Huan Zhang

Being a molecular biologist and marine ecologist, I have been employing modern tools of multi-disciplinary science, especially molecular biology techniques, to study fish population genetics, adaptive gene evolution, phytoplankton community structure and phytoplankton-mediated processes such as predator-prey interactions at the base of the marine food web both within laboratory cultures and at sea. Collaborating with Dr. Hans Dam’s research group, I have extended my work to elucidate the molecular nature of phytoplankton toxin adaptation in marine grazers. We focus on the adaptation of copepods to paralytic shellfish poisoning toxins (Na+ channel blockers STX, TTX). We have discovered a novel mutation in the sodium channel of the marine copepod Acartia hudsonica. This discovery opens a way for us to study the adaptation of copepods to toxic dinoflagellate Alexandrium fundyense and has profound implications for the fate of toxic phytoplankton blooms and for understanding adaptive responses of grazers.

I have also been collaborating with various other research groups at the University of Connecticut, such as Dr. Robert B. Whitlatch to develop genetic markers to study the genetic conspecificity of the worldwide populations of the putatively invasive ascidian Didemnum vexillum and determine its native region, Dr. Robert Mason to develop a mer-lux bioreporter to study the mechanisms of bacteria in the uptaking of inorganic mercury (HgII) and methylmercury (CH3HgII); Dr. George McManus to develop molecular markers for detecting ciliate diet algae; and Dr. Charles Yarish to detect the distribution of an invasive macro alga in the Northeast coast, and the macro algae as potential vectors for toxic micro algae. I have coworked with groups of the other research institutes, e.g., Dr. Wayne Litaker at NOAA to detect the geographic distribution and genetic variation of dinoflagellate Karlodinium veneficum in the US east coast, Dr. Dr. Allen Place at the Center of Marine Biotechnology, University of Maryland Biotechnology Institute to measure the genetic deviation and toxin production from various strains of K. veneficum which were potentially linked with fish-killing events. Furthermore, I have being coworking with Dr. Guangxing Liu at Ocean University of China to study marine copepod diet. Using state-of-the-art next generation sequencing techniques, we have uncovered that copepods collected from the field ate various kinds of organisms such as jellyfish larvae, eelgrass spores and many other phytoplankters, which have never being reported before.

Research Projects

Investigate a novel way of light harvesting by marine phytoplankton: proton-pump rhodopsin in dinoflagellates. – NSF

Identification of Phytoplankton Collected from Long Island Sound 2013. – CTDEP

Costs and Advantages of a novel sodium channel mutation in copepods. – NSF

Proof of concept on using spliced leader to study dinoflagellate gene expression in the ocean. – NSF


Peer-Reviewed Journal Articles (*: corresponding author)

  1. Jing X, Lin S, Zhang H, Koerting C, Yu Z. 2017. Utilization of urea and expression profiles of related genes in the dinoflagellate Prorocentrum donghaiense. PLoS ONE 12(11): e0187837.
  2. Yi X, Huang Y, Zhuang Y, Chen H, Yang F, Wang W, Xu D, Liu G, Zhang H*. 2017. In situ diet of the copepod Calanus sinicus in coastal waters of the South Yellow Sea and the Bohai Sea. Acta Oceanologica Sinica, 36(6): 68–79.
  3. Cui Y, Zhang H, Lin S. 2017. Enhancement of Non-photochemical Quenching as an Adaptive Strategy under Phosphorus Deprivation in the Dinoflagellate Karlodinium veneficum. Front. Microbiol. 8:404. doi: 10.3389/fmicb.2017.00404.
  4. Kraemer G, Yarish C, Kim JK, Zhang H, Lin S. 2017. Life history interactions between the red algae Chondrus crispus (Gigartinales) and Grateloupia turuturu (Halymeniales) in a changing global environment. Phycologia 56 (2), 176–185.
