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Diatom Morphogenesis
Scrivener Publishing
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Beverly, MA 01915-6106
Diatoms: Biology and Applications
Series Editors: Richard Gordon (dickgordoncan@xplornet.com) and Joseph Seckbach

(Joseph.Seckbach@mail.huji.ac.il)
Scope: The diatoms are a single-cell algal group, with each cell surrounded by a silica shell. The shells
have beautiful attractive shapes with multiscalar structure at 8 orders of magnitude, and have several
uses. 20% of the oxygen we breathe is produced by diatom photosynthesis, and they feed most of the
aquatic food chain in freshwaters and the oceans. Diatoms serve as sources of biofuel and electrical
solar energy production and are impacting on nanotechnology and photonics. They are important
ecological and paleoclimate indicators. Some of them are extremophiles, living at high temperatures or
in ice, at extremes of pH, at high or low light levels, and surviving desiccation. There are about 100,000
species and as many papers written about them since their discovery over three hundred years ago. The
literature on diatoms is currently doubling every ten years, with 50,000 papers during the last decade
(2006-2016). In this context, it is timely to review the progress to date, highlight cutting-edge discoveries,
and discuss exciting future perspectives. To fulfill this objective, this new Diatom Series is being launched
under the leadership of two experts in diatoms and related disciplines. The aim is to provide a comprehensive
and reliable source of information on diatom biology and applications and enhance interdisciplinary
collaborations required to advance knowledge and applications of diatoms.
Publishers at Scrivener
Martin Scrivener (martin@scrivenerpublishing.com)
Phillip Carmical (pcarmical@scrivenerpublishing.com)
Diatom Morphogenesis
Edited by
Vadim Annenkov,
Limnological Institute, Siberian Branch of Russian Academy of Sciences
Joseph Seckback
The Hebrew University of Jerusalem, Israel
and
Richard Gordon
Gulf Specimen Marine Laboratory & Aquarium, Panacea, FL, USA
and Wayne State University, Detroit, MI, USA
This edition first published 2022 by John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA and Scrivener
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Library of Congress Cataloging-in-Publication Data
ISBN 978-1-119-487951
Cover image: Colored scanning electron micrographs (SEMs) of a morphogenetic sequence of the diatom Fragillaria capucina
var. mesolepta by Dr. Mary Ann Tiffany, Biology Department, San Diego State University, USA
Sample taken from Lake Murray (a freshwater San Diego Reservoir) on 3/18/2000.
Cover design by Russell Richardson
Set in size of 11pt and Minion Pro by Manila Typesetting Company, Makati, Philippines
Printed in the USA
10 9 8 7 6 5 4 3 2 1
Contents
Preface xv
Part 1: General Issues 1
1 Introduction for a Tutorial on Diatom Morphology 3
Kalina Manoylov and Mohamed Ghobara
1.1 Diatoms in Brief 3
1.2 Tools to Explore Diatom Frustule Morphology 7
1.3 Diatom Frustule 3D Reconstruction 12
1.3.1 Recommended Steps to Understand the Complex Diatom
Morphology: A Guide for Beginners 13
1.4 Conclusion 15
Acknowledgements 15
References 15
2 The Uncanny Symmetry of Some Diatoms and Not of Others: A Multi-Scale
Morphological Characteristic and a Puzzle for Morphogenesis 19
Janice L. Pappas, Mary Ann Tiffany and Richard Gordon
2.1 Introduction 20
2.1.1 Recognition and Symmetry 21
2.1.2 Symmetry and Growth 24
2.1.3 Diatom Pattern Formation, Growth, and Symmetry 25
2.1.4 Diatoms and Uncanny Symmetry 27
2.1.5 Purpose of This Study 28
2.2 Methods 28
2.2.1 Centric Diatom Images Used for Analysis 28
2.2.2 Centric Diatoms, Morphology, and Valve Formation 34
2.2.3 Image Entropy and Symmetry Measurement 36
2.2.4 Image Preparation for Measurement 37
2.2.5 Image Tilt and Slant Measurement Correction for Entropy Values 38
2.2.6 Symmetry Analysis 39
2.2.7 Entropy, Symmetry, and Stability 40
2.2.8 Randomness and Instability 42
2.3 Results 43
2.3.1 Symmetry Analysis 43
2.3.2 Valve Formation—Stability and Instability Analyses 49
v
vi Contents
2.4 Discussion 51
2.4.1 Symmetry and Scale in Diatoms 55
2.4.2 Valve Formation and Stability 56
2.4.3 Symmetry, Stability and Diatom Morphogenesis 57
2.4.4 Future Research—Symmetry, Stability and Directionality in Diatom
Morphogenesis 58
References 59
3 On the Size Sequence of Diatoms in Clonal Chains 69
Thomas Harbich
3.1 Introduction 70
3.2 Mathematical Analysis of the Size Sequence 73
3.2.1 Alternative Method for Calculating the Size Sequence 73
3.2.2 Self-Similarity and Fractal Structure 75
3.2.3 Matching Fragments to a Generation Based on Known Size
Indices of the Fragment 76
3.2.4 Sequence of the Differences of the Size Indices 78
3.2.5 Matching Fragments to a Generation Based on Unknown Size
Indices of the Fragment 80
3.2.6 Synchronicity of Cell Divisions 81
3.3 Observations 82
3.3.1 Challenges in Verifying the Sequence of Sizes 82
3.3.2 Materials and Methods 83
3.3.3 Investigation of the Size Sequence of a Eunotia sp. 84
3.3.4 Synchronicity 86
3.4 Conclusions 87
Acknowledgements 88
Appendix 3A L-System for the Generation of the Sequence of Differences
in Size Indices of Adjacent Diatoms 88
Appendix 3B Probability Consideration for Loss of Synchronicity 89
References 91
4 Valve Morphogenesis in Amphitetras antediluviana Ehrenburg 93
Mary A. Tiffany and Bonnie L. Hurwitz
4.1 Introduction 93
4.2 Material and Methods 94
4.3 Observations 94
4.3.1 Amphitetras antediluviana Mature Valves 94
4.3.2 Amphitetras antediluviana Forming Valves 96
4.3.3 Amphitetras antediluviana Girdle Band Formation 101
4.4 Conclusion 101
Acknowledgments 102
References 102
Glossary 104
Contents vii
Part 2: Simulation 105

5 Geometric Models of Concentric and Spiral Areola Patterns
of Centric Diatoms 107
Anton M. Lyakh
5.1 Introduction 107
5.2 Set of Common Rules Used in the Models 109
5.3 Concentric Pattern of Areolae 109
5.4 Spiral Patterns of Areolae 110
5.4.1 Unidirectional Spiral Pattern 111
5.4.2 Bidirectional Spiral Pattern 113
5.4.3 Common Genesis of Unidirectional and Bidirectional Spiral Patterns 113
5.5 Conversion of an Areolae-Based Model Into a Frame-Based Model 114
5.6 Conclusion 114
Acknowledgements 114
References 115
6 Diatom Pore Arrays’ Periodicities and Symmetries in the Euclidean Plane:
Nature Between Perfection and Imperfection 117
Mohamed M. Ghobara, Mary Ann Tiffany, Richard Gordon and Louisa Reissig
6.2 Materials and Methods 122
6.2.1 Micrograph Segmentation 123
6.2.2 Two-Dimensional Fast Fourier Analysis and Autocorrelation
Function Analysis 123
6.2.3 Lattice Measurements and Recognition 123
6.2.4 Accuracy of 2D ACF-Based Calculations 125
6.2.5 The Perfection of the Unit Cell Parameters Between Different
Parts (Groups of Pore Arrays) of the Same Valve
and the Same Micrograph 126
6.3 Results and Discussion 126
6.3.1 Toward Standardization of the Methodology for the Recognition
of 2D Periodicities of Pore Arrays in Diatom Micrographs 126
6.3.1.1 Using Two-Dimensional Fast Fourier Transform Analysis 126
6.3.1.2 Using Two-Dimensional Autocorrelation Function 131
6.3.1.3 The Accuracy of Lattice Parameters’ Measurements
Using the Proposed 2D ACF Analysis 134
6.3.2 Exploring the Periodicity in Our Studied Micrographs
and the Possible Presence of Different Types of 2D Lattices
in Diatoms 137
6.3.2.1 Irregular Pore Scattering (Non-Periodic Pores) 137
6.3.2.2 Linear Periodicity of Pores in Striae (1D Periodicity) 138
6.3.2.3 The Different 2D Lattices in Diatom Pore Arrays 140
6.3.3 How Perfectly Can Diatoms Build Their 2D Pore Arrays? 146
6.3.3.1 Variation of the 2D Lattice Within the Connected
Pore Array of the Valve 146
viii Contents
6.3.3.2 Comparison of 2D Lattice Parameters and Degree

of Perfection of Distinct Pore Array Groups in the Same
Micrograph and Valve but With Different Rotational
or Reflection Symmetry 148
6.3.3.3 The Perfection of 2D Lattices of Diatom Pore Arrays
Compared to Perfect (Non-Oblique) 2D Bravais Lattices 148
6.3.4 Planar Symmetry Groups to Describe the Whole Diatom Valve
Symmetries and Additionally Describe the Complicated
2D Periodic Pore Arrays’ Symmetries 149
6.3.4.1 Rosette Groups 150
6.3.4.2 Frieze Groups 151
6.3.4.3 Wallpaper Groups 153
6.4 Conclusion 153
Acknowledgment 154
Glossary 154
References 155
7 Quantified Ensemble 3D Surface Features Modeled as a Window
on Centric Diatom Valve Morphogenesis 159
Janice L. Pappas
7.1.1 From 3D Surface Morphology to Morphogenesis 160
7.1.2 Geometric Basis of 3D Surface Models and Analysis 163
7.1.3 Differential Geometry of 3D Surface 163
7.1.4 3D Surface Feature Geometry and Morphological Attributes 165
7.1.5 Centric Diatom Taxa Used as Exemplars in 3D Surface Models
for Morphogenetic Analysis 166
7.1.6 Morphogenetic Descriptors of Centric Diatoms in Valve Formation
as Sequential Change in 3D Surface Morphology 166
7.1.7 Purposes of This Study 167
7.2 Methods 168
7.2.1 Measurement of Ensemble Surface Features and 3D Surface
Morphology: Derivation and Solution of the Jacobian, Hessian,
Laplacian, and Christoffel Symbols 168
7.2.1.1 The Jacobian of 3D Surface Morphology 168
7.2.1.2 Monge Patch 169
7.2.1.3 First and Second Fundamental Forms and Surface
Characterization of the Monge Patch 169
7.2.1.4 3D Surface Characterization via Gauss and Weingarten
Maps and the Fundamental Forms 170
7.2.1.5 Peaks, Valleys, and Saddles of Surface Morphology
and the Hessian 170
7.2.1.6 Smoothness as a Characterization of Surface Morphology
and the Laplacian 171
7.2.1.7 Point Connections of 3D Surface Morphology
and Christoffel Symbols 171
Contents ix
7.2.1.8 Protocol for Using Centric Diatom 3D Surface Models

and Their Ensemble Surface Features in Valve
Formation Analysis 173
7.3 Results 174
7.4 Discussion 184
7.4.1 Ensemble Surface Features and Physical Characteristics
of Valve Morphogenesis 186
7.4.2 Factors Affecting Valve Formation 187
7.4.3 Diatom Growth Patterns—Buckling and Wave Fronts 187
7.4.4 Valve Formation, Ensemble Surface Features, and Self-Similarity 189
7.4.5 Diatom Morphogenesis: Cytoplasmic Inheritance
and Phenotypic Plasticity 189
7.4.6 Phenotypic Variation and Ensemble Surface Features:
Epistasis and Canalization 190
7.5 Conclusions 190
Acknowledgment 191
References 191
8 Buckling: A Geometric and Biophysical Multiscale Feature
of Centric Diatom Valve Morphogenesis 195
Janice L. Pappas and Richard Gordon
8.2 Purpose of Study 197
8.3 Background: Multiscale Diatom Morphogenesis 198
8.3.1 Valve Morphogenesis—Schemata of Schmid and Volcani
and of Hildebrand, Lerch, and Shrestha 198
8.3.2 Valve Formation—An Overview at the Microscale 199
8.3.3 Valve Formation—An Overview at the Meso- and Microscale 200
8.3.4 Valve Formation—An Overview at the Meso- and Nanoscale 200
8.4 Biophysics of Diatom Valve Formation and Buckling 201
8.4.1 Buckling as a Multiscale Measure of Valve Formation 201
8.4.2 Valve Formation—Cytoplasmic Features and Buckling 202
8.4.3 Buckling: Microtubule Filaments and Bundles 203
8.4.4 Buckling: Actin Filament Ring 204
8.5 Geometrical and Biophysical Aspects of Buckling
and Valve Formation 205
8.5.1 Buckling: Geometry of Valve Formation as a Multiscale Wave Front 205
8.5.2 Buckling: Valve Formation and Hamiltonian Biophysics 207
8.5.3 Buckling: Valve Formation and Deformation Gradients 208
8.5.4 Buckling: Multiscale Measurement With Respect to Valve Formation 210
8.5.5 Buckling: Krylov Methods and Association of Valve Surface
Buckling With Microtubule and Actin Buckling 210
8.6 Methods 211
8.6.1 Constructing and Analyzing 3D Valve Surface and 2D Microtubule
and Actin Filament Models 211
x Contents
8.6.2 Krylov Methods: Associating Valve Surface With Microtubule

and Actin Filament Buckling 212
8.7 Results 212
8.8 Conclusion 216
References 223
9 Are Mantle Profiles of Circular Centric Diatoms a Measure
of Buckling Forces During Valve Morphogenesis? 231
Janice L. Pappas and Richard Gordon
9.2 Methods 233
9.2.1 Background: Circular Centric 2D Profiles and 3D Surfaces
of Revolution 236
9.3 Results 238
9.3.1 Approximate Constant Profile Length Representing Approximate
Same Sized Valves 239
9.3.2 Change in Profile Length Representing Size Reduction During
Valve Morphogenesis 240
9.3.2.1 Inferences About Complementarity and Heterovalvy 242
9.3.3 Are Profiles Measures of Buckling Forces During Valve
Morphogenesis? 243
9.4 Discussion 245
9.4.1 Laminated Structures and Mantle Buckling Forces Affecting
the Valve Profile 247
9.5 Conclusion 248
Acknowledgement 248
References 248
Part 3: Physiology, Biochemistry and Applications 251

