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<div class=Section1>
<p class=MsoNormal><i><span style='font-family:"Arial","sans-serif"'>Nature </span></i><span
style='font-family:"Arial","sans-serif"'>(2017)<o:p></o:p></span></p>
<p class=MsoNormal><span style='font-family:"Arial","sans-serif"'>doi:10.1038/nature21415<o:p></o:p></span></p>
<p class=MsoNormal><span style='font-family:"Arial","sans-serif"'>Published
online 15 March 2017<o:p></o:p></span></p>
<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:15.0pt;
margin-left:0in'><span style='font-size:8.0pt;font-family:"Arial","sans-serif";
letter-spacing:-.4pt;font-weight:normal'><o:p> </o:p></span></h1>
<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:15.0pt;
margin-left:0in'><span style='font-size:18.0pt;font-family:"Arial","sans-serif";
letter-spacing:-.4pt'>Phytoplankton can actively diversify their migration
strategy in response to turbulent cues<o:p></o:p></span></h1>
<p class=MsoNormal style='margin-left:0in;text-indent:-.25in;mso-list:l2 level1 lfo1'><![if !supportLists]><span
lang=ES style='font-size:10.0pt;font-family:Symbol'><span style='mso-list:Ignore'>·<span
style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><span style='font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html?WT.ec_id=NATURE-20170316&spMailingID=53632654&spUserID=MjA1NTE2ODQxMAS2&spJobID=1122609480&spReportId=MTEyMjYwOTQ4MAS2#auth-1"><span
class=fn><span lang=ES style='color:windowtext;text-decoration:none'>Anupam
Sengupta</span></span></a></span><span class=comma><span lang=ES
style='font-family:"Arial","sans-serif"'>,</span></span><span lang=ES
style='font-family:"Arial","sans-serif"'> </span><span style='font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html?WT.ec_id=NATURE-20170316&spMailingID=53632654&spUserID=MjA1NTE2ODQxMAS2&spJobID=1122609480&spReportId=MTEyMjYwOTQ4MAS2#auth-2"><span
class=fn><span lang=ES style='color:windowtext;text-decoration:none'>Francesco
Carrara</span></span></a></span><span style='font-family:"Arial","sans-serif"'>
<span lang=ES>&<span class=apple-converted-space> </span></span></span><span
style='font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html?WT.ec_id=NATURE-20170316&spMailingID=53632654&spUserID=MjA1NTE2ODQxMAS2&spJobID=1122609480&spReportId=MTEyMjYwOTQ4MAS2#auth-3"><span
class=fn><span lang=ES style='color:windowtext;text-decoration:none'>Roman
Stocker</span></span></a></span><span lang=ES style='font-family:"Arial","sans-serif"'><o:p></o:p></span></p>
<p style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:15.0pt;
margin-left:0in'><span lang=ES style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<p style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:15.0pt;
margin-left:0in'><span style='font-family:"Arial","sans-serif"'>Marine
phytoplankton inhabit a dynamic environment where turbulence, together with
nutrient and light availability, shapes species fitness, succession and
selection</span><sup><span style='font-size:9.0pt;font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref1"
title="Margalef, R. Life-forms of phytoplankton as survival alternatives in an unstable environment. Oceanol. Acta 1, 493–509 (1978)"
id=ref-link-1><span style='color:windowtext;text-decoration:none'>1</span></a>,<span
class=apple-converted-space> </span><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref2"
title="Smayda, T. J. & Reynolds, C. S. Community assembly in marine phytoplankton: application of recent models to harmful dinoflagellate blooms. J. Plankton Res. 23, 447–461 (2001)"
id=ref-link-2><span style='color:windowtext;text-decoration:none'>2</span></a></span></sup><span
style='font-family:"Arial","sans-serif"'>. Many species of phytoplankton are
motile and undertake diel vertical migrations to gain access to nutrient-rich
deeper layers at night and well-lit surface waters during the day</span><sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref3"
title="Bollens, S. M., Rollwagen-Bollens, G., Quenette, J. A. & Bochdansky, A. B. Cascading migrations and implications for vertical fluxes in pelagic ecosystems. J. Plankton Res. 33, 349–355 (2011)"
id=ref-link-3><span style='color:windowtext;text-decoration:none'>3</span></a>,<span
class=apple-converted-space> </span><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref4"
title="Schuech, R. & Menden-Deuer, S. Going ballistic in the plankton: anisotropic swimming behavior of marine protists. Limnol. Oceanogr. Fluids Environ. 4, 1–16 (2014)"
id=ref-link-4><span style='color:windowtext;text-decoration:none'>4</span></a></span></sup><span
