Algae: Anatomy, Biochemistry, and Biotechnology

© 2010 Taylor & Francis Group, LLC

ALGAE BARSANTI • GUALTIERI

SECOND

EDITION

K13023

“ ... stands out for its in-depth information on structural and mechanical anatomy,

with flagella as the most prominent example. The meticulous and elegant

drawings of algal apparatuses and their mechanics make it easy to understand

complex structures and functions, as well as constitutes another outstanding

feature of this book.”

—Senjie Lin, Marine Sciences, University of Connecticut, Groton, The Quarterly

Review of Biology, Vol. 81, December 2006

“ ... the authors concentrate on highlighting interesting and illuminating topics,

with the idea of inciting the sort of wonder and curiosity that will encourage

further outstanding research.”

—Willem F. Prud’homme van Reine, Blumea, 2006, Vol. 51, No.3

A single-source reference on the biology of algae, Algae: Anatomy, Biochem￾istry, and Biotechnology, Second Edition examines the most important taxa

and structures for freshwater, marine, and terrestrial forms of algae. Its com￾prehensive coverage goes from algae’s historical role through its taxonomy and

ecology to its natural product possibilities.

The authors have gathered a significant amount of new material since the

publication of the first edition. This completely revised second edition contains

many changes and additions including the following:

• All revised and rewritten tables, plus new figures, many in color

• A fascinating new chapter: Oddities and Curiosities in the Algal World

• Expanded information on algal anatomy

• Absorption spectra from all algal divisions, chlorophylls, and accessory

pigments

• Additional information on collection, storage, and preservation of algae

• Updated section on algal toxins and algal bioactive molecules

The book’s unifying theme is on the important role of algae in the earth’s

self-regulating life support system and its function within restorative models

of planetary health. It also discusses algae’s biotechnological applications,

including potential nutritional and pharmaceutical products. Written for students

as well as researchers, teachers, and professionals in the field of phycology and

applied phycology, this new full-color edition is both illuminating and inspiring.

Biological Sciences

K13023_Cover.indd 1 1/15/14 12:40 PM

© 2010 Taylor & Francis Group, LLC

ALGAE

SECOND EDITION

ANATOMY, BIOCHEMISTRY,

AND BIOTECHNOLOGY

© 2010 Taylor & Francis Group, LLC

ALGAE

LAURA BARSANTI • PAOLO GUALTIERI

Istituto di Biofisica

Pisa, Italy

SECOND EDITION

ANATOMY, BIOCHEMISTRY,

AND BIOTECHNOLOGY

CRC Press

Taylor & Francis Group

6000 Broken Sound Parkway NW, Suite 300

Boca Raton, FL 33487-2742

© 2014 by Taylor & Francis Group, LLC

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Version Date: 20130827

International Standard Book Number-13: 978-1-4398-6733-4 (eBook - PDF)

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© 2010 Taylor & Francis Group, LLC

Alla Lilli, perché è sempre la mia mamma, anche se mi fa diventare

matto come quando ero piccino [(ti voglio bene, mamma!)]

Paolo

To my mom, Silvana (1926–2008), and my dad, Renzo (1928–2009),

because I know they are still watching over me, and to Bernard

(1952–2011) pour son voyage en solitaire (nakupenda tarepanda)

Laura

vii

© 2010 Taylor & Francis Group, LLC

Contents

Preface............................................................................................................................................ xiii

Authors.............................................................................................................................................xv

Chapter 1 General Overview ........................................................................................................1

Definition......................................................................................................................1

Classification ................................................................................................................2

Occurrence and Distribution........................................................................................2

Structure of Thallus—Cytomorphological Types........................................................6

Unicells and Unicell Colonial Type.........................................................................8

Filamentous Type .................................................................................................. 10

Siphonocladous Type............................................................................................. 13

Siphonous Type...................................................................................................... 13

Parenchymatous and Pseudo-Parenchymatous Type............................................. 14

Palmelloid Type..................................................................................................... 15

Nutrition ..................................................................................................................... 16

Reproduction .............................................................................................................. 17

Vegetative and Asexual Reproduction................................................................... 17

Binary Fission or Cellular Bisection................................................................. 17

Zoospore, Aplanospore, and Autospore ........................................................... 18

Autocolony Formation...................................................................................... 18

Fragmentation................................................................................................... 18

Resting Stages................................................................................................... 18

Sexual Reproduction..............................................................................................20

Haplontic or Zygotic Life Cycle........................................................................20

Diplontic or Gametic Life Cycle.......................................................................20

