Susan Brawley, Ph.D.

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321 Hitchner Hall
Orono, ME  04469-5741


Ph.D. University of California

Research interests



  • Brawley, S. H., Nicolas A. Blouin, Elizabeth Ficko-Blean, Glen L. Wheeler, Martin Lohr, Holly Goodson, Jerry W. Jenkins, Crysten E. Blaby-Haas, Katherine E. Helliwell, Cheong Xin Chan, Tara Marriage, Debashish Bhattacharya, Anita S. Klein, Yacine Badis, Juliet Brodie, Yuanyu Cao, Jonas Collén, Simon M. Dittami, Claire M. M. Gachon, Beverley R. Green, Steven J. Karpowicz, Jay W. Kim, Ulrich Johan Kudahl, Senjie Lin, Gurvan Michel, Maria Mittag, Bradley J. S. C. Olson, Jasmyn Pangilinan, Yi Peng, Huan Qiu, Shengqiang Shu, John T. Singer, Alison G. Smith, Brittany N. Sprecher, Volker Wagner, Wenfei Wang, Zhi-Yong Wang, Juying Yan, Charles Yarish, Simone Zäuner-Riek, Yunyun Zhuang, Yong Zou, Erika A. Lindquist, Jane Grimwood, Kerrie Barry, Daniel S. Rokhsar, Jeremy Schmutz, John W. Stiller, Arthur R. Grossman, Simon E. Prochnik. 2017. Insights into the red algae and eukaryotic evolution from the genome of Porphyra umbilicalis (Bangiophyceae, Rhodophyta). Proc. Natl. Acad. Sci. U.S. (Includes 106 page SI Appendix).
  • Wells, M., Potin, P., Craigie, J., Raven, J., Merchant, S., Helliwell, K., Smith, A., Camire, M. & S. H. Brawley. 2017. Algae as nutritional and functional food sources: Revisiting our understanding. J. Appl.Phycol. 29: 949-982. doi:10.1007/s10811-016-0974-5.
  • Kim, J., Brawley, S. H., Prochnik, S., Schmutz, J., Jenkins, J., Stiller, J., & A. R. Grossman. 2016.Genome analysis of Planctomycetes inhabiting blades of the red alga Porphyra umbilicalis. PLoS ONE 11(3): e0151883.
  • Miranda, L., Hutchison, K., Grossman, A. & S. H. Brawley. 2013. Diversity and abundance of    the bacterial community of the red macroalga Porphyra umbilicalis: Did bacterial farmers produce macroalgae? PLoS ONE 8(3) e58269. doi:10.1371/journal.pone.0058269
  • Blouin, N. A. & S. H. Brawley. 2012. An AFLP analysis of clonality in widespread asexual populations of Porphyra umbilicalis (Rhodophyta) with a sensitivity analysis for bacterial contamination. Marine Biology 159, 2723-2729.
  • Stiller, J. W., Perry, J., Rymarquis, L. A., Green, P. J., Prochnik, S., Lindquist, E., Chan, C. X., Yarish, C., Lin, S., Zhuang, Y., Blouin, N. A. & S. H. Brawley. 2012. Major developmental regulators and their expression in two closely related species of Porphyra (Rhodophyta). J. Phycol. 48, 883-896.
  • Chan, C. X., Berg, M., Zäuner, S., Benning, C., Wheeler, G., Levine, I., Grossman, A. R., Blouin, N. A., Prochnik, S. E., Eriksen, R. L., Klein, A. S., Yarish, C., Brawley, S. H. &   D. Bhattacharya. 2012. Analysis of Porphyra membrane transporters demonstrates gene transfer among photosynthetic eukaryotes and numerous sodium-coupled transport systems. Plant Physiology (DOI:10.1104/pp.112.193896).
