Skinner, M.K., Nilsson, E.E., and Bhandari, R.K. (2010). Cell-Cell Signaling in the Testis and Ovary. In: Handbook of Cell Signaling Third Edition (Ralph A Bradshaw and Edward Dennis, Eds.) Academic Press/Elsevier Science Publishing. pp. 2663-2678.

Peer-reviewed Manuscripts (student authors- *undergraduate, ** graduate, ‡Postdoc )

2023 (in progress)

      1. **Chakraborty, S., Anand, S., ‡Faheem, M., Bhandari, R.K. (2023). Integrated metabolomic and transcriptional analyses revealing sex-specific alterations in the liver caused by ancestral bisphenol A consumption.
      2. **Fricault, F.J., Wang, X., Aravamudhan, S., Bhandari, R.K. (2023). Effects of aluminum oxide and cerium oxide nanoparticles on human lung alveolar cells in vitro.
      3. Bhandari, R.K. and Nakamura, M. (2023). Synthetic androgen suppressed the female reproductive axis in tilapia.
      4. **Doldron, M.S., ‡Wang, X., Jia, Z., Bhandari, R.K. (2023). Effects of delta-9-tetrahydrocannabinol exposure on human bronchial epithelial cells in vitro.
      5. Bhandari, R.K., ‡Wang, X., vom Saal, F.S., Tillitt, D.E. (2023)  Transgenerational inheritance of environmentally induced DNA methylation patterns in primordial germ cells by testicular somatic cells.
      6. **Feng, Y., ‡Wang, X., *Vassall, M., Bhandari, R.K. (2023). Delta 9-THC induced transcriptional alterations in the brain and gonads of medaka.
      7. **Feng, Y., ‡Wang, X., Bhandari, R.K. (2023). Sperm methylome predicts the effects of paternal preconception THC exposure on offspring health.
      8. **Guzman, E., ‡Wang, X., Bhandari, R.K. (2023). Bisphenol A-induced transgenerational dysbiosis of gut microbiota in medaka.
      9. *Weidman, D., *Mathewson, A., Bhandari, R.K.(2023). Mitigation of glyphosate-induced developmental defects in medaka embryos by vitamin-C co-treatments.
      10. Wang X.,Song, X., Bhandari, R.K. (2023). miRNA expression dynamics during epigenetic reprogramming of primordial germ cells.
      11. Wang, X., Bhandari, R.K. (2023). Environmental chemical induced single nucleotide polymorphism in germ cells and sperm.
      12. Wang, X., Bhandari, R.K. (2023). A dynamic pattern of inheritance of epigenetic memory by germline cells and soma across generations.
      13. Wang, X., Feng. Y., Bhandari, R.K. (2023). Aberrant genome DNA methylation in sperm persisted for eight years.