  5. Zhuang Y, Yang F, Xu D, Chen H, Zhang H*, Liu G. 2017. Spliced leader-based analyses reveal the effects of polycyclic aromatichydrocarbons on gene expression in the copepod Pseudodiaptomuspoplesia. Aquatic Toxicology 183: 114–126.
  6. Yang F, Xu D, Zhuang Y, Huang Y, Yi X, Chen H, Liu G, Zhang H. 2016. Characterization and analysis of ribosomal proteins in two marine calanoid copepods. Chinese Journal of Oceanology and Limnology 34, (6) 1258–1268.
  7. Yang F, Xu D, Zhuang Y, Yi X, Huang H, Chen H, Lin S, Campbell DA, Sturm NR, Liu G, Zhang H*. 2015. Spliced leader RNA trans-splicing discovered in copepods. Sci.Rep. 5, 17411; doi: 10.1038/srep17411.
  8. Lin S, Cheng S, Song S, Zhong X, Lin X, Li W, Li L, Zhang Y, Zhang H, et al. 2015. The Symbiodinium kawagutii genome illuminates dinoflagellate gene expression and coral symbiosis. Science 350, 691-694. DOI: 10.1126/science.aad0408.
  9. Cui Y, Lin X, Zhang H, Lin L and Lin S. 2015. PhnW-PhnX Pathway in Dinoflagellates Not Functional to Utilize Extracellular Phosphonates. Front. Mar. Sci. 2:120. doi: 10.3389/fmars.2015.00120
  10. Chen L, Zhang H, Finiguerra M, Bobkov Y, Bouchard C, Avery DE, Anderson PAV, Lin S and Dam HG. 2015. A novel mutation from gene splicing of a voltage-gated sodium channel in a marine copepod and its potential effect on channel function. JEMBE 469: 131-142.
  11. Zhuang Y, Zhang H, Hannick L, Lin S. 2015. Metatranscriptome profiling reveals versatile N-nutrient utilization, CO2 limitation, oxidative stress, and active toxin production in an Alexandrium fundyense bloom. Harmful Algae 42, 60-70.
  12. Guo Z, Zhang H, Lin S. Light-Promoted Rhodopsin Expression and Starvation Survival in the Marine Dinoflagellate Oxyrrhis marina. PLoS ONE 9(12): e114941. 2014. doi:10.1371/journal.pone.0114941
  13. WANG X-X, HUANG B-Q, ZHANG H. 2014. Phosphorus deficiency affects multiple macromolecular biosynthesis pathways of Thalassiosira weissflogii. Acta Oceanol. Sin. 33(4), 1-7. DOI: 10.1007/s13131-014-0413-x
  14. Yi XY, Zhang H and Liu GX. 2014. Primers to block the amplification of symbiotic apostome ciliate 18S rRNA gene in a PCR-based copepod diet study. Chinese Journal of Oceanology and Limnology 32(3) : 515-521. Doi:10.1007/s00343-014-3129-z
  15. Santoferrara LF, Guida S, Zhang H, McManus GB .2014. De Novo Transcriptomes of a Mixotrophic and a Heterotrophic Ciliate from Marine Plankton. PLoS ONE 9(7): e101418. doi:10.1371/journal.pone.0101418
  16. Huang Y, Xu D, Chen H, Zhang H, Lin S, Liu G. 2014. PCR-based in situ dietary analysis of Calanus sinicus Brodsky in Yellow River estuary and adjacent waters. Chinese Journal of Oceanology and Limnology 44(3): 83-91 (in Chinese with English Abstract).
  17. Wang L, Zhuang Y, Zhang H, Lin X and Lin S. 2014. DNA barcoding species in Alexandrium tamarense complex using ITS and proposing designation of five species. Harmful Algae 31:100–113.
  18. Guo Z., Zhang H., Liu S. and Lin S. 2013. Biology of the Marine Heterotrophic Dinoflagellate Oxyrrhis marina: Current Status and Future Directions. Microorganisms 1:33-57; doi:10.3390/microorganisms1010033.