style='font-family:"Arial","sans-serif"'>. Disruption of this migratory
strategy by turbulence is considered to be an important cause of the succession
between motile and non-motile species when conditions turn turbulent</span><sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref1"
title="Margalef, R. Life-forms of phytoplankton as survival alternatives in an unstable environment. Oceanol. Acta 1, 493–509 (1978)"
id=ref-link-5><span style='color:windowtext;text-decoration:none'>1</span></a>,<span
class=apple-converted-space> </span><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref5"
title="Sullivan, J. M., Swift, E., Donaghay, P. L. & Rines, J. E. B. Small-scale turbulence affects the division rate and morphology of two red-tide dinoflagellates. Harmful Algae 2, 183–199 (2003)"
id=ref-link-6><span style='color:windowtext;text-decoration:none'>5</span></a>,<span
class=apple-converted-space> </span><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref6"
title="Berdalet, E. et al. Species-specific physiological response of dinoflagellates to quantified small-scale turbulence. J. Phycol. 43, 965–977 (2007)"
id=ref-link-7><span style='color:windowtext;text-decoration:none'>6</span></a></span></sup><span
style='font-family:"Arial","sans-serif"'>. However, this classical view
neglects the possibility that motile species may actively respond to turbulent
cues to avoid layers of strong turbulence</span><sup><span style='font-size:
9.0pt;font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref7"
title="Lozovatsky, I., Lee, J., Fernando, H. J. S., Kang, S. K. & Jinadasa, S. U. P. Turbulence in the East China Sea: the summertime stratification. J. Geophys. Res. Oceans. 120, 1856–1871 (2015)"
id=ref-link-8><span style='color:windowtext;text-decoration:none'>7</span></a></span></sup><span
style='font-family:"Arial","sans-serif"'>. Here we report that phytoplankton,
including raphidophytes and dinoflagellates, can actively diversify their
migratory strategy in response to hydrodynamic cues characteristic of
overturning by Kolmogorov-scale eddies. Upon experiencing repeated overturning
with timescales and statistics representative of ocean turbulence, an
upward-swimming population rapidly (5–60</span><span style='font-family:
"Cambria Math","serif"'> </span><span style='font-family:"Arial","sans-serif"'>min)
splits into two subpopulations, one swimming upward and one swimming downward.
Quantitative morphological analysis of the harmful-algal-bloom-forming
raphidophyte<span class=apple-converted-space> </span><i>Heterosigma
akashiwo</i><span class=apple-converted-space> </span>together with a
model of cell mechanics revealed that this behaviour was accompanied by a
modulation of the cells’ fore–aft asymmetry. The minute magnitude
of the required modulation, sufficient to invert the preferential swimming
direction of the cells, highlights the advanced level of control that
phytoplankton can exert on their migratory behaviour. Together with
observations of enhanced cellular stress after overturning and the typically
deleterious effects of strong turbulence on motile phytoplankton</span><sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif"'><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref5"
title="Sullivan, J. M., Swift, E., Donaghay, P. L. & Rines, J. E. B. Small-scale turbulence affects the division rate and morphology of two red-tide dinoflagellates. Harmful Algae 2, 183–199 (2003)"
id=ref-link-9><span style='color:windowtext;text-decoration:none'>5</span></a>,<span
class=apple-converted-space> </span><a
href="http://www.nature.com/nature/journal/vaop/ncurrent/full/nature21415.html#ref8"
title="Thomas, W. H. & Gibson, C. H. Effects of small-scale turbulence on microalgae. J. Appl. Phycol. 2, 71–77 (1990)"
id=ref-link-10><span style='color:windowtext;text-decoration:none'>8</span></a></span></sup><span
style='font-family:"Arial","sans-serif"'>, these results point to an active
adaptation of<span class=apple-converted-space> </span><i>H. akashiwo</i><span
class=apple-converted-space> </span>to increase the chance of evading
turbulent layers by diversifying the direction of migration within the
population, in a manner suggestive of evolutionary bet-hedging. This migratory
behaviour relaxes the boundaries between the fluid dynamic niches of motile and
non-motile phytoplankton, and highlights that rapid responses to hydrodynamic
cues are important survival strategies for phytoplankton in the ocean.<o:p></o:p></span></p>
<p class=MsoNormal><o:p> </o:p></p>
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