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in Flow |
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in Flow |
Instructors: Lauren Mullineaux (Biology) and Stace Beaulieu (AOPE)
| Organisms living on the seafloor interact with flow. This dynamic environment
presents them with challenges and opportunities in terms of physical integrity,
feeding, larval settlement and post-settlement transport.
Effects of flow on benthic organisms have been studied predominantly from one of two viewpoints, either oceanographic (i.e. Jumars, Eckman, Butman, etc.) or biomechanical (i.e. Vogel, Denny, Koehl, etc.). This course will explore both approaches, bringing in discussion of physical processes, including Reynold’s number, boundary-layer fluid dynamics, drag and particle transport. Classes will be conducted in lecture and discussion format with directed readings, student presentations, problem-solving exercises using Matlab, and a short paper. |
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| Here is our class on 29 Feb 2000 running the flume experiment at CRL (click on thumbnails). |  |
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Syllabus
Class will meet Tuesdays 13:00 - 14:30 in Redfield 204.
Readings will be available in Redfield 206 for photocopying.
| Week # | Date | Topic | Reading | Assignment | Instructor(s)
(and Student) |
| 1 | 1 Feb | Introduction and description of the benthic boundary layer | Nowell, A.R.M. and Jumars, P.A., 1984. Flow environments of aquatic benthos. Annual Review of Ecology and Systematics, 15: 303-328. | Beaulieu and Mullineaux | |
| 2 | 8 Feb | Small-scale recruitment of benthic invertebrates: larvae as particles | Butman, C., 1987. Larval settlement of soft-sediment
invertebrates: the spatial scales of pattern explained by active habitat
selection and the emerging role of hydrodynamics. Oceanogr. Mar. Biol.
Annu. Rev. 25: 113-165.
Gross, T., F. Werner and J. Eckman, 1992. Numerical modeling of larval settlement in turbulent bottom boundary layers. Journal of Marine Research 50: 611-642. Mullineaux, L.S. and E. Garland, 1993. Larval recruitment in response to manipulated field flows. Marine Biology116: 667-683. |
Choose class period for student presentation | Mullineaux |
| 3 | 15 Feb | Small scale recruitment: behavior and chemical cues | Pawlik, J. and C. Butman, 1993. Settlement of
a marine tube worm as a function of current velocity: Interacting effects
of hydrodynamics and behavior. Limnol. Oceanogr. 38: 1730-1740.
Snelgrove, P., J. Grassle and C. Butman, 1998. Sediment choice by settling larvae of the bivalve, Spisula solidissima (Dillwyn), in flow and still water. J. Expt. Mar. Biol. Ecol. 231: 171-190. Tamburri, M., C. Finelli, D. Wethey and R. Zimmer-Faust, 1996. Chemical induction of larval settlement behavior in flow. Biological Bulletin 191: 367-373. |
Mullineaux | |
| 4 | 29 Feb | Flume demonstration at the RCRC | Muschenheim, D.K., Grant, J. and Mills, E.L., 1986.
Flumes
for benthic ecologists: theory, construction, and practice.
Marine Ecology
Progress Series, 28: 185-196.
Nowell, A.R.M. and Jumars, P.A., 1987. Flumes: theoretical and experimental considerations for simulation of benthic environments. Oceanogr. Mar. Biol. Annu. Rev., 25: 91-112. |
Matlab problem set handed out | Mullineaux and Beaulieu |
| 5 | 7 Mar | Suspension feeding at the scale of an individual organism | Johnson, A.S., 1988. Hydrodynamic study of the
functional morphology of the benthic suspension feeder Phoronopsis viridis
(Phoronida). Marine Biology, 100: 117-126.
Rubenstein, D.I. and Koehl, M.A.R., 1977. The mechanisms of filter feeding: some theoretical considerations. American Naturalist, 111(981): 981-994. Also, take a look at: Patterson, M.R., 1992. A chemical engineering view of cnidarian symbioses. American Zoologist, 32(4): 566-582. |
Problem set due | Beaulieu (and Sarah Rothberg) |
| 6 | 14 Mar | Suspension feeding at the scale of a colony or group of organisms | McFadden, C.S., 1986. Colony fission increases
particle capture rates of a soft coral: advantages of being a small colony.
Journal
of Experimental Marine Biology and Ecology, 103: 1-20.
Kaandorp, J.A. 1999. Morphological analysis of growth forms of branching marine sessile organisms along environmental gradients. Marine Biology 134: 295-306. Also, take a look at: Helmuth, B.S.T., Sebens, K.P. and Daniel, T.L., 1997. Morphological variation in coral aggregations: branch spacing and mass flux to coral tissues. Journal of Experimental Marine Biology and Ecology, 209(1-2): 233-259. |
Beaulieu (and Rob Jennings) | |
| 7 | 28 Mar | Effects of benthic organisms on particle transport in the benthic boundary layer | Muschenheim, D.K., 1987. The dynamics of near-bed
seston flux and suspension-feeding benthos. J. Mar. Res., 45: 473-496.
O'Riordan, C.A., Monismith, S.G. and Koseff, J.R., 1993. A study of concentration boundary-layer formation over a bed of model bivalves. Limnol. Oceanogr., 38(8): 1712-1729. Also, take a look at: Frechette, M., Butman, C.A. and Geyer, W.R., 1989. The importance of boundary-layer flows in supplying phytoplankton to the benthic suspension feeder, Mytilus edulis. Limnol. Oceanogr., 34(1): 19-36. |
Decide topic for short paper (SEE BELOW) | Beaulieu (and Mario Sengco) |
| 8 | 4 Apr | Hydrodynamic forces imposed on benthic organisms | Witman, J.D. and Suchanek, T.H., 1984. Mussels
in flow: drag and dislodgement by epizoans. Mar. Ecol. Prog. Ser.,
16: 259-268.