10 The Effect of the Silica Cell Wall on Diatom Transport and Metabolism 253
Mark Hildebrand
Publications by and about Mark Hildebrand 254
11 Diatom Plasticity: Trends, Issues, and Applications on Modern and Classical
Taxonomy, Eco‑Evolutionary Dynamics, and Climate Change 261
Lawrence Victor D. Vitug
11.2 Model Species: Phaeodactylum tricornutum 262
11.3 Transformation Mechanisms of P. tricornutum 263
11.4 Future Advances in the Phenotypic Plasticity on P. tricornutum 263
11.4.1 Genomic and Molecular Mechanisms in Diatom Phenotypic
Plasticity 263
11.4.2 Biogeography of Diatoms 263
11.4.3 Eco-Evolutionary Dynamics Approach on Diatoms Phenotypic
Plasticity 264
Contents xi
11.4.4 Adaptive Behavior and Evolutionary Changes in Diatoms Linking

to Diatom Plasticity 265
11.4.5 Climate Change and Phenotypic Plasticity 265
11.5 Conclusion 265
References 265
12 Frustule Photonics and Light Harvesting Strategies in Diatoms 269
Johannes W. Goessling, Yanyan Su, Michael Kühl and Marianne Ellegaard
12.2 Light Spectral Characteristics and Signaling 274
12.2.1 Variation of Light Regimes 274
12.2.2 Light Perception and Signaling 275
12.3 Photosynthesis and Photo-Protection in Diatoms 276
12.3.1 Pigment-Based Light Absorption 276
12.3.2 Molecular Photo-Protection Mechanisms 276
12.3.3 Intracellular Structural Adaptation in Response to Light 277
12.3.4 Motility as a Unique Photo-Protection Mechanism 278
12.4 Frustule Photonics Related to Diatom Photobiology 279
12.4.1 An Extracellular Structure With Optical Properties 279
12.4.2 Intraspecific and Intra-Individual Variation
of Frustule Periodicity 281
12.4.3 Photonic Crystal Properties 281
12.4.4 Light Confinement and Focusing 282
12.4.5 Scattering and Dispersion of Light 283
12.4.6 Attenuation of UV Light for Photo-Protection 283
12.5 Frustule Photonics in Light of Niche Differentiation 285
12.6 Conclusion 291
References 292
13 Steps of Silicic Acid Transformation to Siliceous Frustules:
Main Hypotheses and Discoveries 301
Vadim V. Annenkov, Elena N. Danilovtseva and Richard Gordon
13.2 Penetration of the Boundary Layer: The Diatom as an Antenna
for Silica 303
13.3 Getting Past the Cloud of Extracellular Material 304
13.4 Adsorption of Silica Onto the Outer Organic Coat of the Diatom 305
13.5 Getting Past the Silica Frustule or Through Its Pores 306
13.6 Getting Past the Inner Organic Coat, the Diatotepum 306
13.7 Transport of Silica Across the Cell Membrane 307
13.8 Cytoplasm Storage and Trafficking of Silica
to the Places of Synthesis of the Frustule Parts 309
13.9 Transport and Patterning of Silica Across the Silicalemma 311
13.10 Precipitation and Morphogenesis of the Nascent Valve Within
the Silicalemma 314
13.11 Thickening of the Valve Within the Silicalemma 319
xii Contents
13.12 Exteriorization of the Valve 321

13.13 Future Work Needed 321
13.14 Conclusion 323
References 326
14 The Effects of Cytoskeletal Inhibitors on Diatom Valve Morphogenesis 349
Yekaterina D. Bedoshvili and Yelena V. Likhoshway
14.2 Cytoskeleton and Its Role in Cell Morphogenesis 350
14.3 Abnormalities of Diatom Valve Morphogenesis Induced by Cytoskeleton
Inhibitors 352
14.4 Conclusion 358
Acknowledgment 360
References 360
15 Modeling Silicon Pools in Diatoms Using the Chemistry Toolbox 365
Argyro Spinthaki and Konstantinos D. Demadis
15.1 Diatoms 365
15.2 “Silicon Pools” Biology 366
15.3 Silica Particle Formation From Silicic Acid 366
15.4 Stabilization of “Soluble” Silica Species (Monosilicic and Disilicic Acids) 370
15.4.1 Cationic Polymers 370
15.4.2 Neutral (Uncharged) Polymers 372
15.4.3 Zwitterionic Polymers 373
15.4.4 Blends of Cationic/Anionic Polymers 375
15.5 Chemical Mechanisms 376
15.6 Conclusions/Perspectives 377
Acknowledgments 378
References 378
16 The Mesopores of Raphid Pennate Diatoms: Toward Natural Controllable
Anisotropic Mesoporous Silica Microparticles 383
Mohamed M. Ghobara, Richard Gordon and Louisa Reissig
16.2 Morphology and Very Fine Ultrastructure of Diatom Frustules 386
16.3 Synthetic Mesoporous Silica 391
16.4 The Potential of Raphid Pennates’ Mesoporous Bio-Silica, Similarities,
and Dissimilarities Compared With Synthetic MSM/Ns 393
16.4.1 The Current Potential of Diatom Porous Silica in Applications 393
16.4.2 Why Should We Be Interested in the Mesoporous Silica
of Raphid Pennate Frustules if the Frustules of Other Species
With Larger Pores Work? 393
16.4.3 Similarities and Dissimilarities Compared With Synthetic
MSM/Ns 394
16.5 Our Ability to Control the Diatom Frustule’s Ultrastructure 396
16.5.1 Physicochemical Parameters Alteration Approach 397
16.5.2 Genetic Engineering Approach 398
Contents xiii
16.6 Conclusion 399

Acknowledgment 399
References 399
Glossary 408
Index 411
Preface
Diatoms comprise a large, unicellular eukaryotic algal group that thrives mainly in aque-
ous environments: in fresh water, and in ponds, lakes and oceans. They may be attached to
benthic substrates, in moist habitats or in floating debris and on macrophytes, and as phy-
toplankton; they form a substantial basis of aquatic food webs. They are ubiquitous, being
distributed among various ecological locations. Among this group are some extremophiles
with varying features, such as living in high temperatures, surviving desiccation, or in ice
and at extreme ranges of pH. Some 20%–30% of the oxygen we breath is produced by dia-
tom photosynthesis.
Vegetative cells of diatoms are diploid (2N), and meiosis can take place, producing male
and female gametes fusing to zygotes which grow to auxospores.
One of their specific features is that their chemical composition includes siliceous (glassy)
cell walls (frustules). Their exoskeleton is made of two halves called “valves” that fit inside
one another, secured by silica “girdle bands”.
Diatoms’ fine structure is very impressive as revealed by transmission electron micro-
scope, scanning microscope, and atomic force micrographs. The appearance of their cells
is strikingly unique, and their shells are beautiful attractive shapes, with 60,000 to 200,000
species.
Why Valve Morphogenesis is Important?

Because there is so much detail in their silica wall shapes, spanning 8 orders of magni-
tude, diatoms are model organisms for single-cell morphogenesis. The problem of single
cell morphogenesis has a long history, as yet unsolved, and perhaps diatoms rather than
desmids and ciliates will now lead the way, especially given their 200 million years fossil
record. This may further be because diatoms serve as a source of biofuel, food supplements
and lipids and serve as significant material for nanotechnology. Thus, they are of very wide
interest.
This volume focuses on the morphogenesis of diatoms, namely, the formation of their
shape and the initial developmental steps.
The chapters were contributed by experts on morphological diatoms. The authors stem
from the USA, Russia, Denmark, Germany, Greece, Israel, and Portugal.
xv
xvi Preface
Topics Addressed in This Volume

Topics include computer simulation of morphogenesis, silicic acid to silica frustules, inhibi-
tion in valve morphogenesis, pores within frustules, mesopores of pennate diatoms, frustule
photonics and light harvesting, clonal chains, silica cell wall, geometric models of centric
diatoms, morphology, surface features, buckling of valve morphogenesis, on mantle pro-
files, genetic-biochemical approaches, modeling silicon pools, valve morphogenesis, dia-
tom teratology in taxonomy, phenotypic plasticity, geometric and morphometric analysis,
silica morphogenesis in sister algae, and the uncanny symmetry of some diatoms.
This volume is the third book in the series Diatoms: Biology and Applications. The first
book, Diatoms: Fundamentals and Applications appeared in 2019, and was edited by Joseph
Seckbach and Richard Gordon. The second book, Diatom Gliding Motility, was published in
September 2021 and is edited by Stanley A. Cohn, Kalina M. Manoylov and Richard Gordon.
We would like to thank the authors, the reviewers, the guest editor (Vadim V. Annenkov),
and our publisher Martin Scrivener of Massachusetts, USA.
Joseph Seckbach
Hebrew University Jerusalem, Israel
September 2021
Part 1
GENERAL ISSUES
1
Introduction for a Tutorial on Diatom Morphology
Kalina Manoylov1* and Mohamed Ghobara2
1
Dept. of Biological & Environmental Sciences, Georgia College and State University,
Milledgeville, GA, United States
2
Department of Physics, Freie Universitat Berlin, Berlin, Germany
Abstract
Diatoms are an exceptionally successful group of unicellular microalgae with a large contribution
of global primary production in aquatic environments and contributing a significant amount of
oxygen to both hydro- and atmospheres. They are fascinating throughout their life and even after
death, thanks to their unique cell walls made from ornamented silica. The diatoms include centric
species, which may have radial or polar symmetry, and pennates, which include araphid, mon-
oraphid, and biraphid species. Several applications have utilized diatomite, i.e., the fossil form of
diatom frustules. To date, many diatoms’ secrets have been understood; however, there are still
more hidden. Thus, there is a need for more research on diatom basic biology and applications.
Seeking this goal, more people should be encouraged to work on diatoms. Often novice research-
ers are overwhelmed by the terminology associated with the diverse morphology, the discrepancy
between expected features for published descriptions, and the actual observation of those complex
3D organisms, which can be a barrier for more progress. Here, we provide a brief introduction to
the beginners with a guide to approach the complex diatom morphology focusing on the tools that
can be used for its study.
Keywords: Diatom morphology, tutorial, LM and SEM, frustule morphology
1.1 Diatoms in Brief