      1. **Chakraborty, S., ‡Anand, S., **Coe, S.T., *Reh, B., Bhandari, R.K. (2023). The PCOS–NAFLD Multi-disease phenotype occurred in medaka fish four generations after the removal of bisphenol A exposure. Environ. Sci. Technol. 57 (34), 12602-12619.
      2. **Chakraborty, S., *Dissanayake, M., *Godwin, J., ‡Wang, X., Bhandari, R.K. (2023). Ancestral BPA exposure caused defects in the liver of medaka for four generations. Science of the Total Environment.856 (1):159067.
      3. **Fatema, K., Auditi, T.I., Biswas, S., Binte, S.A., Uddin, M.H., Sumon, K.A., Goswami, C., Bhandari, R.K., Rashid, H. (2023). Investigations of hemato-biochemical and histopathological parameters, and growth performance of walking catfish (Clarias batrachus) exposed to PET and LDPE microplastics.Environmental Toxicology and Pharmacology, 102: 104250
      4. Liang, F. Dong, Z., Ye, J., Hu, W., Bhandari, R.K., Mai, K., Wang, X. (2023). In vivo DNA methylation editing in zebrafish. Epigenetics 18 (1), 2192326.
      5. Bhandari, R.K. (2023). Epigenetic Inheritance in Aquatic Organisms. In Epigenetics in Aquaculture (eds F. Piferrer and H.-P. Wang).
      6. Liang, Z.; Guo, Y.; Pi, D.; Li, X.; Li, B.; Huang, Y.; Song, X.; Bhandari, R.K.; Wang, X. (2023). Transcriptome Analysis of the Developmental Effects of Bisphenol F Exposure in Chinese Medaka (Oryzias sinensis). International Journal of Molecular Sciences. 24, 10898.
      7. *Vassall, M.; **Chakraborty, S.; **Feng, Y.; ‡Faheem, M.; ‡Wang, X.; Bhandari, R.K. (2023). Transcriptional Alterations Induced by Delta-9 Tetrahydrocannabinol in the Brain and Gonads of Adult Medaka. Journal of  Xenobiotics. 2023, 13, 237-251.
      8. *Killian, D.; ‡Faheem, M.; *Reh, B.; Wang, X.; Bhandari, R.K. (2023). Effects of Chronic Roundup Exposure on Medaka Larvae. J. Xenobiot. 13: 500–508. jox13030032.


      1. ‡Lee, Y.H., Kim, M-S, Wang, M., Park, H.G., Bhandari, R.K., Wu, R.S.S., Lee, J-S. (2022). Epigenetic adaptation to ocean acidification in copepods. Nature Climate Change: S41558
      2. *Reh, B., **Feng, Y., ‡Wang, X., Bhandari, R.K. (2022). Effects of potassium perchlorate exposure on primordial germ cells of medaka fish. , Aquatic Toxicology): 106283.
      3. **Hasan M., Sumon, K.A., Siddiquee, M.M., Bhandari, R.K., Hossain, M.D., Rashid, H. (2022). Thiamethoxam affects the developmental stages of banded gourami (Trichogaster fasciata). Toxicology Reports 9: 1233-1239
      4. **Numan, M., Guo, W., Choi, S.L., ‡Wang, X., Baldrich, P., Du, B., Jin, W.., Bhandari, R.K.,, Mayers, B., Ligaba-Osena, A. (2022). Analysis of miRNAs responsive to long-term Calcium deficiency in tef (Eragrostis tef (Zucc.) trotter). Plant Direct. 6(5): e400.
      5. Nakamura, M., Nozu, R., Nakamura, S., Higa, M.,  Bhandari,  R.K., Kobayashi, Y., Horiguchi, R., Komatsu, R., Kojima, Y., Murata, R., Soyano, K., Ogawa, S., Hirai, T., Matsubara, H., Tokumoto, T.,  Kobayashi, T., Kagawa, H., Adachi, S., Yamauchi, K.,  Nagahama, Y. (2022). Morphological and physiological studies on sex change in tropical fish: Sexual plasticity of the ovaries of hermaphroditic and gonochoristic fish. Galaxea, Journal of Coral Reef Studies. 24 (1): 5-17. [J-Stage]
      6. **Rajagopal, S., Sharma, A., Simlot, A., Mathur, P., Mehta, S., Mehta, S., Naravula, J., Medicherla, K.M., Kumar S, A., Kanga, U., Bhandari, R.K., KaviKishor, P.B, Nair, B.G., Suravajala, P. (2022). Infer-ring bonafide differentially expressed genes and their variants associated with vitamin K deficiency using a systems genetics approach. Genes. (accepted)


      1. **Tatum, S., **Saudi, S., Tettey, F., Bhandari, R.K.,, Bhattarai, N.P. (2021). A novel hydrogel-bronchial epithelial cell spheroids for toxicological evaluation. Biomedical Sciences Instrumentation.57(4):407-411.
      2. Faheem, M., Bhandari, R.K. (2021 ). Detrimental effects of bisphenol compounds on physiology and reproduction in fish. Environmental Toxicology and Pharmacology 81: 103497 [PMID: 32950715]