Denny, M., Gaylord, B., Helmuth, B. and Daniel, T., 1998. The menace of momentum: dynamic forces on flexible organisms. Limnol. Oceanogr., 43(5): 955-968. Also, take a look at: Denny, M.W., Daniel, T.L. and Koehl, M.A.R., 1985. Mechanical limits to size in wave-swept organisms. Ecological Monographs, 55(1): 69-102. |
Beaulieu (and Lara Gulmann) | |
| 9 | 11 Apr | Effects of benthic organisms on momentum flux to the bed: consequences for sediment transport | Carey, D.A., 1983. Particle resuspension in the
benthic boundary layer induced by flow around polychaete tubes. Can.
J. Fish. Aquat. Sci., 40(Suppl. 1): 301-308.
Wright, L.D., Schaffner, L.C. and Maa, J.P.-Y., 1997. Biological mediation of bottom boundary layer processes and sediment suspension in the lower Chesapeake Bay. Mar. Geol., 141: 27-50. Also, take a look at: Nepf, H.M. and Koch, E.W., 1999. Vertical secondary flows in submersed plant-like arrays. Limnol. Oceanogr., 44(4): 1072-1080. |
Short paper due | Beaulieu (and Masami Fujiwara) |
| 10 | 25 Apr | Students review fellow students' papers | Fellow students' papers | Written reviews due | Mullineaux and Beaulieu |
| 11 | 2 May | Large scale recruitment: topographic effects | Epifanio, C., 1988. Dispersal strategies of two
species of swimming crab on the continental shelf adjacent to Delaware
Bay. Marine Ecology Progress Series 49: 243-248.
Mullineaux, L.S. and S.W. Mills, 1997. A test of the larval retention hypothesis in seamount-generated flows. Deep-Sea Res. 44: 745-770. Pineda, J., 1991. Predictable upwelling and the shoreward transport of planktonic larvae by internal tidal bores. Science 253: 548-551. |
Revised paper due
and Matlab problem set handed out |
Mullineaux (and Heidi Fuchs) |
| 12 | 9 May | Post-settlement transport of benthic fauna | Emerson, C.W. and J. Grant, 1991. The control
of soft-shell clam (Mya arenaria) recruitment on intertidal sandflats
by bedload sediment transport. Limnol. Oceanogr. 36: 1288-1300.
Palmer, M.A., 1988. Dispersal of marine meiofauna: a review and conceptual model explaining passive transport and active emergence with implications for recruitment. Marine Ecology Progress Series 48: 81-91. Sigurdsson, J.B., C.W. Titman and P.A. Davies, 1976. The dispersal of young post-larval bivalve molluscs by byssus threads. Nature 262: 386-387. |
Problem set due | Mullineaux |
Books referred to during the course:
Denny, M.W., 1988. Biology and the mechanisms of the wave-swept environment.
Princeton University Press, Princeton, 329 pp.
Denny, M.W., 1993. Air and water: the biology and physics of life's
media. Princeton University Press, Princeton, 341 pp.
Mann, K.H. and Lazier, J.R.N., 1991. Dynamics of marine ecosystems:
biological-physical interactions in the oceans. Blackwell Scientific Publications,
Boston, 466 pp.
Schlichting, H., 1979. Boundary-layer theory, seventh edition. McGraw-Hill
Book Company, New York, 817 pp.
Tennekes, H. and Lumley, J.L., 1972. A first course in turbulence.
MIT Press, Cambridge, 300 pp.
Van Dyke, M., 1982. An album of fluid motion. Parabolic Press, Stanford,
176 pp.
Vogel, S., 1994. Life in moving fluids: the physical biology of flow.
Princeton University Press, Princeton, 467 pp.
Wildish, D. and Kristmanson, D., 1997. Benthic suspension feeders
and flow. Cambridge University Press, Cambridge, 409 pp.
Grades will be based on:
2 problem sets = 1/3; paper (plus review) = 1/3; oral presentation
= 1/6; participation = 1/6.
Term Paper Assignment: Choose a science topic that is: (1) relevant to the course, (2) recently or currently controversial. Describe the two (or more) sides of the controversy, and explain why the topic, and its resolution, is important. Critically evaluate the evidence for each point of view. If you think the evidence for one side is better, say which and explain why. If you think the question is unresolved or unresolvable, also explain why. Keep the length to about 5 pages single spaced.
Paper Review: All students will read all 6 term papers. Each student will be assigned one paper to review formally. For the formal review, write a short (~1-page) evaluation based on the following criteria (note that we are using a fairly loose definition of ‘controversy’; any paper that investigates alternative explanations of an unresolved issue qualifies): (1) clarity of presentation of the topic, (2) documentation of the controversy from the literature (this does not need to involve hoards of references; for some topics the issues can be well represented with just a few), (3) balance of presentation, (4) explanation of the scientific (basic or applied) importance of the topic, (5) critical analysis of the controversy (this is the most important part).
Panel Discussion: Each student will be responsible for leading a short (<15 min) discussion of the paper he/she reviewed formally. This discussion should include a brief summary and critique of the paper. Fellow panelists (including the paper’s author) are free to concur, disagree, or add comments. The intent of this panel discussion is to offer constructive criticism and positive reinforcement. The author will use the written critiques (including one from the instructor) and the panel discussion as the basis for revising the paper.