Diatoms are unicellular, eukaryotic, microscopic algae (range from 1.5 µm to 5 mm
in length, or diameter [1.9]), which maintain large population numbers and contrib-
ute considerably to the carbon and oxygen cycle on a global scale [1.8]. This ecologi-
cally successful group of algae is present in all aquatic habitats e.g. [1.1, 1.2] and even
extends to humid terrestrial places. In aquatic habitats, diatoms are present in the pho-
tic zone, i.e., the region of water that light strongly penetrates, as well as in the benthic
zone, i.e., the lowest level of water adjacent to the bottom with dim light conditions,
depending on water column height and water’s turbidity. Diatoms can exist as planktonic
*Corresponding author: kalina.manoylov@gcsu.edu
Vadim Annenkov, Joseph Seckback and Richard Gordon (eds.) Diatom Morphogenesis, (3–18) © 2022 Scrivener
Publishing LLC
3
4 Diatom Morphogenesis
(i.e., suspended in the water column), benthic (i.e., living near the bottom), epiphytic
(i.e., adhered to aquatic plants [1.19], Figures 1.2c–d), or epizoic (i.e., adhered to a wide
range of marine organisms such as crustaceans, mollusks, and vertebrates [1.19, 1.38]), or
epilithic (i.e., attached completely or partially to submerged rocks). The adhesion ability
of some diatoms is related to their mucilage secretion from specialized areas within their
rigid cell walls (such as examples shown in Figures 1.1d and 1.2c–d). Some diatoms can
form colonies in different arrangements such as chains and ribbons (examples shown in
Figures 1.1 and 1.2).
Diatoms are a unique group of microalgae for several reasons, but one of the most
notable and unique differences is the glass cell walls they possess [1.45]. This cell wall is
called the “frustule” and is composed of amorphous hydrated silica that gives it unique
properties. In general, the frustule is composed of two pieces that fit together like a petri-
dish, meaning that the lower part of the frustule, called the hypotheca, sits inside of the
upper part of the frustule, called the epitheca. The frustule volume extends by adding
strips of silica called girdle bands (cingulum) to the mantle, i.e., the curved edge of the
valve. It should be noted that there are plenty of frustule morphologies that vary between
taxa.
Diatoms reproduce both asexually (visible in Figure 1.6) and sexually. Most of the time,
they reproduce asexually via binary fission through adding new hypovalves to the parent
valves. Those new hypovalves are synthesized inside the silica deposition vesicle (SDV).
Only after the new hypovalves have completely synthesized and the protoplast cleavage, as
well as the exocytosis of siliceous parts, has occurred, the final splitting apart will occur,
leaving two daughter diatoms in place. Because the SDV forms inside of each new cell
(a) (b) (c)
(d) (e) (f)
Figure 1.1 Living diatoms as observed under LM, brightfield. (a) Two living cells of Actinoptychus senarius
(Ehrenberg) Ehrenberg at the valve view. (b) The valve view of a single living cell of Coscinodiscus wailesii
Gran and Angst. (c) The girdle view of a single living cell of Coscinodiscus granii L.F. Gough. (d) Two living
cells of Achnanthes brevipes C. Agardh at the girdle view attached to each other with a prolonged stalk for the
attachment to the substrate. (e) A living colony of Stephanopyxis turris (Greville) Ralfs with visible linking
spines. (f) A living colony of Odontella longicruris (Greville) M.A. Hoban with discoid chloroplasts. Copyright
reserved Mary Ann Tiffany, used with her permission. The identification was carried out by Mary Ann Tiffany.
All the scale bars are 50 µm.
Tutorial on Diatom Morphology 5
(a) (b)
10 µm
10 µm
10 µm 10 µm
(c) (d)
(h)
10 µm
10 µm
(f)
10 µm
10 µm
(e) (g)
Figure 1.2 Live centric (a, b) and pennate (c–h) diatoms. (a, b) Pleurosira laevis (Ehrenberg) Compère shown
from girdle view, frustules with numerous girdle bands in straight filaments with discoid chloroplasts, chains
connected with mucilage pads released from ocelli; in (b), visible diameter size restoration within the chain;
(c, d) Epiphytic diatoms on Cladophora glomerata (Linnaeus) Kützing, in (c) focus on Cocconeis spp. With
visible one flat C-shaped plastid; in (d) focus on Rhoicosphenia spp.; (e) Cymbella sp. partial valve and girdle
views, visible chloroplast bridge connecting the chloroplast plates; (f) Eunotia cf. camelus Ehrenberg in girdle
view with visible discoid chloroplasts; (g) Amphora ovalis (Kützing) Kützing with H shaped chloroplast;
(h) Rhoicosphenia sp. girdle view with visible lobes of the plastid. Scale bars, 10 µm. These micrographs were
obtained and identified by KMM.
before splitting into two, each new cell creates a new interior of the petri-dish structure.
What this means is that the cell that originally contained the upper part of the petri dish
(the epitheca) remains the same size, whereas the cell that originally contained the lower
part of the petri-dish (the hypotheca) becomes smaller, since it has now built a smaller
hypovalve to fit into it. Repeated cell division, therefore, leads to some part of the resulting
(a) (b)
3 5
4
1
Figure 1.3 Specific diatom morphology gleaned from images with whole and partial valves views of Navicula
oblonga (Kützing) Kützing; (a) live linear-lanceolate cell with visible two plates like brown chloroplasts,
visible linear striae, and proximal raphe ends deflected slightly toward the secondary side. (b) Valve view
after cleaning, axial area is linear, widening toward the central area and about twice the width of the raphe.
The central area orbicular. The raphe is lateral, becoming filiform near the proximal ends, which are simple.
Central striae do not reach valve edge. These micrographs were obtained and identified by KMM.
Details shown:
1. Central area is more or less orbicular and two to three times wider than the axial area. Proximal raphe ends
are simple and barely wider than the raphe. Striae are finely lineate and the individual areolae are difficult to
distinguish.
2. Round, subsidiary vacuoles on each side of the nucleus visible behind the glass cell wall and chloroplasts;
axial area outlines by lineate striae.
3. Terminal bent striae (terminal striae convergent at the margins and bent back toward the central area).
Striae are radiate next to the axial area.
4. Voigt discontinuity identifies the secondary side of the valve morphogenesis. Ontogeny in diatoms varies
with morphology; in Naviculoid diatoms, the secondary side shows the completion of silica deposition around
the raphe.
5. Distal raphe positioned on the broad, rounded apices and curved toward the primary side of the valve in the
opposite direction when compared to the proximal raphe ends. Scale bars, 10 µm.
population becoming smaller and smaller. Were asexual reproduction the only method by
which diatoms reproduces, this could lead the population eventually to become vulnerable
to dying out, but diatoms are ingenious and have gotten around this problem. At some
point, sexual reproduction is initiated by a number of steps, including meiotic divisions
to produce male and female gametes. These cells can find each other, fuse to form a zygote
and create a structure known as an auxospore, out of which a new large cell of the diatom
species will form, restoring its optimal size, which also depends on the environmental cir-
cumstances surrounding the auxospores. Some new research proposes chemical communi-
cation with pheromones between the male and female gametes [1.20].
Frustule morphogenesis, deposits SDVs and needs more research with new tools.
However, it has been established that the silica morphogenesis of centric species will begin
at the center of the valve, and it begins by creating a primary rib in pennate species [1.21].
Completion of the sternum around the raphe slit morphologically can be identified with
the Voight discontinuity (Figure 1.3b). From that onset within the mother frustule, the
silica will continue to form outward to complete the shape as well as inward to create more
layers, with the oldest silica being on the most outside layer [1.46]. The silicic acid (or its
anions) is taken from the environment, condensed, associated with proteins synthesized by
the endoplasmic reticulum and packaged in a globular vesicle in the Golgi apparatus. Then
finally, these vesicles (silica deposition vesicles) are transported by microtubules, likely in a
genetically predetermined pattern, and delivered to the new valve interface. These are not
the only groups that pull silicic acid (an inorganic compound contains silicon) out of the
water and use it to make a frustule, but diatoms do it uniquely.
Diatom frustules are porous with multilayer, multiscalar porosity, a property that is
unique for each species, giving frustules their beautiful ornamentation [1.17]. The major
bigger pores within the valves are called “areolae” and usually arranged in rows known as
“striae”, which could be either branched or not. In the most general way, diatoms can be
divided into centric and pennate diatoms, which are classified based on the valve symme-
try. Centric diatoms are radially symmetric and lack raphes. Pennate diatoms usually have
bilateral symmetry and there can be no, one, or two raphes. Pennate diatoms can further
be classified based on variations in the position of the raphe on valve. The raphe is used for
motility [1.4] and attachment [1.12]. Sometimes, the frustules are also covered in spines,
which can allow some species to hook together and form chains (Figure 1.1e).
The frustule’s morphological features of diatoms are required for identification.
Specialized terminology has been collected in [1.5–1.7, 1.15, 1.16], and a general guide to
the literature is in [1.10]. Characters continue to be discovered and new descriptive termi-
nologies are proposed [1.23].
1.2 Tools to Explore Diatom Frustule Morphology

The beauty of diatoms was missed until the early, curious microscopists started observing
ambiguous glassy microorganisms under their optical microscopes in the 18th century [1.22,
1.38]. Although the light microscope (LM) helped us to reveal the diatoms’ world, diatom
frustules also helped the microscopists in developing and testing the quality and resolution
of their optical microscopes [1.24, 1.25]. Since the nineteenth century, several works have
been published on diatoms, its morphology, and taxonomy by remarkable workers including
Kützing, Schmidt, Ehrenberg, Grunow, Hustedt, Krammer, Lange-Bertalot, and more (see
references in Round et al. [1.38]). They described both living cells and clean frustules exten-
sively using LM. The unique structure of diatom frustules under LM, with a variety of shapes
and symmetries, has captured a wide interest; however, most of the diatom’s real art, at the
nanoscale, was kept hidden. The limitations for observing frustule ultrastructure, especially
details below 200 nm, were solved after the invention of the electron microscope [1.26].
In 1936, the transmission electron microscope (TEM) was used to capture the first micro-
graph of a diatom frustule [1.26, 1.27], using it as a test object for the quality and resolution
of TEM. After that TEM was used to explore diatom ultrastructure. Following, the scanning
electron microscope (SEM) was invented and used extensively as a more effective tool for
exploring frustules morphology and ultrastructure [1.24, 1.28, 1.36, 1.38].
The details observed using the SEM and TEM reflected the beauty of diatoms when
many hidden details became observable. For instance, some bright striae under an optical
microscope appear as arrays of fine pores under the electron microscope (Figure 1.5a).
It was, to some extent, a kind of revolution for diatom classification and taxonomy with the
morphological details that became available down to 15 nm with SEM and below 10 nm
with TEM (Figure 1.5b). Nowadays, the observation of diatom frustule morphology and
ultrastructure using LM, SEM, and TEM became routine work for people working on ecol-
ogy, environment, forensic, nanotechnological, and other applications that concern frustule
ultrastructure, monitoring diatom species, and taxonomy.
Although 2D information can be collected from LM and TEM and the 3D-shape
appeared under SEM, the information about the surface topology, internal ultrastructure,
and siliceous element relationships within diatom frustules was missing. Therefore, more
tools were evolved and involved in the exploration and understanding of the 3D complex
ultrastructure of the frustule, which could be the reason for their various natural features,
including unique photonic, mechanical, and hydrokinetic properties [1.9, 1.45]. The new
tools include the atomic force microscope (AFM) and the focused ion beam SEM (FIB-
SEM) [1.32, 1.34, 1.35, 1.41].
(g)
(e) (f)
(c) (d)
(a) (b)
(h) (i) (j) (k) (l)

(m)
Figure 1.4 Cleaned diatoms in valve (g, h–j, m–r, u, v) and girdle views (a–f, k, l, s, t, w, x). (a–e, g)
Rhoicosphenia spp., frustules are clavate and strongly flexed, one valve is concave with long raphe branches and
the other valve convex with shortened raphe, different depth pseudosepta visible; (f, k, l) Gomphonema spp.
showing valve heterogeneity; (h) Gomphonella olivacea (Hornemann) Raben. (i) Planothidium lanceolatum
(Bréb. Ex Kütz.) Lange-Bert, rapheless valve shown with asymmetrical central area containing depression;
(j) Geissleria cascadensis (Sovereign) Stancheva and S. A. Spaulding, valves elliptic, with cuneate apices, coarse
areolae, three pairs of annulae are present at each apex; (m) Planothidium delicatulum (Kütz.) Round and
Bukht. Rapheless valve shown, lacking a central area and two middle striae spaced distantly. (Continued)
(q) (r)
(p)
(o)
(n)
(s) (t) (u) (v) (w) (x)
Figure 1.4 (Continued) Cleaned diatoms in valve (g, h–j, m–r, u, v) and girdle views (a–f, k, l, s, t, w, x).
(n) Gomphonema sp. valve heteropolar wider in the middle, axial area narrow, central area irregular outlined
by two shortened striae and opposite to a single striae finishing with an isolated pore, striae parallel toward the
headpole, radiate toward the foot pole; (o) Amphora ovalis, dorsal fascia visible and dorsal striae interrupted
transapically by intercostal ribs; (p) Gomphonema micropus Reichardt lanceolate valve with headpole widely
drawn out and wider than foot pole, striae radiate, central area unilaterally rectangular with shortened
central stria, on the opposite side longer striae finishing with a stigmoid; (q) Navicula genovefae Fusey valve
linear-lanceolate with rostrate broadly rounded apices, punctate striae radiate and curved, becoming nearly
parallel at the apices, less dense around the well-defined central area; (r) Cocconeis placentula Ehrenb. Valves
elliptic, striae radiate and interrupted by a hyaline ring positioned close to the valve margin, siliceous bridges
(imbriae extending from valvocopula) visible; (s) Amphora pediculus (Kütz.) Grunow focus from dorsal site
of two frustules; (t, u) Caloneis sp. on girdle view striae continue on valve mantle, on the linear valve view
with rounded apices, axial area is narrow, broadening to a transverse fascia; (v) Navicula cryptocephala Kütz.
Valve lanceolate with protracted apices and visible large, circular central area; (w) Mastogloia pseudosmithii
Sylvia S. Lee, E. E. Gaiser, Van de Vijver, Edlund, and S. A. Spaulding, evenly sized partecta (chambers on
the valvocopula) on both valves; (x) Navicula cf. tripunctata (O.F. Müll.) Bory. Scale bar, 10 µm. These
micrographs were obtained and identified by KMM.
In 1992, the first observation of diatoms using an AFM has been done [1.33]. In general,
AFM is used as an advanced tool to explore diatom ultrastructure providing information
in the Z-direction, with the ability to understand the surface topology of the frustule parts
with a nanoresolution. For instance, AFM observations of Coscinodiscus sp. clean valves
revealed a distinct dome topology for the cribellum, which was not observed before [1.34].
At the beginning of the current century, AFM was used in several works for understand-
ing the nanoscale ultrastructure and topology of frustule surfaces in a 3D manner. Today,
AFM is also used to explore the organic envelope, micromechanical properties, and to
(a) (b)
200 nm
HV mag spot WD det HPW 9/25/2017 5 µm
20.00 kV 20 000 × 5.0 8.3 mm LFD 20.7 µm 5:19:49 PM
Figure 1.5 (a) SEM of a single cleaned partially open frustule, two overlapping valves, of Nitzschia palea
(Kützing) W. Smith, and scale bar is 5 µm. The rows of pores (striae) that observed here cannot be observed
under LM for this species. (b) TEM of a close-up in Navicula sp. valve showing the hymenate pore occlusions
that will not be observed under SEM; scale bar is 200 nm. These micrographs were obtained and identified
by MG.
Epivalve of the
parent cell
Hypovalve of
the new upper
daughter cell
Hypovalve of
the new lower
Hypovalve of daughter cell
the parent cell
3 µm
Figure 1.6 A cross-section at the center of Coscinodiscus sp. cell collected and treated while binary fission
process was in progress, fabricated and captured by FIB-SEM. Reproduced from Xing et al. [1.42] under
a Creative Commons Attribution 4.0 International license.
understand the biomineralization processes of diatom frustules [1.35]. Luís et al. [1.35] can
be considered a good review for starting AFM studies on diatom frustules.
Furthermore, diatom valves seem to have a complex inner ultrastructure that can-
not be understood completely by observing the internal and external view of a given
valve surface using the previously mentioned tools. Although the multilayer, multisca-
lar porosity can be observed easily using such techniques, the internal anatomy and
relations of the siliceous elements of the frustule cannot be understood [1.41]. It was
usual to wish that the observation of a broken valve or girdle band at the right site and
right angle would help, otherwise, the complex inner structure remained unseen [1.41].
Thus, another advanced method was required for understanding the inner structures and
spatial relationships of the siliceous elements of a given diatom frustule. The FIB-SEM
was introduced as a solution for such a problem by cutting the diatom frustule parts at
nanoresolution to reveal the inner complex ultrastructure of a given valve or frustule
(Figure 1.6) [1.41]. Suzuki et al. [1.40] was the first work introduced using FIB-SEM for
making a cross-section in diatoms. Only a few articles are available using FIB-SEM and
the field is still growing. The acquired data using FIB-SEM could be used to reconstruct
Table 1.1 A summary of the major tools used to study diatom frustule morphology and its
ultrastructure.
LM TEM SEM AFM FIB-SEM
The date of Anonymous, 1703 Krause, 1936 Mid of 1960s [1.24] Linder et al., Suzuki et al., 2001
first known [1.22] [1.27] 1992 [1.33] [1.40]
observation of
diatoms using
the tool
Up-to-date The maximum Up-to-date, the The details less than Recently, the Having SEM as the
resolution resolution of highest TEM 15 nm was not resolution microscope
the common resolution resolved under can be below part of the
compound could be most of SEMs. 1 nm. device. Thus,
optical down to 50 Recently, an outbreak the resolution is
microscope picometer or has been achieved, dependent on
can be around even lower and the resolution this SEM.
200 nm. [1.29]. of SEM could be
Recently, the below 1 nm [1.39].
resolution
was enhanced
(down to 97
nm) using
special kind of
lenses [1.37].
When we should Observation of Observation Observation of the Observation Understanding