      1. ‡Wang, X., Bhandari, R.K.,(2020 ). The Dynamics of DNA methylation during epigenetic reprogramming of primordial germ cells in medaka (Oryzias latipes), Epigenetics 15: 483-498 [PMID: 31851575]
      2. ‡Wang, X., Bhandari, R.K., (2020 ). DNA methylation reprogramming in medaka fish, a promising animal model for environmental epigenetics research. Environmental Epigenetics.6: dvaa008.
      3. **Song, X., ‡Wang, X., Bhandari, R.K. (2020). Developmental abnormalities and epigenetic effects in medaka (Oryzias latipes) embryo induced by triclosan exposure. Chemosphere.261: 127613
      4. Bhandari, R.K., ‡Wang, X., vom Saal, F.S., Tillitt, D.E. (2020). Transcriptome analysis of testis reveals the effects of developmental exposure to bisphenol A or 17α-ethinylestradiol in medaka. Aquatic Toxicology. 225: 105553
      5. Leet, J.K., Richter, C.A., Cornman, R.S., Behringer, J.P., Bhandari, R.K., Nicks, D.K.,  Candrl, J.S, Zajicek, J.L., Blazer, V.S., Tillitt, D.E. (2020). Effects of early life stage exposure of largemouth bass to atrazine or a model estrogen (17α-ethinylestradiol). PeerJ. 8:e9614
      6. ‡Wang, X., **Song, X., Bhandari, R.K. (2020). Distinct expression patterns of seven microRNAs during early embryonic development in medaka (Oryzias latipes).Gene Expression Patterns. 37: 119133
      7. *Thayil, A.J., ‡Wang, X., Bhandari, P., vom Saal, F.S., Tillitt, D.E., Bhandari, R.K. (2020). Bisphenol A and 17 alpha-ethinylestradiol-induced Transgenerational Gene Expression Differences in the Brain-Pituitary-Testis Axis of Medaka, Oryzias latipes. Biology of Reproduction. 103(6): 1324–1335
      8. Faheem, M., Bhandari, R.K. (2020). Detrimental effects of bisphenol exposure on physiology and reproduction of fish. A literature review. Environmental Toxicology and Pharmacology.81: 103497


      1. ‡Wang. X., Bhandari. R.K. (2019). DNA methylation dynamics during epigenetic reprogramming of medaka embryo. Epigenetics. 14(6):611-622. [PMID: 31010368]
      2. ‡Wang, X., *Hill, D., Tillitt, D.E., Bhandari, R.K (2019). Bisphenol A and 17alpha-ethinylestradiol induced transgenerational differences in expression of osmoregulatory genes in the gill of medaka (Oryzias latipes). Aquatic Toxicology. 211: 227-234. [PMID: 31048106]
      3. **Smith, C.S., *Vera, M.K.M., Bhandari, R.K. (2019). Developmental and epigenetic effects of Roundup and glyphosate exposure on medaka (Oryzias latipes). Aquatic Toxicology, 210:215-226. [PMID: 30875550]
      4. **Cleary, J.C., Tillitt, D.E., vom Saal, F.S., Nicks, D.K., Claunch, R.A, Bhandari, R.K. (2019). Transgenerational effects of developmental Atrazine exposure on reproductive axis of medaka. Environmental Pollution, 251: 631-650. [PMID: 31108297]
      5. Bhandari, R.K., Taylor, J.A., *Sommerfeld-Sager, J.,  Tillitt, D.E., Ricke, W.A., vom Saal, F.S. (2019). Estrogen receptor 1 expression and methylation of Esr1 promoter in mouse fetal prostate mesenchymal cells induced by gestational exposure to bisphenol A or ethinylestradiol. Environmental Epigenetics, 5(3): dvz012. doi: 10.1093/eep/dvz012 [PMID: PMC6705189].
      6. **Khanal, S., Bhattarai, S.R., Sarkar, J., Bhandari, R.K., McDonald, J., Bhattarai, N.P. (2019). Nano-fiber integrated microcapsules: A nano-in-micro platform for 3D cell culture. Scientific Reports. [PMID: 31562351]