  19. Zhang H*, Campbell DA, Sturm NR, Dungan CF and Lin S. 2013. Identification and characterization of signal recognition particle RNA in the dinoflagellates Karenia brevis and the perkinsid Perkinsus marinus. Protist 164: 748–761. (Published: August 28, 2013)
  20. Shi X, Zhang H, Lin S (2013) Tandem Repeats, High Copy Number and Remarkable Diel Expression Rhythm of Form II RuBisCO in Prorocentrum donghaiense (Dinophyceae). PLoS ONE 8(8): e71232. doi:10.1371/journal.pone.0071232 (Published: August 19, 2013)
  21. Qiu D, Huang L, Liu S, Zhang H, Lin S (2013) Apical Groove Type and Molecular Phylogeny Suggests Reclassification of Cochlodinium geminatum as Polykrikos geminatum. PLoS ONE 8(8): e71346. doi:10.1371/journal.pone.0071346 (published 19 Aug 2013).
  22. Zhang H*, Finiguerra M, Dam HG, Huang Y, Xu D, Liu, G and Lin S. 2013. An improved method for achieving high-quality RNA for copepod gene transcriptomic studies. J. Exp. Mar. Biol. Ecol. 446: 57–66.
  23. Castillo-Medina RE, Islas-Flores T, Thomé PE, Iglesias-Prieto R, Lin S, Zhang H and Villanueva MA. 2013. The PsbO homolog from Symbiodinium kawagutii (Dinophyceae) characterized using biochemical and molecular methods. Photosynthesis Research (DOI 10.1007/s11120-013-9856-8).
  24. Zhuang, Y., Zhang, H. and Lin, S. 2013. Polyadenylation of 18S rRNA in algae. J. Phycol. 49:570–579.
  25. Zhang H*, Zhuang Y, Gill J, and Lin S. 2013. Proof that dinoflagellate spliced leader (DinoSL) is a useful hook for fishing dinoflagellate transcripts from mixed microbial samples: Symbiodinium kawagutii as a case study. Protist 164: 510–527.
  26. Zhuang Y, Zhang H and Lin S. 2013. Cyclin B gene and its cell cycle-dependent differential expression in the toxic dinoflagellate Alexandrium fundyense Atama Group I/Clade I. Harmful Algae 26: 71–79.
  27. Kuo RC, Zhang H*, Zhuang Y, Hannick L, Lin S. 2013. Transcriptomic study reveals widespread spliced leader trans-splicing, high gene numbers and potential complex carbon fixation mechanisms in the euglenoid alga Eutreptiella sp.. PLoS ONE 8(4): e60826. doi:10.1371/journal.pone.0060826.
  28. Guo Z, Liu S, Hu S, Li T, Huang Y, Liu G, Zhang H*, Lin S. 2012. Prevalent Ciliate Epibiosis on Copepods: High Genetic Diversity and Wide Biogeography Detected by Small Subunit Ribosomal RNA Gene. PLoS ONE 7(9): e44847. doi:10.1371/journal.pone.0044847).
  29. Udonna Ndu, Robert Mason, Zhang H, Lin S, and Pieter Visscher. 2012. The Effect of Inorganic and Organic Ligands on the Bioavailability of Methylmercury as Determined by a mer-lux Bioreporter. Appl. Environ. Microbiol. 78(20):7276. DOI: 10.1128/AEM. 00362-12.
  30. Stefaniak L, Zhang H, Adriaan Gittenberger, Kirsty Smith, Kent Holsinger, Lin S and Whitlatch RB. 2012. Determining the native region of the putatively invasive ascidian Didemnum vexillum Kott, 2002. Journal of Experimental Marine Biology and Ecology 422–423: 64–71.