use? the presence of the fine outer ultrastructure of the 3D the inner
or absence of porosity including most topology ultrastructure
diatoms in a (mesopores) porosity. of a diatom of diatom
sample. present in Observation of frustule or its frustule or its
Identification of some genera, the overall 3D components. parts by cutting
diatoms on the like raphid ultrastructure of Measuring forces cross-sections
genus level. pennates the frustule or related with through it.
Enumeration (Figure 1.5b). different parts. both living Observation of
of diatom Observation of Identification at diatoms and the siliceous
frustules thin cross- the species and its cleaned elements
for different sections in subspecies level. frustules. structural
purposes. a valve or a relations within
girdle band. the frustule.
Observation Observation of
of the the whole 3D
cytoplasmic ultrastructure
components of the frustule
of thin cross- via the 3D
sections of reconstruction.
living cells
(living cells
anatomy).
(Continued)
Table 1.1 A summary of the major tools used to study diatom frustule morphology and its
ultrastructure. (Continued)
LM TEM SEM AFM FIB-SEM
The disadvantages The observations Only the tiniest The samples must The frustules This technique
for most of the parts of be coated with a must fix to sometimes
ultrastructure the valve, conductive layer, the substrate needs more
details will be like pore which in turn before sophisticated
limited. occlusions, could change the measuring. preparation
Either the girdle will be nano texture of the A very sensitive of the samples
view or the observed. frustule silica and tool with and more
valve view will The high energy probably pore sizes, complicated sophisticated
be available. electron beam thus the thickness precautions work to
may damage and smoothness to follow reconstruct the
some sensitive of the conductive to get the frustule or its
samples, so layer should be desired parts, however
it should be optimized and be results. it worth.
used wisely. thin as possible Related with the
without getting presence of the
nanoparticles on device, which
the top. usually is not
The high energy available for all
electron beam may research groups.
also damage some
sensitive samples
The regular resolution
keep the pore
occlusions of
very fine porosity
(below 10 nm)
hidden.
the overall 3D geometry of diatoms to carry out further computational simulations nec-
essary for diatom nanotechnology applications.
Finally, all the techniques mentioned were summarized in Table 1.1 to help beginners
and students choose between different tools on-demand.
1.3 Diatom Frustule 3D Reconstruction

Toward the complete understanding of the 3D structure of a given diatom frustule, a com-
prehensive 3D model can be created from the data collected from different characterization
techniques. This approach, which is designated as the 3D reconstruction of diatom frustules,
can be used for different purposes but is mainly for computer modeling. Oncoming tools
for the 3D reconstruction of diatom frustules are FIB-SEM [1.32, 1.42] and digital holo-
graphic microscopy (DHM) combined with SEM [1.30]. The combination of DHM and
SEM or AFM might give the ability to model and visualize microscopic 3D objects with a
high resolution in all directions [1.30]. Hildebrand et al. [1.32] introduced the ability for
the 3D reconstruction of subcellular architecture using FIB-SEM with new insights into the
architecture and synthesis process of both the siliceous and organic components inside the
cell. Xing et al. [1.42] is an inspiring reference for the 3D reconstruction of diatom frustule
using the 2D image series resulting from FIB-SEM.
1.3.1 Recommended Steps to Understand the Complex Diatom Morphology:

A Guide for Beginners
1. Fresh samples, if available, which means the sample is not diatomite (i.e., the fossil
form of diatom frustules) [1.31], allow evaluation of the physiological state of the pop-
ulation at the time of collection. The features of living cells might help in the identifi-
cation process, through the descriptions of colonial forms, cell attachment, extruded
materials, plastids, and nucleus position (for instance see Figures 1.1 and 1.2). Since dia-
tom frustules are rigid silica, they appear from two different viewpoints, the girdle view
(from the side) and the valve view (from the front) under LM Figure 1.4. Some genera
are easily identifiable in girdle view, such as Mastogloia and Amphora pediculus (Kütz.)
Grunow, others are not. Lately, confocal microscopy uses are very promising [1.3, 1.13],
also combining microscopic and molecular information allowed reclassification of a
population in culture annotated as a radial centric species related to Leptocylindrus dan-
icus Cleve, as an araphid pennate species in the staurosiroid lineage, within the genus
Plagiostriata [1.14].
2. The selection of the suitable cleaning procedure, depending on the source of diatoms, is a
crucial step to extract the siliceous frustule parts. Routinely, diatoms are identified and enu-
merated without their protoplasts where striae morphology & number, raphe shape, and
other morphological characteristics could be verified for identification (details in Figures
1.3b and 1.4). Among the best guides for the cleaning process could be Wang et al. [1.43].
More gentle cleaning methods (e.g., H2O2 method) should be considered in case we need
to preserve mesopores (i.e., fine pores with diameters ranging from 50 to 3 nm that present
on pore occlusions, which cover the areolae from inside or outside [1.48]) especially within
genera, such as raphid pennate diatoms (Figure 1.5b).
3. Preparation of permanent slides should be carried out for LM observations using a suit-
able high refractive index mounting medium such as Naphrax and Hyrax [1.44]. Figure 1.4
considers a good example of observing cleaned frustules fixed in a permanent slide under
LM.
4. Using LM to observe the 2D morphology of the cleaned valve and girdle bands to
assign the most similar genus and, if possible, the species. The overall morphology,
striae, valve symmetry, ribs, and the presence of raphe will be clear under LM; however,
the pores, the spines, and special structures within the valve and girdle band might
be not observable due to their smaller size. Though, in other few cases, the areolae
might be observable even before the frustule cleaning procedure, especially for large
taxa (Figure 1.1b). Valve outline, valve ends, shapes of sturdy, and hyaline silica (not
penetrated with striae or other openings) should be recorded. Classical diameter in
centric diatoms from valve view or length and frustule depth form girdle view allows
verifications of descriptions from literature. Moreover, the frustule silicification rate
can be observed under LM reflects information about the environmental condition that
surrounds the living cells.
5. Using SEM, after suitable sample preparation and coating with a conductive material
[1.38], to observe the ultrastructure of the valve and girdle bands and define the species
accordingly. Under SEM, the valve will have two different viewpoints, the internal and
external view. Some important criteria should be recorded using SEM including:
• the areolae structure from the internal and external valve view,
• striae (the periodic rows of areolae) shape,
• striae count in 10 microns or fibulae (in Nitzschioid diatoms [1.11], folds
in Surirelloid diatoms [1.18]),
• sternum size and shape (in Fragilaroid and Naviculoid pennate diatoms),
• nodule zone or annulus shape and its number (if more than one present),
• raphe structure and shape of raphe at the end and in the middle of a valve,
• the pore occlusions (if observable),
• the presence of spine or other projections like fultoportulae and labiate
processes,
• the shape and porosity of the girdle bands should also be recorded.
* For calculating the averages and ranges of the measured morphological data at least
10 specimens should be measured. Acid cleaned material allows observation of details but
often causes change or collapse of the frustule’s three-dimensional ultrastructure, thus the
selection of the most suitable cleaning procedure might be a crucial step to observe the dia-
tom ultrastructure properly. It worthy to be noted that the observation of the frustule parts
using SEM might be sufficient for genus and species identification without the need of LM;
however, some important descriptions for identification are based on the LM appearance
of the valve.
6. Observing the ultrastructure using TEM [1.38], if necessary and missing details did not
appear under SEM, such as the hymenate pore occlusions of raphid pennate (see, for exam-
ple, Figure 1.5b).
7. Study the nanotopography of the internal and external surface of the valve using AFM
[1.34]. Some features within the valve could be observed through SEM, or even optical
microscope if large enough, such as the frustule surface topology, which could be flat or
rather undulated, wrinkled, or containing groves; however, other nanotopographical fea-
tures cannot be revealed without a tool like AFM such as the dome shape of the cribellum
in [1.34].
8. To understand the relationships of frustule elements and the inner ultrastructure of a
given valve, or girdle bands, FIB-SEM should be considered (see example in Figure 1.6).
For more advanced understanding, one of the methods mentioned for 3D reconstruction
should be used [1.41] such as the 3D reconstruction of the 2D image series resulting from
FIB-SEM. Through these techniques, a clear understanding of, for example, the multilayer
nature of the valve could be obtained.
Afterward, all the ultrastructure details from inside and outside become well-known.
1.4 Conclusion
Diatom morphology could be an interesting topic for students; however, many challenges
in the field usually keep them away from getting a deeper understanding. This is a major
opportunity, since only 15,000–20,000 species have been identified [1.46], out of very large
number of extant and fossil species. Recently, the developed tools, especially AFM and
FIB-SEM, can make the complex 3D ultrastructure of diatoms more accessible; however,
very few attempts have been recorded in literature. For both the beginners and profession-
als, many unrevealed secrets of diatom morphology and morphogenesis are still kept hid-
den, which needs more progress in the field. The study of diatom morphology could also
inspire material scientists and engineers especially as the functions beyond morphology are
revealed. A comprehensive tutorial suitable for different specialists could be helpful to put
the beginners on the right track. Here, we only provided a potential introduction for such
tutorial.
Acknowledgements
We would like to express our gratitude to the editors for their understanding and help; we
are grateful for the meaningful suggestions of reviewers. KMM’s work on this project was
supported by funding from EPD, State of Georgia, USA. MG is very grateful to Dr. Mary
Ann Tiffany for giving him the micrographs in Figure 1.1.
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2
The Uncanny Symmetry of Some Diatoms and Not
of Others: A Multi-Scale Morphological
Characteristic and a Puzzle for Morphogenesis*
Janice L. Pappas1†, Mary Ann Tiffany2 and Richard Gordon3,4
1
Department of Mathematics, Michigan Math and Science Scholars, University of Michigan,
Ann Arbor, MI, United States
2
Dept. of Biology, San Diego State University, San Diego, CA, United States
3
Gulf Specimen Marine Laboratory & Aquarium, Panacea, FL, United States
4
C.S. Mott Center for Human Growth & Development, Department of Obstetrics & Gynecology,
Wayne State University, Detroit, MI, United States
Abstract
Organism symmetry via body plan is the balanced arrangement of structural features (internally
or externally) that is definable in terms of dimension (2D or 3D) and geometry (contours, sur-
faces, boundaries). Symmetry is evident at the endpoint of morphogenesis and at each scale, from
genes to cells to organism to population or higher taxonomic level. Symmetry may change from
juvenile to adult stages as well as between sexual and asexual reproduction. The arrangement of
morphological features reiterates symmetry antecedents so that at any given time, degrees of sym-
metry may be directed, random or chaotic outcomes signifying degrees of instability throughout
morphogenesis.
With their geometric surface ornamentation, diatoms exhibit a high degree of rotational and/or
reflective symmetry depending on the ontogenetic path, either commencing with an annulus or a
sternum for centrics and pennates, respectively. Sometimes this symmetry is remarkable, whence
we call it “uncanny symmetry”. Using centric diatom examples, surface features are used in uncanny
rotational symmetry assessment via digital image processing techniques and include analysis of veg-
etative valve formation stage, initial valves, and abnormally developed cells. Quantitatively, entropy
as digital image information content is related to symmetry states and is used to determine random
and chaotic instability in morphogenesis.
Keywords: Rotational symmetry, reflective symmetry, centric diatoms, diatom morphogenesis,
fluctuating asymmetry, uncanny symmetry, entropy, developmental instability
*Dedicated to the memory of the late Antone G. Jacobson and his life pursuing the puzzle of morphogenesis.
†
Corresponding author: jlpappas@umich.edu
Vadim Annenkov, Joseph Seckback and Richard Gordon (eds.) Diatom Morphogenesis, (19–68) © 2022 Scrivener
Publishing LLC
19
2.1 Introduction
Morphology is a conduit to assessing evolutionary relationships as well as ecological inter-
actions among organisms. Different aspects of morphology provide information on size,
shape, color, pattern of the external form of an organism, as well as internal anatomical fea-
tures. Morphology is informative regarding commonalities among organisms at the species
level in population studies and among larger taxonomic groupings from generic to phylum
level in macroevolutionary studies. At the phylum level, body plan (bauplan) assessment
enables morphological analysis across taxonomic levels, and characteristics of phylogenetic
importance can be gleaned as comparative markers of continuity in arrangement of struc-
tural form.
A body plan is the group of structural characteristics of a phylum that is shared via
development [2.41]. Genetic and embryogenetic processes manifest themselves through-
out development, culminating in the adult form as the end member of morphogenesis.
The connection among morphogenetic forms throughout development is exemplified via
symmetry and symmetry breaking [2.3, 2.19, 2.24]. It is symmetry that enables assessment
of body plans not only among developmental stages within a given phylum but also among
phyla exhibiting commonality in some aspect of structural form.
Symmetry as an external morphological characteristic is determined via perception
or observation of the “balance” of an organism. That is, we perceive or observe whether
the organism exhibits some evenly distributed “likeness” of form, albeit as a reflective
[2.159] or rotational [2.159] impression. To determine the “balance” or “likeness” of a
given form, we necessarily perceive or observe the geometry of the organism explic-
itly or implicitly as an impression. To arrive at the point of recognition of the form as
an identifiable entity, we perceive such characteristics as degree of curvedness (round-
ness) or angularity (sharpness) as well as distinct surface peaks, valleys, or saddles (local
maxima or minima as extrema), if present. Surface geometry plays a key role in our
perception of the quality of form [2.104–2.108]. Contours and edges are the pieces of
information we gather and connect to the extrema or points (dots) that enable us to con-
coct a contiguous picture or image about the form we perceive or observe. This provides
the initial point of determination in the assessment of symmetry of a given organism.
The boundary shape of an organism as a composite of contours is instrumental in sym-
metry determination.
The product of morphogenesis culminates in a fixed or adult shape in which the organ-
ism will exhibit specific external and internal symmetries, usually having symmetries
common to a higher taxonomic group. Structural aspects of morphology such as sym-
metry are evident from the morphogenetic process of each succeeding step in develop-
ment. Arrangement of morphological features reiterate internal and external symmetry
antecedents so that to some extent, symmetry is a directed process with regard to mor-
phogenesis [2.54]. Symmetry is not only evident at the endpoint of morphogenesis but
also at each scale of the materials involved, from genes to organelles to cells to tissues and
beyond, in the morphogenetic process. That is, symmetry is hierarchical and scale depen-
dent. Changes from juvenile to adult stages as well as differences between sexual and
asexual reproduction will exhibit symmetry characteristics at each stage of development.
Diatoms’ Uncanny Symmetry 21
Matching symmetry states to morphogenetic stages may inform the developmental

process.
Symmetry is, then, a compilation of perception or observation as well as the arrange-
ment of structural features (internally or externally) defining dimension (3D) and geometry
(contours, surfaces, and boundaries) in terms of “balance” and “likeness”, and this compila-
tion can be evaluated at any step in the morphogenetic and developmental processes of an
organism at the individual, population, or higher taxonomic grouping level.
2.1.1 Recognition and Symmetry