      1. **Jandegian CM, Deem SL, Bhandari RK, Holliday CM, Nicks D, Rosenfeld CS, Selcer KW, Tillitt DE, Vom Saal FS, Vélez-Rivera V, Yang Y, Holliday DK. (2017). Corrigendum to “Developmental exposure to bisphenol A (BPA) alters sexual differentiation in painted turtles (Chrysemys picta)” [Gen. Comp. Endocr. 216 (2015) 77-85]. General and Comparative Endocrinology. 247:223. [PMID: 28454885]


      1. Bhandari, R.K. (2016). Medaka as a model for studying environmentally induced transgenerational inheritance phenotypes.Environmental Epigenetics 2:1-9. (Invited perspective) [PMID: 29492282]


      1. Bhandari R.K., vom Saal, F.S., Tillitt, D. E.  (2015). Transgenerational effects from early developmental exposures to bisphenol A or 17α-ethinylestradiol in medaka, Oryzias latipesScientific Reports, 5 : 9303. [PMID: 25790734]
      2. Bhandari RK, Deem SL, Holliday DK, Jandegian CM, Kassotis CD, Nagel SC, Tillitt DE, Vom Saal FS, Rosenfeld CS (2015). Effects of the environmental estrogenic contaminants bisphenol A and 17α-ethinylestradiol on sexual development and adult behaviors in aquatic wildlife species. General and Comparative Endocrinology, 214:195-219. [PMID: 25277515]
      3. Skinner, M.K., Bhandari, R.K., Haque, M.M., Nilsson, E.E. (2015). Environmentally induced epigenetic transgenerational inheritance of altered SRY genomic binding during gonadal sex determination. Environmental Epigenetics 1: 1-10.  doi: 10.1093/eep/evv004 [PMID: 27175298]
      4. **Pandit, N.P., Bhandari, R.K., Kobayashi, Y., Nakamura, M. (2015). High temperature-induced Sterility in the Female Nile Tilapia,Oreochromis niloticus. General and Comparative Endocrinology, 213:110-117. [PMID: 25745814]
      5. **Jandegian, C.M., Deem, S.L, Bhandari, R.K., Holliday, C.M., Nicks, D., Rosenfeld, C.S., vom Saal, F.S., Tillitt, D.E., Vélez-Rivera, V, Selcer, K.W, Yang, Y, and Holliday, D.K. (2015). Developmental exposure to bisphenol A disrupts sexual differentiation in painted turtles (Chrysemys picta). General and Comparative Endocrinology, 216:77-85. [PMID: 25863134]


      1. Skinner MK, Guerrero-Bosagna C, Haque M, Nilsson E, Bhandari RK, McCarrey JR (2013) Environmentally induced transgenerational epigenetic reprogramming of primordial germ cells and the subsequent germ line. PLoS One. 8(7):e66318. [PMID: 23869203]
      2. Paul-Prasanth B*, Bhandari RK*, Kobayashi T, Horiguchi R, Kobayashi Y, Nakamoto M, Shibata Y, Sakai F, Nakamura M, Nagahama Y. (2013). Estrogen oversees the maintenance of the female genetic program in terminally differentiated gonochorists. Scientific Reports, 2013 Oct 7; 3:2862. (*Equally contributing first authors)
      3. *Miura, S., Bhandari, R.K., Kobayashi, Y. and Nakamura, M. (2013). Estrogen favors differentiation of ovary in the ambisexual gonads of Anemone fish. J Experimental Zoology: A Ecology Genetics Physiology, 319(10):560-568. [PMID: 24039239]