  31. Lin X, Zhang H, Cui Y, Lin S. 2012. High sequence variability, diverse subcellular localizations, and ecological implications of alkaline phosphatase in dinoflagellates and other eukaryotic phytoplankton. Frontiers in Aquatic Microbiology 3, Article 235 (2012). doi:10.3389/fmicb.2012.00235.
  32. Lin X, Zhang H*, Huang BQ and Lin S. 2012. Alkaline phosphatase gene sequence characteristics and transcriptional regulation by phosphate limitation in Karenia brevis (Dinophyceae). Harmful Algae 17, 14-24. doi.10.1016/j.hal.2012.02.005.
  33. Haska CL, Yarish C, Kraemer G, Blaschik N, Whitlatch R, Zhang H and Lin S. 2012. Bait worm packaging as a potential vector of invasive species. Biological Invasions. 14, 481-493.
  34. Miranda L, Zhuang Y, Zhang H* and Lin S. 2012. Phylogenetic analysis guided by intragenomic SSU rDNA polymorphism refines classification of “Alexandrium tamarense” species complex. Harmful Algae 16, 35–48. doi:10.1016/j.hal.2012.01.002.
  35. Lin X, Zhang H, Huang BQ and Lin S. 2011. Alkaline phosphatase gene sequence and transcriptional regulation by phosphate limitation in Amphidinium carterae (dinophyceae). J. Phycol. 47, 1110–1120.
  36. Zhang H*, Campbell DA, Sturm NR, Dungan CF and Lin S. 2011. Spliced leader RNAs, mitochondrial gene frameshifts and multi-protein phylogeny expand support for the genus perkinsus as a unique group of alveolates. PLoS One 6(5):e19933. doi:10.1371/journal.pone.0019933
  37. Zhang H*, Dungan CF and Lin S. 2011. Introns, Alternative Splicing, Spliced Leader trans-Splicing and Differential Expression of pcna and cyclin in Perkinsus marinus. Protist 162, 154-167. doi:10.1016/j.protis.2010.03.003).
  38. Lin S, Zhang H, Zhuang Y, Tran B, and Gill J. 2010. Spliced leader–based metatranscriptomic analyses lead to recognition of hidden genomic features in dinoflagellates. PNAS USA 107, 20033–20038. doi/10.1073/pnas.1007246107.
  39. Lin S and Zhang H. 2010. Dinoflagellate meta-transcriptomics enabled by spliced leader. In C. K. Ho (ed.) Proceedings of 13th International Conference on Harmful Algae. Pp 166-170.
  40. Hou Y, Zhang H, Miranda L and Lin S. 2010. Serious overestimation in quantitative PCR by circular (supercoiled) plasmid standard: microalgal pcna as the model gene. PLoS ONE 5(3): e9545. doi:10.1371/journal.pone.0009545.
  41. Morita M, Futami K, Zhang H, Kubokawak, Ojima Y, Okamoto N. 2009. Evolutionary analysis of amphioxus myc gene. J Tokyo Univ. Mar Sci & Tech. 5, 11-16.
  42. Zhang H, Campbell DA, Sturm NR, and Lin S. 2009. Dinoflagellates spliced leader RNA genes display a variety of sequences and genomic arrangements. Mol. Biol. Evol. 26, 1757–1771.
  43. Lin S, Zhang H, Hou Y, Zhuang Y, and Miranda L. 2009. High-Level Diversity of Dinoflagellates in the Natural Environment, Revealed by assessment of Mitochondrial cox1 and cob Genes for Dinoflagellate DNA Barcoding. Appl. Environ. Microbiol. 75, 1279–1290.
  44. Lin S, Sandh G, Zhang H, Cheng J, Perkins K, Carpenter EJ and Bergman B. 2009. Two flavodoxin genes in Trichodesmium (Oscillatoriales, Cyanophyceae): Remarkable sequence divergence and possible functional diversification. J. Exp. Mar. Biol. Ecol. 371, 93–101.