Object recognition is the term applied to our understanding of how to perceive or observe
a three-dimensional (3D) object and interpret it from its two-dimensional (2D) projection.
Perception studies with Attneave’s cat [2.9] or Biederman’s cup [2.13] as iconic images to
discern the process of recognition utilize the geometric aspect of objects as points connected
with straight edges or partial contours, respectively (cf. visual tracking of eye movements
[2.4]). These studies showed that recognition is initiated from perception of outlines rather
than just points on an object. Information acquisition is accomplished via this perception
or observation. One perspective is the preference for the simplest interpretation over others
to infer a structured whole, and this forms the basis of structural information theory [2.80].
Another perspective involves preference for a minimum (not necessarily simple) descrip-
tion forming the basis of algorithmic information theory [2.21]. In any case, information
theory is at the heart of object recognition. Perceived or observed information may occur
as an object that is recognized. Initial recognition may occur of a whole scene as an object.
In contrast, individual objects in the scene and their boundary and surface attributes such
as shape, transparency, pattern, location, size, or texture may be perceived, and perception
violations of individual objects in a scene may be recognized (Figure 2.1, after Mezzanotte’s
scenes, in [2.14]). Symmetry as a mode of recognition can apply to the scene as an object or
the parts of this object in relation to the boundary and surface attributes that comprise the
scene. Information about symmetry is acquired with respect to an external or anatomical
viewpoint.
Body shape is important in symmetry determination of organisms. Shape deformation
relies on a point-based approach with regard to object recognition requiring identifiable
equivalences on multiple shapes. The shape decomposition approach represents an implicit
reference to contours in recognition as the conduit to assessing symmetry [2.127] (Figures
2.2a–c). Shape decomposition relies on the delamination of surface boundary height via
contours with respect to object recognition and implicitly influences shape in symmetry
assessment [2.72] (Figures 2.2d–f).
For reflective (also called bilateral, mirror, mirror-image, line, plane, or left/right) sym-
metry [2.159], shape is considered to be the most ubiquitous among organisms and is espe-
cially noticeable in animals and plants. Colored symmetry [2.132] can also occur, as in the
left/right asymmetry of otherwise bilaterally symmetric organisms [2.41, 2.144]. However,
symmetry is present as more than rotational or reflective in organisms (Figure 2.3).
In 2D, dihedral (rotational plus reflective) [Weisstein 2002], translational, and glide (reflec-
tive plus translational) [2.159] symmetries are evident. The 2D and 3D quality of whole
organisms or their parts lends itself to the related symmetries of knots (including chirality)
Figure 2.1 A drawing composed by JLP that is reminiscent of one of Mezzanotte’s scenes [2.15]. Although
the scene is the whole perceived object, individual objects diatom (fish, jelly fish, boat, oars, waves, and shore)
and their attributes such as shape, transparency, location, size, and texture and perspective of the scene may be
viewed as perception violations.
(a) (b) (c)
(d) (e) (f)
Figure 2.2 Shape perception of Arachnoidiscus ehrenbergii from image ProvBay5_12lx450. (a–c) Shape
deformation via an increasing number of points defining valve features. (d–f) Shape decomposition via an
increasing number of contours defining valve features.
[2.154], helices/spirals (including handedness) [2.159], and compound or multiple spi-

rals (a variant of knot symmetry), [2.159] as well as scale (fractal or dilation) [2.159] and
related conformal symmetry [2.159] with associated Möbius transformations [2.159].
Symmetries associated with loop networks [2.36, 2.88, 2.89, 2.159, 2.164] are present at
various scales in organisms as well. See Figure 2.3 for examples of all symmetries on the
diatom valve surface. Note that 1) rotational and reflective symmetries apply to the whole
valve; 2) dihedral, translational, and glide symmetries apply to pores on the valve face;
3) knot, helical, compound spiral, and conformal symmetries apply to valve central areas;
4) scale symmetry applies to the replication of five-fold symmetry within a Triceratium
valve face; 5) loop networks are seen in the pores on the valve surface of the Trigonium and
Triceratium valves depicted.
Symmetry has implications for evolutionary processes in animals and plants [2.54].
Symmetry may be adaptive so that active organisms or those in rapidly flowing water [2.45]
will have reflective symmetry while sedentary organisms in low flow shear regimes will
have rotational or dihedral symmetry. Organisms with large surface to volume ratios or
repetitive parts tend to have translational symmetry (Figure 2.4). Organisms that exhibit
energy efficiency have scale symmetry [2.158]. In general, symmetry is a dynamical prop-
erty of organisms.
(a) (b) (c) (d) (e)
(f) (g) (h) (i) (j)
(k) (l) (m) (n)
Figure 2.3 Diatom valve face symmetries. (a) Arachnoidiscus ehrenbergii-rotational; (b) Biddulphia sp.-
reflective; (c) Coscinodiscus sp. microstructure-translational; (d) Asterolampra marylandica-dihedral; (e)
Coscinodiscus sp. microstructure-glide; (f) Trigonium americanum-helical/spiral central area; (g) abnormal
Cyclotella meneghiniana-multiple helices/spirals central area; (h) Coscinodiscus sp.-helical/spiral; (i)
Actinoptychus splendens-knot (interlooping valve structure); (j) Arachnoidiscus ornatus-knot (interlocking
rings under valve ribs); (k) Triceratium pentacrinus fo. quadrata-scale; (l) initial valve of Biddulphia sp. 3D
valve surface-conformal symmetry or Möbius transformation; (m) Trigonium dubium-loop network; (n)
Triceratium bicorne-loop network. (All SEMs by Mary Ann Tiffany). See “Recognition and symmetry” section
in the text.
Figure 2.4 Double translational symmetry at two scales exhibited in a Paralia sulcata chain colony of
repetitive cells. Light micrograph by Mary Ann Tiffany.
2.1.2 Symmetry and Growth

Morphogenesis has been characterized as a system of detached movement of ensembles of
cells in animals in contrast to lack of detached movement via ensembles of cells in plants.
That is, locomotion is one way to associate a specific kind of symmetry to motile organisms
in contrast to sessile organisms which may exhibit another kind of symmetry [2.59]. Modes
of growth contribute to the recognition of specific types of symmetry. Branching growth
exhibits continuous symmetry decrease and increase in size during growth [2.47]. Stepwise
growth and size reduction are modes of discrete growth where size increases or decreases
over time. Growth changes are concomitant with symmetry changes as potential indicators
of stress associated with development, morphogenesis, cytogenesis, epigenesis [2.154], and
embryogenesis [2.43, 2.149], depending on evolutionary or ecological conditions. Levels
of environmental or teratological stress are said to influence fitness and adaptability of an
organism, although this connection has not been quantitatively or definitively formalized
[2.53, 2.54, 2.35, 2.68].
Developmental instability has been associated with fluctuating asymmetry which is gen-
erally considered as small random deviations from perfect reflective (bilateral) symmetry
[2.7, 2.53, 2.60, 2.87, 2.94, 2.128, 2.152]. Instability is presumed to be associated with either
side of an organism and developmental processes, and genetics or environmental condi-
tions may influence the size of the fluctuations in asymmetry [2.53]. No explicit or quantita-
tive relation has been given connecting fluctuating asymmetry with instability as a measure
of stress. Assessment using fluctuating asymmetry characterizations have been made on a
population (species) level and has not been carried out with individuals or at interspecific
or higher taxonomic levels [2.53, 2.54].
2.1.3 Diatom Pattern Formation, Growth, and Symmetry

Diatoms are unicellular microalgae that are known for their exquisitely ornamented
hydrated silica frustule (cell wall) with geometric intricacies. At a molecular level, orna-
mentation on the valve surface emanates from silicic acid precipitation [2.47] and interac-
tion with proteins in the silica deposition vesicle (SDV) and silica deposition through the
SDV membrane (known as the silicalemma) [2.50, 2.142]. Valve surface patterning occurs
as a diversity of geometric structures [2.98]: the multitude of species-specific forms with
various symmetries. Anatomically, a diatom frustule is composed of two close-fitting thecae
(valves) subtended by cingulae (girdle bands) [2.119, 2.150, 2.151]. During reproduction,
the cell undergoes expansion as two daughter cells are created, with each daughter half-cell
acquiring each half of the mother cell and one half (valve) of the frustule [2.119]. Each new
valve is constructed within an SDV within each daughter cell. Because one valve fits inside
the other (like petri dishes) and each daughter cell builds its new valve within the perimeter
of the valve it inherited, one daughter cell is usually smaller in size than the mother cell
(but cf. [2.117]), and the size diminution process continues until a developmental limit is
reached according to the MacDonald-Pfitzer rule [2.90, 2.111, 2.112], which may correlate
with a minimal organelle content [2.97]. At this point, size is restored via sexual reproduc-
tion and auxosporulation and from this, an initial cell is produced which in turn produces
the vegetative cell [2.119].
Diatom symmetry formation is closely tied to pattern formation. Models of pattern
formation have been developed that consider diffusion limited precipitation in non-
equilibrium conditions where silica diffuses within the thin SDV depositing silica initially on
the midring in centric diatoms or the midrib in pennate diatoms [2.47, 2.109]. This process
has been observed to take about 10 minutes and is followed by thickening of the valve sur-
face, which occupies an additional 6 hours [2.47]. Alternatively, pattern formation has been
modeled as phase separation of two hypothesized liquids [2.82, 2.138] so that the repetitive
honeycomb and pore patterns formed in some diatoms occur because of the pH regime and
molecular reaction during silica deposition [2.76]. The third phase, the actual solid precipi-
tated silica, is not included in this model [2.82]. In an alternate computer simulation, albeit
confined to pore occlusions, a single fluid is involved (cf. concentrating “mother liquor”
[2.47]), with solid silica particles free to “redissolve” after precipitation [2.161, 2.162]. This
is similar to a simulation of diatom costae that permitted surface diffusion of silica particles
along the precipitate surface, after they adhered to it, to approximate sintering [2.47]. The
balance between precipitation and sintering has been simulated, using the language of “diffu-
sion limited” versus “reaction limited” [2.25]. The main difference is whether silica particles
continued to move in the medium [2.25] or along the surface of the precipitate [2.47]. Pore
occlusion simulations [2.25] parallel simulation of precipitation with a centric diatom’s mid-
ring [2.47]. Spine formation has been modelled as “a combination of membrane and cyto-
skeletal activities creating a mold for silica deposition” [2.11], where the silica presumably
sinters. A model has been proposed that regards the SDV, in conjunction with the micro-
filament ring at its perimeter [2.50], as a tensegrity structure that may explain the shape
of polygonal centric diatoms [2.48]. All of these models presume that the SDV membrane
constrains the silica precipitation, the latter at best somehow enlarging the SDV, by unstated
mechanisms. However, the opposite effect, of the precipitated silica altering the shape of the
SDV, not just its size, can also be conceived.
Pattern formation diatom studies on cellular organelles during mitotic stages of ontog-
eny preceding valve formation indicate changes in organelle location and symmetry (cell
organization) at the subcellular level as well [2.114, 2.123, 2.155]. For example, during inter-
phase the arrangement of vacuoles on either side of the central area with respect to micro-
tubules and the microtubule organizing center [2.113, 2.122] and spindle formation via the
position of the microtubules are indicative of reflective symmetry. As cytokinesis proceeds,
at metaphase chromosomes assemble around the spindle during metaphase, while at the
end of anaphase complete cleavage of the spindle occurs. The position and cleavage of the
spindle exhibit reflective symmetry just prior to the formation of new daughter cells [2.28]
(cf. [2.1]).
Each of the modeling systems proposed for diatom valve formation and morphogenesis
are incomplete in that they either have not been found to be present during all parts of the
process or indirectly attempt to account for morphogenetic processes from a chemical/
molecular or cytogenetic viewpoint. As has been emphasized, each “new model of diatom
morphogenesis”, which, although highly attractive, is a model, awaiting to be substantiated
by TEM-micrographs of sufficiently prepared cells, and experiments undertaken with the
living cell” [2.124]. This remains the case. Experiments such as manipulation of salinity
have morphogenetic effects, which can be interpreted at the level of silica precipitation
[2.157], and the presence of (probably) sodium chloride crystals inside the silicalemma
[2.46] is consistent with easy access of small ions into the SDV. The same goes for pH [2.58].
Gene regulation of silica transporters may occur at multiple levels [2.120], perhaps corre-
sponding to multiscalar effects on morphogenesis.
In any case, no model so far accurately simulates the mature silica structure at all scales.
This is not surprising, since multiscalar diatom silica structures span an extraordinary 8
orders of magnitude [2.37], and it would seem unlikely that the same mechanism prevails
at all size scales (although nested hexagons made of silica spheres of decreasing sizes have
been postulated, but not computer simulated, to account for three levels of structure in
Coscinodiscus [2.138, 2.139]. The formation of these nanospheres has also been simulated
[2.81]. A seven-state cellular automaton model for pennate diatom morphogenesis sets the
striae and pores at specific spacings, by unspecified mechanisms [2.12]. At a lower scale,
organic microrings of diameter 6 nm may correspond to fultoportulae [2.75]. By looking
at each of the models as potential partial mechanisms for producing different aspects of
symmetry during different morphogenetic stages and scales, the models may be implic-
itly useful in addressing how symmetry becomes evident during morphogenesis and how
symmetry may be instrumental in understanding stability changes during morphogenesis.
However, we presently have no global models that work at all scales. “Better models are
needed, therefore, but for this we need a better formulation of the problems we are trying
to solve” [2.93].
None of these models for diatom pattern formation involve the linked chemical reactions
of Turing reaction-diffusion processes [2.148] which may lead to instabilities that have been
invoked for multicellular morphogenesis [2.91, 2.92, 2.99]. However, Turing models have
recently been simulated for multilayer situations [2.153], producing patterns somewhat
similar to arrays of diatom pores so that such models are still under consideration [2.43].
In a dynamical system, changes in structural stability may be related to changes in
symmetry via pattern formation changes not only over time but also at hierarchical spa-
tial scales [2.166]. Pattern formation as an indicator of symmetry states may be evident as
self-similarity as well [2.96]. The role of symmetry in developmental processes is discern-