Diplohaplontic or Sporic Life Cycles................................................................20

Summaries of the 11 Algal Phyla...............................................................................22

Cyanobacteria........................................................................................................22

Glaucophyta...........................................................................................................24

Rhodophyta ...........................................................................................................25

Chlorophyta ...........................................................................................................29

Charophyta ............................................................................................................ 32

Haptophyta ............................................................................................................ 33

Cryptophyta........................................................................................................... 35

Ochrophyta ............................................................................................................ 35

Cercozoa—Chlorarachniophyceae........................................................................39

Myzozoa—Dinophyceae .......................................................................................39

Euglenozoa—Euglenophyceae .............................................................................. 41

Endosymbiosis and Origin of Eukaryotic Photosynthesis......................................... 42

Suggested Reading .....................................................................................................46

Chapter 2 Anatomy .....................................................................................................................49

Cytomorphology and Ultrastructure ..........................................................................49

Outside the Cell.....................................................................................................49

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© 2010 Taylor & Francis Group, LLC

Type 1—Simple Cell Membrane ......................................................................49

Type 2—Cell Surface with Additional Extracellular Material ........................50

Type 3—Cell Surface with Additional Intracellular Material

in Vesicles.........................................................................................................60

Type 4—Cell Surface with Additional Extracellular and Intracellular

Material............................................................................................................. 62

Flagella and Associated Structures.......................................................................66

Flagellar Shape and Surface Features ..............................................................68

Flagellar Scales.................................................................................................68

Flagellar Hairs..................................................................................................70

Flagellar Spines ................................................................................................72

Internal Features of the Flagellum ........................................................................72

Axoneme...........................................................................................................72

Paraxial Rod .....................................................................................................73

Other Intraflagellar Accessory Structures........................................................ 74

Transition Zone................................................................................................. 75

Basal Bodies.....................................................................................................79

Root System......................................................................................................82

How Algae Move...................................................................................................93

Swimming.........................................................................................................93

Movements Other than Swimming...................................................................99

Buoyancy Control ...........................................................................................100

How a Flagellum Is Built: The Intraflagellar Transport................................. 102

How a Flagellar Motor Works........................................................................ 103

How a Paraxial Rod Works ............................................................................ 104

The Photoreceptor Apparata ............................................................................... 104

Types of Photoreceptive Systems............................................................................. 106

Type I................................................................................................................... 106

Type II ................................................................................................................. 108

Type III................................................................................................................ 109

Photoreceptive Proteins....................................................................................... 111

Fundamental Behavioral and Physiological Features.......................................... 111

Sampling Strategies........................................................................................ 112

Trajectory Control........................................................................................... 113

Signal Transmission........................................................................................ 114

An Example: Photoreceptor and Photoreception in Euglena.............................. 114

Chloroplasts.................................................................................................... 118

The Nucleus, Nuclear Division, and Cytokinesis...........................................126

Ejectile Organelles and Feeding Apparata ..................................................... 132

Suggested Reading ................................................................................................... 137

Chapter 3 Photosynthesis.......................................................................................................... 141

Light ......................................................................................................................... 141

Photosynthesis.......................................................................................................... 144

Light-Dependent Reactions...................................................................................... 145

PSII and PSI: Structure, Function, and Organization ......................................... 153

ATP Synthase ...................................................................................................... 155

ETC Components ................................................................................................ 155

Electron Transport: The Z-Scheme..................................................................... 157

Proton Transport: Mechanism of Photosynthetic Phosphorylation..................... 158

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© 2010 Taylor & Francis Group, LLC

Pigment Distribution in PSII and PSI Super-Complexes of Algal Division ....... 160

Light-Independent Reactions.................................................................................... 160

RuBisCO.............................................................................................................. 166

Calvin–Benson–Bassham Cycle.......................................................................... 167

Carboxylation ...................................................................................................... 167

Reduction............................................................................................................. 167

Regeneration........................................................................................................ 167

Photorespiration................................................................................................... 168

The Energy Relationships in Photosynthesis: The Balance Sheet........................... 168

Suggested Reading ................................................................................................... 170

Chapter 4 Working with Light .................................................................................................. 173

How Light Behaves .................................................................................................. 173

Scattering.................................................................................................................. 173

Absorption................................................................................................................ 174

Interference............................................................................................................... 175

Reflection.................................................................................................................. 175

Refraction ................................................................................................................. 177

Dispersion................................................................................................................. 178

Diffraction................................................................................................................ 178

Field Instruments: Use and Application................................................................... 181

Radiometry............................................................................................................... 181