  • Chan, C. X., N. Blouin, Y.Y. Zhuang, S. Zauner, S. Prochnik, E. Lindquist, S. Lin, C. Benning, M. Lohr, C. Yarish, E. Gantt, A. Grossman, S. Lu, K. Muller, J. Stiller, S. Brawley, D. Bhattacharya. 2012. Porphyra (Bangiophyceae) transcriptomes provide insights into red algal development and metabolism. J. Phycol. 48, 1328-1342. 7
  • Muhlin, J. F., Coleman, M. A., Rees, T. A. V. & S. H. Brawley. 2011. Modeling of reproduction             in the intertidal macrophyte Fucus vesiculosus L. and implications for spatial subsidies in         the nearshore environment. Mar. Ecol. Prog. Ser. 440, 79-94 (DOI: 10.3354/meps09350).
  • Johnson, L. A., Brawley, S. H. & W. H. Adey. 2011. Factors explaining the continued spread of  Fucus serratus in the Canadian Maritimes. Biological Invasions DOI: 10.1007/s10530-01109976-z.
  • Blouin, N., Brodie, J., Pu, X., Grossman, A. & S. H. Brawley. 2011. Porphyra: A marine crop shaped by stress. Trends in Plant Science 16, 29-37.
  • CABI. 2010. Littorina littorea [original text by S. Brawley]. In: Invasive Species Compendium. Wallingford, UK: CABI. [26 published pages].
  • Gantt, E. and 18 intermediate authors & S. H. Brawley. 2010. Porphyra: Complex life histories in a harsh environment. P. umbilicalis, the genomics’ project. In: J. Seckbach & D. Chapman (eds), Red Algae in the Genomic Age (v. 13, Cellular Origins, Life in Extreme Habitats and Astrobiology), Springer, pp 129-148.
  • Muhlin, J. & S. H. Brawley. 2009. Recent versus relic: Discerning the genetic signature of Fucus vesiculosus (Heterokontophyta; Phaeophyceae) in the northwestern Atlantic. J. Phycol. 45, 828-837.
  • Brawley, S. H., J. A. Coyer, A. M. H. Blakeslee, G. Hoarau, L. E. Johnson, J. E. Byer, W. T. Stam & J. L. Olsen. 2009. Historical invasions of the intertidal zone of Atlantic North America associated with distinctive patterns of trade and emigration. Proc. Natl. Acad. Sci., U.S.A. 106, 8239-8244 (doi: 10.1073/pnas.0812300106).
  • Brawley, S. H., J. Pusey, B. J. W. Cole, L. E. Gott and S. A. Norton. 2008. A revolutionary model to improve science education, teachers, and scientists. Maine Policy Review 17, 68-81.
  • Schmidt, P. S., E. A. Serrão, G. A. Pearson, C. Riginos, P. D. Rawson, T. J. Hilbish, S. H. Brawley, G. C. Trussell, E. Carrington, D. S. Wethey, J. W. Grahame, F. Bonhomme, and D. M. Rand. 2008. Ecological genetics in the North Atlantic intertidal: Environmental gradients, replicated clines, and adaptation at specific loci. Ecology 89, S91-S107.
  • Muhlin, J. F., C. R. Engel, R. Stessel, R. A. Weatherbee & S. H. Brawley. 2008. The influence of coastal topography, circulation patterns and rafting in structuring populations of an intertidal alga. Molec. Ecol. 17, 1198-1210.
  • Koester, J., S. H. Brawley, L. Karp-Boss and D. G. Mann. 2007. Sexual reproduction in the marine centric diatom Ditylum brightwellii (Bacillariophyta). Eur. J. Phycol. 42, 351-366.
  • Blouin, N. B., X. G. Fei, J. Peng, C. Yarish, and S. H. Brawley. 2007. Seeding nets with neutral spores of the red alga Porphyra umbilicalis (L.) Kützing for use in integrated multi-trophic aquaculture (IMTA). Aquaculture 270, 77-91.
  • Blouin, N., B. L. Calder, B. Perkins, and S. H. Brawley. 2006. Sensory and fatty acid analyses of
    two Atlantic species of Porphyra (Rhodophyta). J. Appl. Phycol. 18, 79-85.
