Quantifying diatom aspirations: Mechanical properties of chain‐forming species

Diatoms developed a variety of mechanisms to form chain-like colonies, resulting in diverse morphologies and bulk mechanical properties. These properties affect translation, rotation, and deformation of colonies in ambient flows as well as their susceptibility to breakage by flow- and grazer-induced forces. Morphological characteristics of diatom chains have been extensively studied, yet no studies have examined their mechanical properties. We studied the flexibility of four morphologically distinct species (Lithodesmium undulatum, Stephanopyxis turris, Lauderia annulata, and Guinardia delicatula) by measuring their deflections when held across a capillary tip in developing pipe flow and applying simple beam theory and a finite-difference analysis of curvature to calculate flexural stiffness. Flexural stiffness varies greatly, with at least four orders of magnitude difference among the examined species (from 1.7 × 10−13 N m2, the most rigid, to 1.3 × 10−17 N m2, the most flexible), but two other species (Melosira nummuloides and a Thalassiosira sp.) were too flexible to measure with our apparatus. Vulnerability to breakage by flow also varied between species and, for species with heavily silicified joints between cells, was enhanced under nutrient depletion. These results highlight yet another attribute underlying the biodiversity of diatoms and their potential for utilizing highly differentiated ecological niches. Quantitative information from this study can now be used in the design of more mechanically realistic models that capture the dynamic coupling between elastic particles and flow to study diatom–flow interactions and their effects on nutrient acquisition, encounter with grazers, aggregate formation, and settling.

[1]  A. Sournia FORM AND FUNCTION IN MARINE PHYTOPLANKTON , 1982 .

[2]  I. Gebeshuber,et al.  Micromechanics in biogenic hydrated silica: Hinges and interlocking devices in diatoms , 2006 .

[3]  E. Boss,et al.  Nutrient fluxes to planktonic osmotrophs in the presence of fluid motion , 1996 .

[4]  A. Alldredge,et al.  Direct observations of the mass flocculation of diatom blooms: characteristics, settling velocities and formation of diatom aggregates , 1989 .

[5]  H. Dam,et al.  Sedimentation of phytoplankton during a diatom bloom : Rates and mechanisms , 1996 .

[6]  M. Pahlow,et al.  Impact of cell shape and chain formation on nutrient acquisition by marine diatoms , 1997 .

[7]  J. Howard,et al.  Flexural rigidity of microtubules and actin filaments measured from thermal fluctuations in shape , 1993, The Journal of cell biology.

[8]  E. Selander,et al.  Grazer‐induced chain length plasticity reduces grazing risk in a marine diatom , 2012 .

[9]  C. Schönenberger,et al.  Nanomechanics of microtubules. , 2002, Physical review letters.

[10]  Nigel Goldenfeld,et al.  Simple viscous flows: From boundary layers to the renormalization group , 2006, physics/0609138.

[11]  A. R. Loeblich,et al.  Evolution of the oceanic plankton , 1973 .

[12]  D. Barkley,et al.  The Onset of Turbulence in Pipe Flow , 2011, Science.

[13]  D. Mann,et al.  The origin and evolution of the diatoms: their adaptation to a planktonic existence , 2007 .

[14]  R. Margalef Life-forms of phytoplankton as survival alternatives in an unstable environment , 1978 .

[15]  V. Smetácek,et al.  Colonies of Phaeocystis globosa are protected by a thin but tough skin , 1999 .

[16]  Richard C. Dugdale,et al.  The effect of nutrient availability and temperature on chain length of the diatom, Skeletonema costatum , 2006 .

[17]  D. Wolf-Gladrow,et al.  The relationship between physical aggregation of phytoplankton and particle flux: a numerical model , 1992 .

[18]  R. Emlet ECHINODERM CALCITE: A MECHANICAL ANALYSIS FROM LARVAL SPICULES , 1982 .

[19]  T. Smayda,et al.  EXPERIMENTAL OBSERVATIONS ON THE FLOTATION OF MARINE DIATOMS. II. SKELETONEMA COSTATUM AND RHIZOSOLENIA SETIGERA , 1966 .

[20]  Y. Hiramoto,et al.  DIRECT MEASUREMENTS OF THE STIFFNESS OF ECHINODERM SPERM FLAGELLA , 1979 .

[21]  L. Fauci,et al.  Nutrient transport and acquisition by diatom chains in a moving fluid , 2008, Journal of Fluid Mechanics.

[22]  Victor Smetacek,et al.  Architecture and material properties of diatom shells provide effective mechanical protection , 2003, Nature.

[23]  Yves Engelborghs,et al.  Dynamical and mechanical study of immobilized microtubules with atomic force microscopy , 1996 .