able through surface pattern and shape changes over time, and these changes may be used
in a morphogenetic dynamical system to explicitly assess the contribution of symmetry to
structural stability.
In our work on morphogenesis with the late Antone G. Jacobson, we conceived of and
repeatedly traversed an intellectual triangular loop of 1) experiments and observations, 2)
computer simulation, and 3) formal mathematics, to converge to an understanding of how
nature did it [2.49, 2.61–2.64]. This loop has not been traversed or ever closed even once
for diatoms. Curiously, in the research with Jacobson, we invoked the bilateral symmetry of
the organism we were simulating, which both reduced computer time and made the sim-
ulation match reality much better. A similar two axis symmetry was placed on a pennate
diatom simulation [2.12]. However, in a real organism there is no global “force” creating its
symmetry. Symmetry is a consequence of morphogenesis, not a cause, and deviations from
symmetry or its perfection are hints to the underlying processes.
2.1.4 Diatoms and Uncanny Symmetry

Microorganisms are generally not well studied in terms of quantified symmetry changes
over time. Diatoms have distinct amorphous silica frustules that exhibit a variety of geo-
metric shapes and surfaces that lend themselves to analyses of symmetries. Diatoms are
pigmented protists that are considered to be a monophyletic phylum. Morphogenesis is
a topic of great interest not only to phycologists but also to nanotechnologists [2.42, 2.44,
2.51, 2.85]. In girdle view, diatoms are asymmetrical because of the parent-daughter cell
division that occurs within the previously formed cell. In valve view, shape and surface are
both open to symmetry considerations.
We have proposed that diatoms possess “uncanny symmetry” [2.134]. By uncanny sym-
metry is meant that (some) diatoms in valve view (may) exhibit near perfect symmetry. It
was shown that by subtracting the rotated image of a given diatom from its original image,
an almost completely blacked out image would result, indicating near perfect matching
of image pixels, suggesting near perfect symmetry [2.134]. Such results were obtained for
Aulacodiscus oregonus and Triceratium formosum var. quinquelobata (Plate 7, Figures 1–4
in [2.134]). “The apparently high degree of perfection of this noncrystalline precipitate
deserves quantification, which may prove comparable with the (small) degree of imperfec-
tion of crystalline snowflakes [2.83]” [2.134].
Centric diatoms exhibit rotational symmetry in shape and surface, but other symme-
tries such as dihedral symmetry are present as well. A case in point is Auliscus (Plate 5,
Figure 1 in [2.134]). From dihedral symmetry, reflective symmetry of this diatom may
be characterized as a 180° rotational symmetry, thereby identifying rotational symmetry
as a starting point in the measurement of uncanny symmetry. That multiple symme-
tries occur simultaneously in centric diatom may be recovered by an uncanny symmetry
measurement.
As suggested in [2.134], we propose to quantify centric diatom symmetry generally and
uncanny symmetry specifically using concepts from information theory and image pro-
cessing methods. Because symmetry involves the acquisition of information regarding the
“balance” of an organism such as a diatom, information theory is used to develop a measure
of centric diatom valve uncanny symmetry.
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hearted fellow, and does not scorn old friends. Many’s the nod and
word he has given me from the billiard-room window at Sidney, when
I have been passing, let who would be there. Everybody is glad of
the prosperity of John Lawe, Esq.”
Somebody having made inquiry about the voyage, Green went on,
“O, that’s the worst part of it. It’s horrid enough, to be sure, to be
cooped up for months on board, and all so solemn and dull, and no
getting out of the way of the clergyman. But it’s not so bad as it used
to be, when they treated such folks as we like so many wild beasts.
They paid the captain so much a head for the people embarked, and
never asked how many he landed; so he starved as many as he
pleased, and stowed them so close that scores were stifled by the
way. It was mighty dull work then for those that got safe; the labour
was so hard, and no liberty. There was little encouragement to go to
the colonies then. But now that they don’t kill one by inches by the
way, it is worth putting up with the passage, for the chance of making
one’s fortune at the end of it.”
“Particularly for them that have friends in power to get fine
situations for them,” said Jerry pertly. “Bob and I are going to have
good care taken of us, I hear. But it’s a great plague that the old
ones are going to be spies over us. It will spoil our sport terribly,
unless we can manage to cut them.”
“That’s better than having them whining and praying after us all
the way from here to the ship, as the old folks mostly do,” said
Green. “When I went before, my father behaved as if he was
following me to the gallows. He knows better now. He gave me the
wink yesterday for a sharp chap that knew how to take care of
myself. He said,—true enough,—that the worst blunder I ever made
was coming back when once I was well off.”
“Aye, aye, Green; a certain person knew how to take care of
herself as well as you. She knew better than to keep herself single
five years for you. ’Tis a fine feather in her cap to have brought you
so far on a fool’s errand.”
Green tried to conceal his visible passion under an appearance of
indifference, while he muttered that a better one than he came for
would follow him out very soon, if the judge did not baulk them of the
sentence they meant to get pronounced upon her.
“Here they come, lads!” he cried, interrupting himself. “All is ready:
our carriage at the door! Put a bold face upon it, boys! Now for it!
Don’t have anything to say to the whiners at the gate. Curse all spoil-
sports! Give them three cheers, boys! Hurra! hurra! hurra!”
And gibing, jeering, laughing, shouting, went the batch of convicts
through a throng of relatives and former companions, and gazing
strangers; some of whom were pale and weeping, others signing and
winking, and more gaping in wonder and pleasure at the scene;
speculating upon whether the largest share of punishment did not
rest with those who were left behind. Bob, and one or two other
scowlers, were almost overlooked in the company of adventurers,
who seemed to be going forth merrily to cheat the law, and seek their
fortunes in a land of plenty.
Chapter IV.
NEW HOMES.
Ellen was the first of the family that arrived at Hobart Town in Van
Diemen’s Land. Next came the convict-ship, which was sent round to
Launceston to disembark its passengers; that port being nearer the
district where the convict labour was to be employed. When the
batch of parish emigrants arrived, a fortnight afterwards, Frank
found, on application to the proper government officer, that his sister
had landed in good health, and had received a high character from
the clergyman and his lady who had come over as superintendents
of the expedition; that Ellen had been forwarded, with a few of her
fellow-passengers, to the district where a service had been procured
for her as dairy-maid on a settler’s farm; and that care had been
taken that her parents and brother should be indentured to farmers
in the same neighbourhood. So far, all was well. Frank could learn
nothing about his brothers, except that they were to be landed at
Launceston, and that Launceston was within thirty miles of the spot
where he was to be located. The officer he was speaking to had
nothing to do with the arrangements respecting convicts: his
business was to take care of emigrant labourers on their arrival.
Castle himself could not help being pleased at the appearance of
things at Hobart Town, when he and Frank took a walk, the evening
after their arrival. The only objections he could think of were, that
there were few shops; that it was not at all likely that the country
inland should be half so civilized as what he saw; and that it was a
thing he had not been used to, to have Christmas fall at the hottest
time of the year, and the trees green all the winter through. It was
now May; and they told him that winter was coming on, and yet that
the woods would look as green as now all the time; and that the
snow, if there was any, would not lie more than a day on any ground
but the mountain tops, and a bleak common here and there. They
told him that for more than three hundred days in the year the sun
would shine all day, and the air be dry and pure, and seldom too hot
or too cold. All this was what he had not been used to, and did not
know how to believe. His son supposed that if it came true, he would
not object; as one of the consequences of such a climate is that
English people have much better health, and live, on the average, a
good deal longer at Van Diemen’s Land than at home. Castle
peevishly laughed at all talk about life and health, when it was, in his
opinion, doubtful whether they might not be starved to death within
three months. His son left this point to be demonstrated by time
rather than by argument; and meanwhile observed that there were
few signs of starvation about Hobart Town, in which, besides the
government residence, there are nearly eight hundred houses, most
of which are surrounded with gardens; the dwellings having been
originally built on separate allotments of land, of a quarter of an acre
each. The streets cross at right-angles, and command fine views of
the neighbouring country, and afford cheering evidences of the
success of the industry which has sought employment there. A dock-
yard is seen on the river’s brink; and corn-mills, tanneries, breweries,
a hat-manufactory, &c., are conspicuous in the midst of the town. An
amphitheatre of green hills rises to the westward, the crowning
summit of which is 4000 feet high; and from these hills descends a
fine stream of water, flowing through the town into the Derwent,
which, with its varying expanse and beautifully wooded bays and
sloping shores, forms the eastern boundary. This view was little
enough like what Castle had fancied in opposition to all that he had
been told. He was for ever picturing to himself a region of wild
woods, or bleak plains covered with snow; and he was now as much
surprised at the sight of meadows, hills, dales, and a thriving town,
with a blue sky overhead, as if he might not have known as much
before. He had complained of his hard lot in being indentured as a
shepherd; and no wonder, while he thought his flocks were to inhabit
a dreary wilderness; but now that he found he had nothing to fear
from storms and snow-drifts, that the pastures were excellent, the
springs plentiful, and the sheep as fine as the world can produce, he
began to think he might be worse off in point of occupation; though
he would give nobody the satisfaction of hearing him say so. His wife
was to be a domestic servant in the same farm where he was
shepherd; and even little Susan was carefully stipulated for; the
labour of children being valuable at almost any age, in a place where
much more assistance is wanted than can be had.
The first part of their journey to the Dairy Plains, (the district where
they were to settle,) was through the very choicest portion of the
island, both as to beauty and fertility. It is not surprising that those
who first surveyed this tract, and took it as a fair sample of the island
at large, should have represented Van Diemen’s Land as a terrestrial
paradise, and been suspected of exaggeration through the
favourableness of their report. The fact is, the island is supposed to
contain about 15,000,000 acres,—one-third of which is considered
arable, another third fit for sheep-pasture, and the rest unprofitable
at present. The country between Hobart Town and Launceston
consists of green hills and fertile plains, among which towns and
villages and solitary dwellings are interspersed. Rivers wind between
their wooded banks, and streams flow down from the high grounds.
Excellent macadamized roads run through the whole district, and
branch off to the growing settlements on either hand of the main
track. It was a great amusement to Frank to compare whatever he
met with that partook of the civilization of his own country with
whatever looked new and strange. Before leaving Hobart Town, he
had been all the more struck with its printing establishments, its
Mechanics’ Institute, its Book Society, and schools, from observing
the strangeness of the natural productions that he met at every step.
In the gardens he beheld tea trees where he had been accustomed
to see lilacs and laburnums; and cotton plants, myrtles, and
geraniums growing as tall as himself, and spreading out into bushes.
The very grass grows differently;—not stringy in the roots and
carpetlike in the surface, as in England; but in tufts. Parrots, instead
of canaries, were the pets of young ladies; and the bandicoot was
offered for sale instead of the rabbit. Cockatoos instead of crows
were to be frightened away from the fields and gardens; and flocks
of pigeons among the stubble looked as much like partridges as
pigeons; only more beautiful,—with their gold-dropped wings,—than
either species in England. On the road, in like manner, the freestone
bridge over the Jordan, the postman on horseback, the tillage and
inclosures, looked British; but the evergreen woods, in the midst of
which arose the peppermint tree to a lofty height;—the herds of
kangaroos coming out of their covert into the dewy plains at sunrise;
—the spotted opossums climbing and descending the forest trees;—
the black swans sailing on the lakes, and uttering cries like the
creaking of an old sign-board;—all appeared foreign, and scarcely
belonging to the people who had settled among them.
A sight of a yet different character met the eyes of the travellers
near the close of the second day, when they were drawing near their
future abode in the province called Norfolk Plains, in the centre of
which lay the Dairy Plains, where Ellen was expecting them. They
had for some time quitted the broad road, and were following a track
along which their waggon proceeded with tolerable convenience. At
last they came to a point beyond which it had not been carried, and
where a gang of labourers was at work roadmaking;—not as in
England, each man intent upon his own heap of stones, free in limb
and thoughtful in countenance;—not as in Ireland, where some are
lounging and all are joking;—but charged with the fetters of felons,