  • Coleman, M. A. and S.H. Brawley. 2005. Variability in temperature and historical patterns in
    reproduction in the Fucus distichus complex (Heterokontophyta; Phaeophyceae): Implications for speciation and the collection of herbarium specimens. J. Phycol. 41, 1110-1119.
  • Coleman, M. and S. H. Brawley. 2005. Spatial and temporal variability in dispersal and
    population genetic structure of a rockpool alga. Mar. Ecol. Prog. Ser. 300, 63-77.
  • Coleman, M. A. and S. H. Brawley. 2005. Are life history characteristics good predictors of genetic diversity and structure? A case study of the interetidal alga Fucus spiralis (Heterokontophyta: Phaeophyceae). J. Phycol. 41, 753-762.
  • Gordon, R. and S. H. Brawley. 2004. Effects of water motion on propagule release from algae with complex life histories. Mar. Biol. 145, 21-29.
  • Calder, E., M. Bagley, and S. H. Brawley. 2004. National Science Foundation Graduate Teaching Fellows promote food science education in K-12 schools in Maine. J. Food Sci. Ed. 2, 58-60.
  • Li, R. and S. H. Brawley. 2004. Improved survival to heat stress in intertidal embryos simultaneously exposed to hypersalinity and the effect of parental thermal history. Mar. Biol. 144, 205-213.
  • Engel, C., S. H. Brawley, K. J. Edwards and E. Serrão. 2003. Isolation and cross-species
    amplification of microsatellite loci from the fucoid seaweeds Fucus vesiculosus, F. serratus and Ascophyllum nodosum (Heterokontophyta, Fucaceae). Molecular Ecology Notes 3,180-182.
  • Berndt, M. L., J. A. Callow, and S. H. Brawley. 2002. Gamete concentrations and timing and success of fertilization in a rocky shore seaweed. Mar. Ecol. Prog. Ser. 226, 273-285.
  • Speransky, V., S. H. Brawley, and M. E. McCully. 2001. Ion fluxes and modification of the extracellular matrix during gamete release in fucoid algae. J. Phycol. 37, 555-573.
  • Brawley, S.H., and K.R. Robinson. 1985. Cytochalasin treatment disrupts the endogenous currents associated with cell polarization in embryogenesis: studies of the role of F-actin in embryogenesis. J. Cell Biol. 100, 1173-84.
  • Speransky, S. and S. H. Brawley. 2000. Gamete release is increased by calm conditions in the coenocytic green alga Bryopsis (Chlorophyta). J. Phycol. 36, 730-739.
  • Brawley, S.H., L.E. Johnson, G.A. Pearson, V. Speransky, R.L. and E. Serrão 1999. Gamete release at low tide in fucoid algae: Maladaptive or advantageous? Amer. Zool. 39: 218-29.
  • Rosemond, A.D., P.J. Mulholland, and S.H. Brawley. 2000. Seasonally shifting limitation of stream periphyton: Response of algal populations and assemblage biomass and productivity to variation in light, nutrients, and herbivores. Can. J. Fish. Aquat. Sci. 57: 66-75.
  • Serrão, E., L. Alice and S. Brawley. 1999. Evolution of the Fucaceae (Phaeophyceae) inferred from nrDNA-ITS. J. Phycol. 35, 382-394.
  • Johnson, L.E. and S.H. Brawley. 1999. Dispersal and recruitment of a canopy-forming intertidal alga: The relative roles of propagule availability and post-settlement processes. Oecologia 117, 517-26.
  • Serrão, E., S.H. Brawley, J. Hedman, L. Kautsky and G. Samuelsson. 1999. Reproductive success in Fucus vesiculosus (Phaeophyceae) in the Baltic Sea. J. Phycol. 35: 254-269.
  • Brawley, S.H. 1999. Submission and retrieval of an aligned set of nucleic acid sequences.
