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Smithsonian at the Poles: Contributions to International Polar

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Environmental and Molecular Mechanisms<br />

of Cold Adapt<strong>at</strong>ion in <strong>Polar</strong> Marine<br />

Invertebr<strong>at</strong>es<br />

Adam G. Marsh<br />

Adam G. Marsh, Professor, College of Marine<br />

and Earth Studies, University of Delaware, 700<br />

Pilot<strong>to</strong>wn Road, Smith Labor<strong>at</strong>ory 104, Lewes,<br />

DE 19958, USA (amarsh@udel.edu). Accepted<br />

28 May 2008.<br />

ABSTRACT. The under-ice environment places extreme selective pressures on polar marine<br />

invertebr<strong>at</strong>es (sea urchins, starfi sh, clams, worms) in terms of <strong>the</strong> low temper<strong>at</strong>ure,<br />

oligotrophic w<strong>at</strong>ers, and limited light availability. Free-swimming embryos and larvae<br />

face inordin<strong>at</strong>e challenges of survival with almost nonexistent food supplies establishing<br />

near starv<strong>at</strong>ion conditions <strong>at</strong> <strong>the</strong> <strong>the</strong>rmal limits of cellular stress th<strong>at</strong> would appear <strong>to</strong><br />

require large energy reserves <strong>to</strong> overcome. Yet, despite <strong>the</strong> long developmental periods<br />

for which <strong>the</strong>se embryos and larvae are adrift in <strong>the</strong> w<strong>at</strong>er column, <strong>the</strong> coastal under-ice<br />

habit<strong>at</strong>s of <strong>the</strong> polar regions support a surprising degree of vibrant marine life. How can<br />

so many animals be adapted <strong>to</strong> live in such an extreme environment? We all recognize<br />

th<strong>at</strong> environmental adapt<strong>at</strong>ions are coded in <strong>the</strong> DNA sequences th<strong>at</strong> comprise a species<br />

genome. The fi eld of polar molecular ecology <strong>at</strong>tempts <strong>to</strong> unravel <strong>the</strong> specifi c imprint<br />

th<strong>at</strong> adapt<strong>at</strong>ions <strong>to</strong> life in a polar habit<strong>at</strong> have left in <strong>the</strong> genes and genomes of <strong>the</strong>se<br />

animals. This work requires a unique integr<strong>at</strong>ion of both fi eld studies (under ice scuba<br />

diving and experiments) and labor<strong>at</strong>ory work (genome sequencing and gene expression<br />

studies). Understanding <strong>the</strong> molecular mechanisms of cold adapt<strong>at</strong>ion is critical <strong>to</strong> our<br />

understanding of how <strong>the</strong>se organisms will respond <strong>to</strong> potential future changes in <strong>the</strong>ir<br />

polar environments associ<strong>at</strong>ed with global clim<strong>at</strong>e warming.<br />

INTRODUCTION: THE NECESSITY OF SCIENCE DIVING<br />

Looking across <strong>the</strong> coastal margins of most polar habit<strong>at</strong>s, one is immedi<strong>at</strong>ely<br />

struck by <strong>the</strong> stark, frozen wasteland th<strong>at</strong> hides <strong>the</strong> transition from land <strong>to</strong><br />

sea bene<strong>at</strong>h a thick layer of snow and ice. Standing on <strong>the</strong> sea ice surface along<br />

any shore line above 70° l<strong>at</strong>itude, it is hard <strong>to</strong> imagine th<strong>at</strong> <strong>the</strong>re is fl uid ocean<br />

anywhere nearby, and even harder <strong>to</strong> think th<strong>at</strong> <strong>the</strong>re is even a remote possibility<br />

of animal life in such an environment. Yet under <strong>the</strong> fi ve meters of solid sea ice, a<br />

rich and active community of marine organisms exists. The real challenge is getting<br />

<strong>to</strong> <strong>the</strong>m.<br />

Scientists studying how <strong>the</strong>se organisms are adapted <strong>to</strong> survive and persist<br />

near <strong>the</strong> poles are limited by <strong>the</strong> logistical constraints in getting access under <strong>the</strong><br />

ice <strong>to</strong> collect animals and plants for study. The sea ice cover establishes an effective<br />

barrier <strong>to</strong> using most of <strong>the</strong> common collection techniques th<strong>at</strong> marine biologists<br />

employ from vessel-based sampling oper<strong>at</strong>ions. The time and effort th<strong>at</strong> is<br />

invested in opening a hole in <strong>the</strong> ice gre<strong>at</strong>ly precludes <strong>the</strong> number of sampling sites

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