  45. Morita M, Futami K, Zhang H , Kubokawa K, Ojima Y, Okamoto N. 2009. Evolutionary analysis of amphioxus myc gene. Journal of the Tokyo University of Marine Science and Technology 5:11-16.
  46. Stefaniak L, Lambert G, Gittenberger A, Zhang H, Lin S and Whitlatch RB. 2009. Genetic conspecificity of the worldwide populations of Didemnum vexillum Kott, 2002. Aquatic Invasions 4, 29-44. DOI 10.3391/ai.2009.4.1.3.
  47. Zhang H and Lin S. 2009. Retrieval of Missing Spliced Leader in Dinoflagellates. PLoS ONE 4(1): e4129. doi:10.1371/journal.pone.0004129.
  48. Yokoyama S, Takashi Tada, Zhang H, and Lyle Britt. 2008. Elucidation of phenotypic adaptations: molecular analyses of dim-light vision proteins in vertebrates PNAS USA, 105, 13480-13485.
  49. Zhang H, Litaker W, Vandersea MW, Tester P and Lin S. 2008. Geographic distribution of Karlodinium veneficum in the US east coast as detected by ITS- ferredoxin Real-Time PCR assay. J. Plankton Res. 30:905-922.
  50. Zhang H and Lin S. 2008. Status of mRNA editing and SL RNA trans-splicing groups Oxyrrhis, Noctiluca, Heterocapsa, and Amphidinium as basal lineages of dinoflagellates J. Phycol. 44:703-711.
  51. Okamura A, Zhang H, Mikawa N, Kotake A, Yamada Y, Utoh T, Horie N, Tanaka S, Oka HP and Tsukamoto K. 2008. Decline in non-native freshwater eels in Japan: ecology and future perspectives. Environmental Biology of Fishes 81:347-358.
  52. Zhang H, Bhattacharya D, Maranda L and Lin S. 2008. Mitochondrial cob and cox1 and their mRNA editing in Dinophysis acuminata from Narragansett Bay, with special reference to the phylogenetic position of Dinophysis. Appl. Environ. Microbiol. 74:1546–1554.
  53. Lin S, Zhang H and Gray MW. 2008. RNA editing in dinoflagellates and its implications for the evolutionary history of the editing machinery. In: Harold C. Smith (ed.), RNA and DNA Editing: Molecular Mechanisms and Their Integration into Biological Systems. John Wiley & Sons, Inc., Hoboken, New Jersey 280-309.
  54. Zhang H, Bhattacharya D and Lin S. 2007. A three-gene dinoflagellate phylogeny suggests reconciliation of Exuviaella with Prorocentrum and a basal position for Amphidinium and Heterocapsa. J. Mol. Evol. 65:463–474.
  55. Zhang H, Hou Y, Miranda L, Campbell DA, Sturm NR, Gaasterland T, and Lin S. 2007. Spliced leader RNA trans-splicing in dinoflagellates. PNAS USA, 104:4618-4623.(cited 121 times)
  56. Lin S, Zhang H, Hou Y, Miranda L, and Debashish B. 2006. Development of a dinoflagellate-oriented PCR primer set leads to the detection of picoplanktonic dinoflagellates from long island sound. Appl. Environ. Microbiol. 78, 5626–5630.
  57. Lin S, Zhang H and Dubois A. 2006. Low abundance distribution of Pfiesteria piscicida in Pacific and Western Atlantic as detected by mtDNA-18S rDNA real-time polymerase chain reaction. J. Plankton Res., 28, 667-681.
  58. Lin S, Zhang H and Jiao N. 2006. Potential utility of mitochondrial cytochrome b and its mRNA editing in resolving closely related dinoflagellates: a case study of Prorocentrum (Dinophyceae). J. Phycol. 42, 646–654 (2006)
  59. Zhang H, Hou Y and Lin S. 2006. Isolation and characterization of PCNA from the dinoflagellate Pfiesteria piscicida. J. Eukaryot. Microbiol.,53, 142–150.