    J. Phycol. 35, 433-37.
  • Speransky, V. S. Speransky, and S. H. Brawley. 1999. Cryoanalytical studies of freezing damage and recovery in Fucus vesiculosus (Phaeophyceae). J. Phycol. 35, 1264-1275.
  • Pearson, G.A., E. Serrão, and S.H. Brawley. 1998. Control of gamete release in fucoid algae: sensing hydrodynamic conditions via carbon acquisition. Ecology 79, 1725-1739.
  • Li, R., S.H. Brawley and T. Close. 1998. Proteins immunologically related to dehydrins in fucoid algae. J. Phycol. 34, 642-50.
  • Pearson, G. and S.H. Brawley. 1998. A model for signal transduction during gamete release in the fucoid alga Pelvetia compressa. Plant Physiol. 118, 305-13.
  • Rosemond, A. D. and S. H. Brawley. 1996. Species-specific characteristics explain the persistence of Stigeoclonium tenue (Chlorophyta) in a woodland stream. J. Phycol. 32, 54-63.
  • Serrão, E. A., L. Kautsky and S. H. Brawley. 1996. Distributional success of the marine seaweed Fucus vesiculosus the brackish Baltic Sea correlates with the osmotic capabilities of Baltic gametes. Oecologia 107, 1-12.
  • Serrão, E. A., G. Pearson, L. Kautsky and S. H. Brawley. 1996. Successful external fertilization in turbulent environments. Proc. Natl. Acad. Sci. U.S.A. 93, 5286-90.
  • Pearson, G. and S. H. Brawley. 1996. Reproductive ecology of Fucus distichus (Phaeophyceae): An intertidal alga with successful external fertilization. Mar. Ecol. Prog. Ser. 143, 211-23.
  • Committee on Biodiversity in Marine Systems. 1995. Understanding Marine Biodiversity. National Academy Press, Washington, 128 pp.
  • Brawley, S.H. and L.E. Johnson. 1993. Predicting desiccation stress in microscopic organisms: the use of agarose beads to determine evaporation rates within and between intertidal microhabitats. J.Phycol. 29, 528-535.
  • Davison, I.R., L.E. Johnson, and S.H. Brawley. 1993. Sublethal stress in the intertidal zone: tidal emersion inhibits photosynthesis and retards development in embryos of the brown alga Pelvetia fastigiata. Oecologia 96, 483-492.
  • Brawley, S.H. 1992. Mesoherbivores. In: Plant-Animal interactions in the Marine Benthos (D. John, S. Hawkins & J. Price, Ed.), Oxford University Press, Oxford, pp 235-263.
  • Brawley, S.H. 1992. Fertilization in natural populations of the dioecious brown alga Fucus ceranoides L. and the importance of the polyspermy block. Marine Biology. 113, 145-157Â
  • Brawley, S.H. and L.E. Johnson. 1992. Gametogenesis, gametes and zygotes: an ecological perspective on sexual reproduction in the algae. British Phycological Journal 27, 233-252.
  • Brawley, S.H. 1991. The fast block against polyspermy in fucoid algae is an electrical block. Develop. Biol. 144, 94-106.
  • Brawley, S.H. and L.E. Johnson. 1991. Survival of fucoid embryos in the intertidal zone depends upon developmental stage and microhabitat. J. Phycol. 27, 179-186.
  • Brawley, S.H. 1990. Polyspermy blocks in fucoid algae and the occurrence of polyspermy in nature. In: Mechanism of Fertilization: Plants to Humans (B.Dale, Ed.). Berlin: Springer- Verlag, pp. 419-431.
  • Brawley, S.H. 1990. The polyspermy block in fucoid algae. In: Cell Walls and Surfaces, Reproduction, Photosynthesis, Experimental Phycology I (W. Wiessner, D.G. robinson, R.C. Starr, Eds.). Berlin: Springer-Verlag, pp. 134-144.
  • Brawley, S.H. and D.M. Roberts. 1989. Calmodulin-binding proteins are developmentally regulated in gametes and embryos of fucoid algae. Develop. Biol. 131, 313-320.