      1. Bhandari, R.K., Schinke, E.N., Haque, MM, and Skinner, M.K. (2012). SRY-induced TCF21 genome-wide targets and cascade of bHLH factors during Sertoli cell differentiation and male sex determination in the rat. Biology of Reproduction, 87: 131-136. [PMID: 23034159]
      2. Bhandari, R.K., Haque, M.M., and Skinner, M.K. (2012). Global genome analysis of the downstream binding targets of testis determining factor SRY and SOX9. PLoS One, 7(9): e43380. [PMID: 22984422]


      1. Bhandari, R.K., Clement, T., Sadler-Riggleman, I., and Skinner, M.K. (2011). Basic helix-loop- helix transcription factor Tcf21 is a downstream target of male sex determining gene Sry. PLoS One, 6(5): e19935. pp1-11. [PMID: 21637323]
      2. Clement, T*., Bhandari, R.K.*, Riggleman, I., and Skinner, M.K. (2011). Sry directly regulates Neurotrophin-3 promoter during male sex determination and testis development. Biology of Reproduction, 85: 277-284. (*Equally contributing first authors). [PMID: 21508350]


      1. **Westring, C.G., Ando, H., Kitahashi,  T., Bhandari, R.K., Ueda, H., Urano, A., Dores, R.M.,  Sher, A.A. and Danielson, P.B. (2008). Seasonal changes in CRF-I and urotensin I transcript levels  in masu salmon:  Correlation with cortisol secretion during spawning. General and Comparative Endocrinology, 155: 126-140.
      2. *Kojima, Y.  Bhandari, R.K.,  Kobayashi, Y.  and Nakamura M. (2008).  A male-to-female sex reversal of the primary male wrasse with estrogen treatments. General and Comparative Endocrinology, 156:628-632. (Undergraduate student)


      1. **Nakamura M.  Alam, M.A., Kobayashi, Y. and Bhandari, R.K.   (2007). Roles of sex hormones in sex change of grouper,Epinephelus merra. Journal of Marine Science and Technology-Taiwan, Special issue 15: 23-27.


      1. Bhandari, R.K., Alam, M.A., Soyano K., and Nakamura, M., (2006). Induction of female-to-male sex change in the honeycomb grouper (Epinephelus merra) by 11-ketotestosterone treatments. Zoological Science, 23: 65-69.
      2. Bhandari, R.K., Nakamura, M., Kobayashi, T., and Nagahama, Y. (2006). Suppression of steroidogenic  enzyme  expression  during  androgen-induced  sex  reversal  in  Nile  tilapia (Oreochromis niloticus). General and Comparative Endocrinology, 145: 20-24.
      3. *Komatsu, T. Bhandari, R.K., Nakamura, M.(2006). Acceleration of precocious spermatoenesis in under-yearling Golden Rabbitfish (Siganus guttatus, Bloch) by GnRHa treatments. Aquaculture, 257: 558-565. (Ph.D. Student)
      4. *Alam, M.A., Bhandari, R.K., Kobayashi, Y., Soyano K., and Nakamura, M., (2006). Induction of sex change within two full moons during breeding season and spawning in grouper. Aquaculture, 255: 532-535. (Ph.D. Student)
      5. *Alam, M.A., Bhandari, R.K., Kobayashi, Y., Soyano, K., and Nakamura, M. (2006). Changes in androgen-producing cell size and circulating 11-ketotestosterone levels during female-to- male sex change in Honeycomb grouper (Epinephelus merra). Molecular Reproduction, and Development. 73(2):206-14. (Ph.D. Student)
      6. **Nakamura M., Kobayashi, Y., Miura, S. Alam, M.A., and Bhandari R.K. (2006). Sex change in coral reef fish. Fish Physiology and Biochemistry, 31: 117-122.