and superintended by an armed taskmaster. As Frank looked upon
these wretches, with their hardened or woful countenances, he felt
indeed that he was not in England, but in one of her penal
settlements,—breathing the air of one of the places where her vice
and misery are deposited. His very soul became sick when, as the
labourers turned to stare at the somewhat uncommon sight of a
waggon full of travellers, he met the eyes of his convict brothers. He
hoped that his companions would not perceive them; but he soon
found that his father did, by his testy complaints of the jolting of the
cart, of cold and heat, and what not. The unhappy mother looked on
her outcast children with as much curiosity as compassion. Bob
turned away, and stooped to his work, never looking up till they were
out of sight; but Jerry waved his cap and shouted, and dared Frank
to a wager which of them would first be free to work for themselves:
whether it would take longest to work out his sentence, or to pay for
Frank’s passage and settlement. This supplied a new theme of
complaint to Castle, who wrought himself up into a passion about his
being a slave, with all his family. Frank, who hated bondage as much
as any man, thought it could hardly be called slavery to contract to
work for one person for a certain time, in return for advantages which
could not otherwise be obtained. If disappointed of these
advantages,—of sufficient food, clothing, shelter, and money wages,
—the contract was void, and no harm done; if not disappointed, the
object was gained. The evil lay, not in their case as labourers; but as
honest men. Felons ought not to be let off so easily, (because their
labour happened to be more valuable than at home,) as to make
disgrace, for which many of them did not care, their only punishment;
their worldly circumstances being actually bettered by their removal
to a new colony. It was not that labourers need be better off than
their family would probably be, four or five years hence; but that
felons ought not to be placed in as good circumstances as the
honest emigrant at the end of the same period.
Frank was not yet aware (as he afterwards became) that, for want
of knowing the rate of wages at the colonies, emigrants often bind
themselves for a much lower rate than they might obtain if they went
free, or if they were properly informed of the existing state of things;
and thus think themselves deceived, and are tempted to break their
contract when they find how matters stand. This evil is to be referred
to the ignorance of emigrant labourers, quite as much as to the close
economy of the settlers, and should induce all who have heard of it
to obtain such information before concluding their bargain as will
save them from repentance afterwards, and guard them against
quarrels on this score with their new masters;—quarrels, which,
always a great evil, are most so in newly settled countries, where all
hands and hearts are wanted to work together for the common good.
As it is, a British artisan jumps at the offer of a plentiful subsistence
and 2s. a-day besides for five years, out of which the expenses of
his removal are to be paid; and for this rate he binds himself. When
he gets to his destination, he finds that this plentiful subsistence,
including meat, bread, beer or spirits, tea, sugar, comfortable
clothing, and a convenient dwelling, costs no more than 2s. a-day,
and that, if free, he might earn, being a good workman, from 7s. to
12s. a-day, or even 15s., if he be a superior mechanic of a scarce
class. It is mortifying to find that he has sold himself, however much
higher than formerly, for less than he is worth in his new position;
and hence arise discontents which embitter the first years of his new
life, if they do not occasion a breach of contract. The friends of a
rational plan of emigration should do their utmost to make known to
as many as it may concern, to what extent labour is wanted in the
colonies,—what is the rate of money wages in each, and what those
money wages will procure. The official information on these points
transmitted from Van Diemen’s Land in 1827, was, that common
labourers earn 3s. per day; common mechanics 7s.; better
mechanics, from 8s. to 12s.; best ditto, from 12s. to 15s.; and
persons of peculiar qualifications, fitted to superintend farms or other
undertakings, 1l. a-day. Since that time, wages are understood to
have risen. The price of wheat is 7s. a bushel; meat, 2d. or 3d. per
lb.; sugar, from 3d. to 6d.; and tea, from 1s. 6d. to 4s. per lb.—No
wonder that, amidst all their gratitude at being well provided for,
many such workmen as Frank are vexed and mortified to find how
much more they might have made of their labour.
Far other feelings, however, than those of discontent were
awakened in Frank by the aspect of his new abode. It was almost in
a state of nature, his employer, Mr. Stapleton, having preceded him
to take possession only a few days before: but it was far from being
a desolate spot in the midst of a waste, as settlers’ farms are wont to
be in colonies where the unwise object is to disperse the inhabitants,
instead of bringing them near to enjoy the advantages of a division of
labour and reciprocity of consumption. The Dutch government at the
Cape of Good Hope formerly forbade settlers to approach within
three miles of each other; and thus effectually prevented the full
improvement of the land, the construction of roads, and the opening
of a market for exchanges. Hence the Dutch settlers at the Cape are
to this day deprived of many advantages of civilized life. They have
too much of whatever they grow, and too little of what they would fain
buy; and are debarred the comforts of society and mutual help.
These evils are likely to be avoided by the method of disposing of
land now adopted by our government in Australia; the land being
sold on terms which make it the interest of the settler to improve his
tract, and to take advantage of a neighbourhood which may relieve
him of some of his produce. Mr. Stapleton, having been obliged to
choose his land carefully, and to pay 9s. an acre for it, (instead of
6d., or nothing at all, like some of the earlier inhabitants,) was not
tempted to wander away into the wilderness, and sit down where he
might happen to like the prospect, or to be smitten with some new
discovery of fish-ponds, woods, and meadows. He made his choice
instead among the lands of a certain district; and selected such, as
to extent and quality, as would on the whole best suit his purposes,
in conjunction with the privileges of a neighbourhood. His land,
though not of the very first quality, was good enough to have fetched
15s. per acre, if it had lain somewhat more to the north or east,
where the country was rapidly becoming better peopled; but it
stretched towards the unoccupied districts at the foot of the western
mountains, and was less valuable than if it had been surrounded by
civilization, instead of only bordering upon it.—It consisted,—not of
jungle and forest ground, where room must be made by the axe
before seed could be sown and sunshine visit it; but of a lightly
timbered and undulating surface of grass land, wanting only a single
burning to render it fit for the plough, or for a new growth of pasture.
The trees were not of the nature of copse and thicket; but growing in
clumps a hundred feet apart, and with clear stems, measuring ninety
feet or more to the lowest branch; thus affording spots for shade and
shelter without interfering with tillage. The boundaries, where not
formed by natural streams, were fixed by marking the trees; and
there was no immediate need of fences where neither man nor beast
was likely to trespass, and where there was at present no live stock
that could be in danger of straying. No one was near who could be
tempted to steal; for none were poor. The wild cattle, which in former
days did great mischief on the grounds of the settler, had long ago
been driven among the mountains, where it was supposed the race
had died out, as none now appeared. The few oxen and horses that
Stapleton brought with him were kept near the dwelling; and the rest
of the stock was not to follow till all was in readiness for its reception.
A rude shed had been hastily constructed for shelter, under a clump
of trees; and a few sawn planks were lying about: by which Frank
saw that the materials of his business were ready for him to begin
upon without delay. Tools and utensils were stowed away in corners,
or heaped under the trees, till their proper places were provided for
them; and a goodly row of casks and packages of provision stood in
the background. Frank had believed that his spirit of enterprise had
died within him under the hardships of his own country; but he now
felt it revive in a moment; and was anything but dismayed at the
prospect of what he had to do in his capacity of carpenter, before the
scene before him could put on the appearance of a snug and well-
managed farmstead. He saw in fancy the day when a little hamlet of
weather-boarded cottages would be sending up their blue smoke
among those trees; when cattle-sheds and sheep-pens would stretch
out behind the dwellings, and the busy forge and creaking timber-
wain would drown the cry of the quail, and scare away the
kangaroos that were now leaping over the plains. He did not forget to
add a very superior workshop and timber-yard to his picture of his
own dwelling; or to imagine his father looking down from among his
flock on the hills, or Ellen within sight, going forth in the bright early
morning with her milk-pail.
As if to answer to his thought, Ellen now appeared. She had stolen
half an hour to run in search of Mr. Stapleton, to ask once more how
soon he thought Frank might possibly arrive. Mr. Stapleton was
almost as eager for the event as herself; but he knew no more, and
was just dismissing her, disappointed, when the waggon was heard
approaching. While she waited a moment, straining her sight to
make out whether it was the right party, before she ran to meet them,
her brother jumped out, and even Castle started up with more
alacrity than he had shown since they landed. Before they could well
greet one another, Stapleton came up to ask where Frank’s tools
were, and to tell him that he was wanted very much indeed. He could
not refuse him permission to go forward one mile, in order to deposit
Castle and his wife at their new abode; but he lent a hand towards
emptying the waggon of his workman’s packages, and gave him
notice that he should be glad to see him back the first possible
moment.
“You will soon find what great people such as we are here,” said
Ellen, laughing. “This is the place to grow proud in. No more
lounging about the fields, Frank; no more leaning over gates
chewing straws, while nobody inquires for one. You will never need
to touch your hat and ask for work here; people will come begging
you to be so very kind as to put up a door for any pay you please.
This is the place to grow proud in.”
Frank observed, with a grave smile, that pride was dangerous to
one in Ellen’s place.
“Well, then, I will be proud of you, and you shall be proud of me;
and no harm can come of that.”
The first time that the brother and sister could obtain a few
minutes’ conversation without being overheard, Frank inquired,
“Now, Ellen, tell me straight forward. How do you like your
change?”
“Why, I more than half like it; but there are some things I do not
like.—It is a fine thing to be so well off, and to know that I shall be so:
and I do not mind the work, though it is rather hard, to be sure; and
my cows are nothing but a credit to me, and I have seen no animals
to be afraid of when I go out milking, though some of them leap
about very strangely indeed; and my mistress makes much of me, as
I told you; and her little worries are not much to be wondered at
when one thinks of the confusion we live in just now; and I dare say
there will be an end of them when we get our soap and candles
made out of the house, and another hand or two to help in the
brewing and washing. And then to think that father and you are so
well off——”
“But tell me what there is that you do not like.”
Ellen almost shuddered when she whispered that her fellow
servant, who ate at the same table, and slept in the same room, and
was her companion almost all day, was a convict, and had been sent
to this country for robbing an aged mistress who had confided in her,
and deserved gratitude instead of treachery from her. To be
compelled to hold daily and hourly intercourse with such a person
was a very great evil, and one which doubled Frank’s anxiety about
his sister. He was glad to hear that there was a probability of the
woman marrying as soon as she could obtain a remission of her
servitude by steady conduct.
A half smile which he perceived on Ellen’s lips when this part of
the story was being told, made him question her further respecting
the evils of her situation, or the trials which she might not be
disposed to consider exactly as evils. The idea in her mind was that
which he suspected,—that she might quit her service before her
convict companion.—Frank looked graver than ever. Who—what—
where was he,—the person that seemed to have made advances in
Ellen’s good graces already? She was eager to explain that there
was no one in particular yet. It was too early for her to have looked
about and settled her mind yet;—but there was this one, and that
one, and the other one, that carried her pail for her, morning and
evening, however busy he might be; or was ready to teach her how
to clean and card wool; or showed her what a pretty little homestead
he was about to have in the neighbourhood, and intimated how
happy she might be as the mistress of it.
“They hinder my work sadly, and their own too,” continued she,
blushing, “for all I tell them that I have nothing to say to any body yet.
I am so afraid any of them should have been convicts, (though I am
sure Harry Moore never was;) and I dare not ask mistress any thing
about them.”
“Ask her, by all means,” said Frank. “Or I will ask your master, if
you wish it. They only can tell us, and it is a point we must find out.
Meantime, keep to your business as quietly as you can. What makes
you so sure that Moore (is not that his name?) was never a convict?”
Ellen could give no better reason than that she could wager her
life upon it. She thought her brother grown very pertinacious on a
sudden, because this was not perfectly satisfactory to him; but Frank
was not pertinacious—only wary and affectionate.
Chapter V.
THE CASTLES AT HOME.
It was very well for a man of Castle’s irritable temper to be made a