  60. Zhang H and Lin S. 2005. Mitochondrial Cytochrome b mRNA Editing in Dinoflagellates: Possible Ecological and Evolutionary Associations? J. Eukaryot. Microbiol., 52, 538–545.
  61. Zhang H and Lin S. 2005. Development of a cob-18S dual-gene Real-Time PCR assay for Pfiesteria shumwayae and quantification of this species in the natural environment. Appl. Environ. Microbiol.71, 7053-7063.
  62. George Mcmanus, Huan Zhang, Senjie Lin, Laura Katz Katarzyna A. Pirog. 2005. In the land of the blind, one-eye is king: ecology of the mixotrophic ciliates Strombidium oculatum and Strombidium stylifer. Environ. Microbiol.71, 7053-7063. DOI: 10.1111/j.1550-7408.2005.05202003_1_59.x (6 Apr 2005 published on line)
  63. Lin S and Zhang H. 2005. Isolation of mitochondrial cytochrome b gene and development of a real-time quantitative PCR assay for detecting Neoparamoeba aestuarina. J. Shellfish Research 24, 733-739.
  64. Zhang H, Bhattacharya D and Lin S. 2005. Phylogeny of Dinoflagellates Based on Mitochondrial Cytochrome b and Nuclear Small Subunit rDNA Sequence Comparisons. J. Phycol. 41, 411–420.
  65. Lin S, Mulholland MR, Zhang H, Feinstein TN, Jochem FJ and Carpenter EJ. 2004. Intense grazing and prey-dependent growth of Pfiesteria piscicida (Dinophyceae). J. Phycol. 40: 1062-1073.
  66. Okamura A, Zhang H, Utoh T, et al. 2004. Artificial hybrid between Anguilla anguilla and A. japonica. J. Fish Biology 64, 1450-1454.
  67. McManus G, Zhang H and Lin S. 2004. Marine planktonic ciliates that prey on macroalgae and enslave their chloroplasts. Limnology & Oceanography 49, 308-313.
  68. Zhang H, Futami K and Okamoto N. 2003 Isolation of c-myc genes from goldfish and their tissue specific expression. J. Fish Biol. 63, 1266-1273.
  69. Zhang H and Lin S. 2003. Complex gene structure of the form II Rubisco in the dinoflagellate Prorocentrum minimum (dinophyceae). J. Phycology 39, 1160-1171.
  70. Lin S, Timothy N. Feinstein, Zhang H and Edward J. Carpenter. 2003. Development of an immunofluorescence technique for detecting Pfiesteria piscicida. Harmful Algae 2, 223-231.
  71. Lin S and Zhang H. 2003. Mitogen-associated protein kinase (MAPK) in Pfiesteria piscicida. Appl. Environ. Microbiol. 69, 343-349.
  72. Zhang H, Futami K, Yamada Y, Horie N, Okamura A, Utoh T, Mikawa N, Oka HP Okamoto N. 2002. Isolation of freshwater and deep-sea type opsin genes from the common Japanese conger J. Fish Biol. 61, 313-324.
  73. Tanaka S, Zhang H, Yamada Y, Okamura A, Horie N, Utoh T, Mikawa N, Oka HP and H. Kurokura. 2002. Inhibitory effect of sodium bicarbonate on the motility of sperm of Japanese eel. J. Fish Biol. 60, 1134-1141.
  74. Lin S, Zhang H, Spencer DF, Norman JE and Gray MW. 2002. Widespread and extensive editing of mitochondrial mRNAs in dinoflagellates. J. Mol. Biol. 320, 727-739.
  75. Tanaka S, Zhang H, Horie N, Yamada Y, Okamura A, Utoh T, Mikawa N, Oka HP and Kurokura H. 2002. Long –term cryopreservation of sperm of Japanese eel. J. Fish Biol. 60,139-146.
  76. Zhang H and Lin S. 2002. Detection and Quantification of Pfiesteria piscicida by Using the Mitochondrial Cytochrome b Gene. Appl. Environ. Microbiol. 68, 989-994.