  • Brawley, S.H. and X.G. Fei. 1988. Ecological studies of G. asiatica and G. lemaneiformis in China. Chinese J. Oceanol. Limnol. 6, 22-34.
  • Brawley, S.H. and E. Bell. 1987. Partial activation of Fucus eggs with calcium ionophores and low- sodium seawater. Develop. Biol. 122, 217-226.
  • Brawley, S.H. and X. G. Fei. 1987. Studies of mesoherbivory in aquaria and in an unbarricaded mariculture farm on the Chinese coast. J. Phycol. 23, 614-623.
  • Brawley, S.H. 1987. A sodium-dependent, fast block to polyspermy occurs in eggs of fucoid algae. Develop. Biol. 124, 390-397. (Featured on journal cover.)
  • Allen, N.S. and S.H. Brawley. 1984. Observations of exocytosis in Fucus vesiculosus gametes using video-enhanced light microscopy: a video report. Cell Motility 4, 25-27.
  • Brawley, S.H., D.F. Wetherell, and K.R. Robinson. 1984. Electrical polarity in embryos of wild carrot precedes cotyledon development. Proc. Natl. Acad. Sci. U.S.A. 81, 6064-6067.
  • Brawley, S.H. and J.R. Sears. 1982. Septal plugs in a green alga. Amer. J. Bot. 69, 455-463.
  • Brawley, S.H. and W. H. Adey. 1982. Coralliophila abbreviata: a significant corallivore! Bull. Mar. Sci. 32, 595-599.
  • Sears, J.R. and S.H. Brawley. 1982. Smithsoniella gen.nov., a possible evolutionary link between the multicellular and siphonous habits in the Ulvophyceae, Chlorophyta. Amer. J. Bot. 69, 1450-1461.
  • Brawley, S.H. and W.H. Adey. 1981. The effect of micrograzers on algal community structure in a coral reef microcosm. Marine Biology 61, 167-177.
  • 1981. Micrograzers may affect macroalgal density. Nature 292, 177.Â
  • Brawley, S.H. and R. Wetherbee. 1981. Algal cytology and ultrastructure. In: Biology of the Seaweeds (Botanical Monograph series) (C. Lobban and M. Wynne, eds.). Oxford: Blackwell Scientific Pub. Ltd., pgs. 248-299.
  • Quatrano, R.S., S.H. Brawley, and W.E. Hogsett. 1979. The control of the polar deposition of a sulfated polysaccharide in Fucus zygotes. In: Determinants of Spatial Organization (S.Subtelny and I. R. Konigsberg, eds.). New York: Academic Press, pgs. 77-96.
  • Brawley, S.H. and R.S. Quatrano. 1979a. Effects of microtubule inhibitors on pronuclear migration and embryogenesis in Fucus distichus (Phaeophyta). J. Phycol. 15, 266-272.Â
  • 1979b. Sulfation of fucoidin in Fucus embryos. IV. Autoradiographic investigations of fucoidin sulfation and secretion during differentiation and the effect of cytochalasin treatment. Develop. Biol. 73, 193-205.
  • Brawley, S.H., R.S. Quantrano, and R. Wetherbee. 1977. Fine-structural studies of the gametes and embryo of Fucus vesiculosus L. (Phaeophyta), III. Cytokinesis and the multicellular embryo. J. Cell Sci. 244, 275-294.
  • Brawley, S. H. and W. H. Adey. 1977. Territorial behavior of three-spot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity. Env. Biol. Fish 2,45-51Â
  • Brawley, S.H., R. Wetherbee, and R. S. Quatrano, 1976a. Fine-structural studies of the gametes and the embryo of Fucus vesiculosus L. (Phaeophyta). I. Fertilization and pronuclear fusion. J. Cell Sci. 20, 233-254.
  • Fine-structural studies of the gametes and embryo of Fucus vesiculosus L. (Phaeophyta). II. The cytoplasm of the egg and young zygote. J. Cell Sci. 20, 255-271.