shepherd, instead of a labourer at home, within sight and hearing of
all the bustle and difficulty occasioned by much pressure of work and
few hands to do it. He could not have borne to be, as his wife said,
driven from pillar to post,—called off from one thing before he had
done it, to do something else to which he was altogether
unaccustomed. It suited him much better to be out upon the downs
after the sheep; though even in that quiet place he had his troubles.
The sheep-walk was too extensive to be under the management of
one person; and Castle’s brother-shepherd was not a very congenial
companion. He was a gentleman convict;—a young man who had
gamed away his little fortune, and then taken to swindling, for which
he had been transported. Being unequal to hard work, and having no
mechanical skill, he was sent out to tend sheep; an employment as
little suitable as might be to his social dispositions and active habits.
The two reluctant companions agreed only in their inclination to
grumble.
“They call this a fine scene,” observed the young man, “but it does
not suit my taste. I had rather see our sheep in the Smithfield pens
than on these downs. Then one misses the London cries, however
much the magpies chatter here. As for the cooing of the doves, it
really depresses the spirits. People talk of the stars being so brilliant
here,—like golden lamps; but I like real lamps better. A row of them
in Pall Mall is worth a hemisphere of stars.”
“I don’t know much about lamplight,” replied Castle, “having been
too poor to burn candles at home, and so going to bed in the twilight;
but this place is so lonesome, I sometimes wonder whether it is in
the world or out of it. All this view is like an old deserted park, to be
sure; but where is the squire’s house, or the church steeple, and the
children coming out of school? There is no public-house far or near;
and no parson or his lady to speak a word to one: only a young man
that comes to read prayers on Sundays in a shed or on the green,
and away again to do the same thing somewhere else. Not such a
thing have I seen since I came as a carriage with ladies in it; and
they say there are no hunts. With all the game there is here, no
scarlet jackets ever come in sight from the woods.”
“That is the worst of it,” responded the other grumbler. “We have
all the dulness of a country life without its solace of amusement. It
was really too tantalizing lately, to see a kangaroo hunt which I could
not join. If they would let me take my turn, I might be of some use to
them as an experienced huntsman. I should like to hunt opossums
till I could get skins enough to make your pretty daughter a cloak
worthy to be worn; and——”
Castle here moved off impatiently, having too much paternal pride
to listen to convict wooing on Ellen’s behalf. The young man followed
him, continuing,
“The snipe-shooting is very choice, I’m told, in the marshes
yonder. I must have leave when winter comes on, to go and try my
luck. But the hunts are the best things,—more spirited perhaps than
you are aware of.”
“Hunts! hunts!” cried Castle. “I see neither deer nor fox. An odd
sort of hunting, if you mean killing any of these leaping things, with
their queer ways. Why, the little ones don’t run beside their dams, as
is natural, but she pops them into her bag, and off she hops, as if
she had only two legs. The first I saw, I thought she had happened
an accident, and had her fore legs cut short; and I thought she got
on wonderfully well considering; and then in a minute appeared a
whole herd of them, with their young in their bags.”
“It is a pretty sight to see them come down from the woods at
sunrise to feed in the plains. Then is the time to hide behind a
thicket, and make sure of one’s game. Which do you prefer, as a
bottom dish, kangaroo or bandicoot? In a pie, properly seasoned, it
is difficult to say which is the best. I have given many a hint down
below that either is much more palatable to me than rations of salt
meat.”
Castle, who thought no man need desire more in the way of diet
than to eat meat every day, looked with contempt on the grimaces of
his companion over his ample supply of beef, wheaten bread, and
cider.
“If you want to hunt,” said Castle, “I wish you would kill off the vile
beasts that have been making havoc among my lambs. I might have
got at one, but I was downright scared with its ugliness.”
“Was it the hyena or the devil?”
“O, the devil, to judge by its looks. It is as big as a middle-sized
dog, with the head of an otter, crowded with teeth. It moved very
slow, but I could do no better than stare at it.”
“They call it the devil here,” replied the gentleman. “You should dig
little pits, and set your dogs upon it when it has fallen in. It will go on
worrying your lambs, unless you keep on the watch.”
“Another thing that puts me out,” observed Castle, “is that the
beasts are one below another here, as if they were bewitched. In
England, we have a horse of one size, and a dog of another, and a
rat of another; and none of them is like the rest; but here we have a
big kangaroo, and a kangaroo the size of a dog, and another no
bigger than a rat; and these last are not real kangaroos. I declare it
makes my hair stand up to see a rat leaping like a real kangaroo: just
as it would to see a mouse shaking its mane and trotting and
cantering like a horse. I have not been used to such freaks, and this
is a country I can’t understand.”
“I hope to understand it better,” replied the convict. “I was always
fond of roving, and in time I may have explored farther than we can
see from these green hills that we both find so dull. What do you
mean to do when you get free?”
“They may settle that that got me bound,” replied Castle, testily.
Then, struck with a sense of his own ingratitude, he added, “To be
sure, if there is no squire’s house, there is no workhouse either; and
if I see no acquaintance, there is nobody to taunt me with misfortune;
but, on the contrary, they make much of me at home. And there’s
——”
“Your daughter.”
“What; my little Susan! Yes, they make a handy little thing of her
already, and——”
“I mean the other handy one, Ellen.”
“She will do well enough, Sir, I assure you. She has a fine spirit
and a steady mind of her own, and a proud brother to take care of
her; and that is better than a broken-down father; though it should go
hard with me but I would protect her, Sir, if there was no one to do it
better.”
So saying, Castle walked off, showing by his manner that he was
not sociably disposed.
His wife was much more altered within a short time by her change
of circumstances than he. The first thing that seemed to affect her
favourably was the use that was made of her little daughter in the
household arrangements. When the farmer’s wife found that her new
domestic was indolent and indifferent, she endeavoured to make the
best of a poor bargain by squeezing as much work as she could out
of Susan. The child was willing enough, and proud to find herself of
so much use; but her mother was jealous on her account, and began
for the first time to show symptoms of tenderness for her. She not
only argued in her defence, but helped her when she was more
disposed to proceed with her work than to “go and play;” words
which had little charm for a child whose associations with play were
those of hunger, scolding, mockery, and all the miseries of pauper
life. When the farm servants rose at daybreak to go forth for the day,
Susan was always ready to jump up at the first word, to replenish the
wallets and fill the cans, though her mother turned round in bed, and
muttered that it was too soon to get up. She needed no reminding
about tending the house-lamb, and feeding the poultry, and dusting
as much of the coarse furniture as she could reach. After breakfast,
if any one would lift her upon the dresser, or lay the utensils and the
bowl of water on the floor, she would wash up without breaking
anything: and she was always at hand to carry messages into field or
farm-yard, or to help with dinner and supper, or to carry letters to the
spot where they were to be deposited in readiness for the postman’s
weekly call; and when not able to do anything better, she could scare
away the crows and cockatoos from the fields and garden. Her
mother thought this a hard life for a little girl: but Susan was stout,
rosy, and merry; and the farmer himself found a few minutes now
and then to take her on his knee and teach her the alphabet, in
preparation for the time when a schoolmaster could be brought
within reach. The first thing Mrs. Castle did heartily was washing up,
one day when Susan had nearly scalded her fingers. She took more
and more of the child’s work from her, and still Susan turned to
something else; so that ere long, both were pretty fully employed;
and in proportion as the once reckless and lazy pauper became
interested in the occupations going on around her,—in proportion as
she bestirred herself to get the baking done while the house was
clear of the men, and the washing over in time to have a chat in the
evening,—she grew like the active and tidy housewife she would
never have become in her own land.
A circumstance which hastened this change was the opportunity
she now had of gratifying one taste,—almost the only taste she ever
had, and which seemed to have died out under the hardships of her
condition: a taste for gardening. When a girl, she had had a garden;
and as long as her husband had owned an acre of ground, she took
possession of a corner of it for her pinks and roses, under pretence
of growing vegetables for the family. From that time to this, nobody
had heard her mention fruit or flowers; but Ellen bore in mind her
love for them now when the remembrance might be turned to some
purpose. She mentioned, in her step-mother’s presence, that her
master was trying what he could do in the management of vines, for
the growth of which the climate was peculiarly favourable; and that
whether he got any wine or not, his trouble would be more than
repaid by his profits from his other fruits. The peaches, to be sure,
were not of the best sort, though so plentiful as to lie rotting on the
ground, after bushels had been thrown to the pigs; but the apricots,
and yet more the raspberries, which grew to such a size and in such
quantity as no English person would believe without seeing them,
were likely to prove a good speculation, being sent to a distance in
the form of jam. Sugar being remarkably cheap in this country, there
was little risk in trying a batch of sweetmeats, which were to be sent
to India for sale by a vessel from Launceston. The idea was caught
up as Ellen expected it would be; and as the farmer and his wife did
not take to the scheme of fruit-growing as heartily as was desired,
the emigrant family tried whether they could not get a garden of their
own. The small part of their wages which they were yet at liberty to
use was applied to the purchase of a plot of ground, and Frank found
time to work in it, and Ellen procured wherewith to stock it, and their
step-mother haunted it early and late, before and after work,—and
Castle himself relaxed his brow, and spoke in a tone that was not
querulous, as he looked round upon that which, however small, was
so much more than he had ever expected to possess again,—a
family property.
“Look at father,” whispered Ellen to her brother. “I have not seen
him and her arm-in-arm since I was no bigger than Susan.”
“He is like a prisoner that has been quite shut up coming for the
first time into the gaol court,” said Frank. “The feel of the air makes
him push his hat up from over his eyes. Only set him quite free, and
he will uncover his brows, and lift up his head like a man.”
“And so he ought,” replied Ellen, “since it is for no fault of his own
that he has been bound down to poverty.”
“Ah! poverty is a cold and dreary prison, Ellen. That puts me in
mind,—have you seen, I wonder, any thing that has surprised you
very much lately,—any thing that you would like to tell me if you were
sure of not being overheard, or of not being thought fanciful?”
“The word ‘prison’ put me in mind of something that I have been
wanting to talk to you about almost this month past.—I don’t know
how to believe my own eyes about it, but I am sure I have seen Jerry
in our farm-yard at night, and lurking about among the sheds before
most of our folks are stirring in the morning.”
“Aye; but was he all alone, Ellen?”
“Bob was never with him, that I could see; but he seemed to me to
be waiting for somebody off our premises, and I thought it must
naturally be me. So twice I ran out to catch him; but once I was
crossed by two of our people that I don’t choose to come in the way
of; and the other time he was whispering with the same two, so that I
dared not go near. How could he get liberty? and what could he be
about?”
“Something very deep, I am afraid,” replied her brother. “As to the
liberty,—it is no difficult matter for convicts who behave pretty
regularly to get hours of liberty at the beginning and end of the day;
and the lads being employed on roadmaking so near, accounts for
our getting a glimpse of them sometimes. But what I am uneasy
about is Jerry’s having so much to say to the convicts at your place
and mine;—for I have seen him at Stapleton’s oftener than you have
among your people. I am afraid of some plot——”
“O, mercy!” cried Ellen; “what sort of a plot?”
“That is more than I can say. Sometimes they plot, I hear, for
nothing worse than to escape; but some have had to do with the
natives, (who are little better than wild beasts), and have brought
them down upon the farms, setting them to steal and even to
murder; for which they pay the poor savage creatures by helping
themselves with their wives.”
Ellen trembled while she asked whether any of the natives could
be in the neighbourhood.—Her brother hoped not, as the
government had declared that they were driven back among the
mountains, where they must soon die out as their wild cattle had
done: but as long as any convicts were disposed to bush-ranging,—
and some did actually escape every year,—he could not, for his part,
feel quite secure. He thought he should speak to Stapleton about it.
Meanwhile, he desired Ellen to drop not a syllable that should alarm
her father, or anybody else.
“I hope, sister,” he continued with some hesitation, “I hope Harry
Moore has no acquaintance, more or less, with Jerry, or any other
such people.”
Ellen’s eyes flashed as they used to do when she was a
passionate little girl at school.
“Harry!” she cried. “Harry Moore have any sneaking doings! Harry
Moore keep bad company! You don’t know Harry a bit better than the
very first day,—the day when you thought he might be a convict
himself.”
“No need to be angry, Ellen. He might just know him enough, you
see, to say ‘How d’ye do?’ when they meet, and to judge how often
Jerry might fairly be here.”
“After all,” said Ellen, sighing, “it is my father’s own son that I flew
off about his being acquainted with; so there is no need for me to be
so proud. No; Harry does not know either of the lads, even by sight;
but I shall tell him what you have been saying, though nobody else,
Frank.”
“Certainly. Conceal nothing that weighs upon your mind from
Harry, any more than if he was your husband already. I look to him to
help me to keep an eye upon Bob, who may be made something of,
they say, little hope as there is for Jerry. Bob works within bounds at
spare hours, instead of roving into the bush, or prowling about the
settlers’ farms, where he has no business. Bob must be saving
money fast, unless he has unseen ways of spending it. He works
hard, and is well paid for his extra labour. He may have the
advantage of me, after all; and settle on a place of his own before
me.”
“Because he got a free passage as a punishment. That is really
very hard, Frank.”
“Harry Moore will be the first at liberty, however, Ellen; and that I
am glad of on your account. I am soon to begin building you a
house, at over hours; and you may depend on my doing my best to
have it all complete by the time the six months are gone.”
“Six months!” cried Ellen.
“Why, I do not mean that you need wait till then. You may fairly
marry as soon as you like;—and many in our own country would be
glad to have that said to them. I only mentioned six months as the
time when Harry would be all his own master. Then I shall hope to
see you milking a cow of your own.—Meantime, till I have found out
more about Jerry, be cautious how you get out of reach of those that
will take care of you.”
Ellen sighed, and smiled, and wondered which was the strangest
world,—the one she had left behind, or the new one which seemed,
after several months, nearly as foreign as when she had entered it.
She had no doubt which was the pleasantest. How could she, when
a vague fear and thorough dislike of some of the people in the
neighbourhood were the only set-off against the prosperity of all
whom she loved, and her own bright prospects with such a husband
as Harry Moore had promised to be?

Diatom Morphogenesis Diatoms Biology and Applications Vadim V Annenkov Full Chapter

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Diatom Morphogenesis (Diatoms:

Biology and Applications) Vadim V.

Diatoms: Biology and Applications

Series Editors: Richard Gordon (dickgordoncan@xplornet.com) and Joseph Seckbach

Wiley Global Headquarters

Limit of Liability/Disclaimer of Warranty

Library of Congress Cataloging-in-Publication Data

Printed in the USA

Part 2: Simulation 105

6.3.3.2 Comparison of 2D Lattice Parameters and Degree

7.2.1.8 Protocol for Using Centric Diatom 3D Surface Models

8.6.2 Krylov Methods: Associating Valve Surface With Microtubule

Part 3: Physiology, Biochemistry and Applications 251

11.4.4 Adaptive Behavior and Evolutionary Changes in Diatoms Linking

13.12 Exteriorization of the Valve 321

16.6 Conclusion 399

Why Valve Morphogenesis is Important?

Topics Addressed in This Volume

Keywords: Diatom morphology, tutorial, LM and SEM, frustule morphology

1.1 Diatoms in Brief

*Corresponding author: kalina.manoylov@gcsu.edu

(a) (b) (c)

(d) (e) (f)

1.2 Tools to Explore Diatom Frustule Morphology

(h) (i) (j) (k) (l)

(s) (t) (u) (v) (w) (x)

When we should Observation of Observation Observation of the Observation Understanding

1.3 Diatom Frustule 3D Reconstruction

1.3.1 Recommended Steps to Understand the Complex Diatom Morphology:

Matching symmetry states to morphogenetic stages may inform the developmental

2.1.1 Recognition and Symmetry

(a) (b) (c)

(d) (e) (f)

[2.154], helices/spirals (including handedness) [2.159], and compound or multiple spi-

(a) (b) (c) (d) (e)

(f) (g) (h) (i) (j)

(k) (l) (m) (n)

2.1.2 Symmetry and Growth

2.1.3 Diatom Pattern Formation, Growth, and Symmetry

self-similarity as well [2.96]. The role of symmetry in developmental processes is discern-

2.1.4 Diatoms and Uncanny Symmetry

THE CASTLES AT HOME.

It was very well for a man of Castle’s irritable temper to be made a

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