  77. Horie N, Utoh T, Yamada Y, Okamura A, Zhang H, Mikawa N, Atsushi Akazawa, Tanaka S and Oka HP. 2002. Development of embryos and larvae in the common Japanese conger Conger myriaster. Fisheries Scis. 68, 972-983.
  78. Futami K, Komiya T, Zhang H and Okamoto N. 2001. Differential expression of mas and two types of c-myc genes in a tetraploid fish, the common carp (Cyprinus carpio). Gene 269, 113-119.
  79. Yamada Y, Zhang H, Okamura A, Tanaka S, Horie N, Mikawa N, Utoh T and Oka HP. 2001. Morphological and histological changes in the swim bladder during maturation of the Japanese eel. J. Fish Biology 58,804-814.
  80. Okamura A, Zhang H, Yamada Y, Tanaka S, Horie N, Mikawa N, Utoh T and Oka HP. 2000. Re-examination of the spermatozoal ultrastructure of eels: observations of the external morphology of spermatozoa in three species. J. of Fish Biology 57, 161-169.
  81. Zhang H, Futami K, Horie N, Okamura A, Utoh T, Mikawa N, Yamada Y, Tanaka S and Okamoto N. 2000. Molecular cloning of fresh water and deep-sea rod opsin genes from Japanese eel Anguilla japonica and their expressional analysis during sexual maturation. FEBS letters 469, 39-43.
  82. Futami K, Takeru Komiya, Zhang H and Okamoto N. 2000. Determination of heterogeneous transcription start points of two c-myc genes from the common carp (Cyprinus carpio). Gene 245, 43-47.
  83. Okamura A, Utoh T, Zhang H, Yamada Y, Horie N, Mikawa N, Tanaka S and Oka HP. 2000. Seasonal changes in maturity in the Conger Eel Conger myriaster at the Pacific coast of Atsumi Peninsula, central Japan. Nippon Suisan Gakkashi 66, 412-416 (2000) (in Japanese).
  84. Futami K, Zhang H and Okamoto N. 2000. Molecular evolution of two c-myc genes of a tetraploid teleosts, the common carp. Protein, Nucleic acid, Enzume 45, 2943-2948 (2000) (in Japanese).
  85. Zhang H, Mikawa N, Yamada Y, Horie N, Okamura A, Utoh T, Tanaka S and Takashi Motonobu. 1999. Detection of foreign eels in the natural waters of Japan by PCR. Fisheries Science 65, 684-686.
  86. Yokoyama S, Zhang H, Radlwimmer FB and Blow NS. 1999. Adaptive evolution of color vision of the Comoran coelacanth (Latimeria chalumnae). PNAS USA 96, 6279-6284. (111 times cited by the related articles)
  87. Zhang H and Motonobu T. 1999. Spawning migration and Visual adaptation of Japanese eel. Kaiyo Monthly special 18, 141-145 (in Japanese).
  88. Yokoyama S and Zhang H: Cloning and characterization of the pineal gland-specific opsin gene of marine lamprey (Petromyzon marinus). Gene 202, 89-93 (1997).
  89. Zhang H and Yokoyama S. 1997. Molecular evolution of the rhodopsin gene of marine lamprey, Petromyzon marinus. Gene 191, 1-6.
  90. Zhang H, Okamoto N and Ikeda Y. 1995. Two c-myc genes from a tetraploid fish, the common carp (Cyprinus carpio). Gene 153, 231-236.
  91. Zhang H, Hang Zhang, Okamoto N and Ikeda Y. 1994. Cloning and sequencing of a c-myc gene from goldfish Carassius auratus. Fisheries Science 60, 707-711.
  92. Zhang H, Okamoto N and Ikeda Y. 1993. Molecular cloning of carp cellular myc (c-myc) cDNA. Gyobyo Kenkyu 28, 111-117.