      1. Bhandari, R.K., Alam, M.A., Higa, M. Soyano, K., Nakamura, M. (2005). Evidence that the estradiol-17β regulates the sex change of honeycomb grouper Epinephelus  merra), a protogynous hermaphrodite fish. Journal of Experimental Zoology, 330A: 497-503.
      2. **Jodo, A.., Kitahashi, T., Taniyama, S., Bhandari, R.K., Ueda, H., Urano, A. and Ando, H. (2005). Seasonal variations in expression of five subtypes of gonadotropin-releasing hormone receptor genes in the brain of masu salmon from immaturity to spawning. Zoological Science, 22: 1331-1338.
      3. **Luo, Q., Ban, M., Ando, H., Kitahashi, T., Bhandari, R.K., McCormick, S.D. and Urano, A. (2005). Distinct effects of 4-nonylphenol and estradiol-17β on expression of estrogen receptor a gene in smolting sockeye salmon. Comparative Biochemistry and Physiology. 140c: 123-130.
      4. *Alam, M.A., Komuro, H., Bhandari, R.K., Nakamura, S., Soyano, K., and Nakamura, M. (2005).  Immunohistochemical evidence identifying the site of androgen production in the ovary of the protogynous grouper Epinephelus merra. Cell and Tissue Research 320: 323-329. (Masters Student)


      1. Bhandari, R.K., Komuro, H., Higa, M.,and Nakamura, M. (2004). Sex inversion of sexually immature honeycomb grouper (Epinephelus  merra)  by  aromatase  inhibitor.  Zoological Science, 21 (3): 305-310.
      2. Bhandari, R.K., Higa, M., Nakamura, S. and Nakamura, M.  (2004). Aromatase inhibitor induces complete sex change in a protogynous honeycomb grouper, Epinephelus merra. Molecular Reproduction and development, 67(3): 303-307.


      1. Bhandari, R.K. Komuro, H., Nakamura, S., Higa, M., and Nakamura (2003).  Gonadal restructuring and   correlative steroid hormone profiles during natural sex change in protogynous honeycomb grouper, Epinephelus merra. Zoological Science, 20: 1399-1404.
      2. Bhandari, R.K., Higa, M., Komuro, H., Nakamura, S. and Nakamura, M. (2003). Treatments with  an  aromatase  inhibitor  induces  complete  sex  change  in  protogynous  honeycomb grouper (Epinephelus merra). Fish Physiology and Biochemistry, 28: 141-142.
      3. Bhandari, R.K., Taniyama, S., Kitahashi, T., Ando, H., Yamauchi, K., Zohar, Y., Ueda, H., and Urano, A. (2003). Seasonal changes of responses to gonadotropin-releasing hormone analog in expression of growth hormone/prolactin/somatolactin genes in the pituitary of masu salmon. General and Comparative Endocrinology, 130: 55-63.
      4. *Miura, S., Komatsu, T., Higa, M., Bhandari, R.K., Nakamura, S. and Nakamura, M. (2003). Gonadal sex differentiation in protandrous anemone fish, Amphiprion clarkii.  Fish Physiology and Biochemistry, 28: 165-166. 


      1. Bhandari, R.K., Ushikoshi, I., Fukuoka, H., Koide, N., Yamauchi, K., and Ueda, H. (2002). Effects of Rhizopus extract administration on somatic growth and sexual maturation in lacustrine sockeye salmon, Oncorhynchus nerka. Fisheries Science, 68 (4): 776-782.

Publication as an undergraduate student

      1. Nahar, Z., Shah, A.K.M.A., Bhandari, R.K., Ali, M.H., Dewan, S. (2000). Effects of different feeds on growth, survival, and production of African catfish,(Clarias gariepinus, Burchell) Bangladesh Journal of Fisheries, 4: 121-126.
      2. Saha, D.K., Shah, A.K.M.A., Islam, M.S., Bhandari, R.K., and Rahman, M.S.(1999). Effects of fertilization on production of Indian major carps. Bangladesh Journal of  Fisheries 22: 33-38.