Measurement Geometries: Solid Angles.................................................................. 181

Radiant Energy......................................................................................................... 182

Spectral Radiant Energy .......................................................................................... 182

Radiant Flux (Radiant Power).................................................................................. 182

Spectral Radiant Flux (Spectral Radiant Power) ..................................................... 182

Radiant Flux Density (Irradiance and Radiant Exitance)........................................ 182

Spectral Radiant Flux Density ................................................................................. 183

Radiance................................................................................................................... 183

Spectral Radiance..................................................................................................... 184

Radiant Intensity ...................................................................................................... 184

Spectral Radiant Intensity........................................................................................ 185

Photometry ............................................................................................................... 185

Luminous Flux (Luminous Power) .......................................................................... 185

Luminous Intensity................................................................................................... 185

Luminous Energy..................................................................................................... 188

Luminous Flux Density (Illuminance and Luminous Exitance) ............................. 188

Luminance................................................................................................................ 188

Lambertian Surfaces................................................................................................ 188

Units Conversion ...................................................................................................... 189

Radiant and Luminous Flux (Radiant and Luminous Power)............................. 189

Irradiance (Flux Density).................................................................................... 190

Radiance .............................................................................................................. 190

Radiant Intensity.................................................................................................. 190

Luminous Intensity.............................................................................................. 190

Luminance........................................................................................................... 190

Geometries........................................................................................................... 190

PAR Detectors.......................................................................................................... 191

The Photosynthesis–Irradiance Response Curve (P vs. E Curve)........................... 193

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© 2010 Taylor & Francis Group, LLC

Photoacclimation...................................................................................................... 196

Suggested Reading ................................................................................................... 197

Chapter 5 Biogeochemical Role of Algae................................................................................. 199

The Role of Algae in Biogeochemistry.................................................................... 199

Limiting Nutrients....................................................................................................200

Algae and the Phosphorus Cycle..............................................................................202

Algae and the Nitrogen Cycle ..................................................................................204

Algae and the Silicon Cycle .....................................................................................209

Algae and the Sulfur Cycle ...................................................................................... 212

Algae and the Oxygen–Carbon Cycles.................................................................... 214

Suggested Reading ................................................................................................... 218

Chapter 6 Algal Culturing ........................................................................................................ 221

Collection, Storage, and Preservation ...................................................................... 221

Culture Types...........................................................................................................224

Culture Parameters...................................................................................................226

Temperature.........................................................................................................227

Light.....................................................................................................................227

pH ........................................................................................................................227

Salinity ................................................................................................................227

Mixing .................................................................................................................228

Culture Vessels.........................................................................................................228

Media Choice and Preparation.................................................................................229

Freshwater Media ................................................................................................230

Marine Media ......................................................................................................230

Seawater Base...........................................................................................................240

Nutrients, Trace Metals, and Chelators.................................................................... 241

Vitamins...................................................................................................................243

Soil Extract...............................................................................................................244

Buffers......................................................................................................................244

Sterilization of Culture Materials ............................................................................245

Culture Methods....................................................................................................... 252

Batch Cultures..................................................................................................... 253

Continuous Cultures............................................................................................ 255

Semicontinuous Cultures.....................................................................................256

Commercial-Scale Cultures ................................................................................ 257

Outdoor Ponds.......................................................................................................... 257

Photobioreactors.......................................................................................................259

Culture of Sessile Microalgae .............................................................................259

Quantitative Determinations of Algal Density and Growth ....................................260

Growth Rate and Generation Time Determinations...........................................264

Suggested Reading ...................................................................................................265

Chapter 7 Algae Utilization ...................................................................................................... 267

Introduction .............................................................................................................. 267

Sources and Uses of Algae.......................................................................................268

Human Food........................................................................................................268

Contents xi

© 2010 Taylor & Francis Group, LLC

Cyanobacteria .................................................................................................268

Rhodophyta..................................................................................................... 271

Ochrophyta (Phaeophyceae)........................................................................... 274

Chlorophyta .................................................................................................... 279

Animal Feed........................................................................................................282

Extracts................................................................................................................286

Agar ................................................................................................................287

Alginates.........................................................................................................288

Carrageenan....................................................................................................289

Fertilizers............................................................................................................. 291

Cosmetics.............................................................................................................293

Functional Foods and Nutraceuticals.......................................................................294

Toxins .................................................................................................................. 301

Selected Reading......................................................................................................305

Chapter 8 Oddities and Curiosities in the Algal World ............................................................309

In the Realm of Darkness.........................................................................................309

Algae–Animal Interaction: Riding a Sloth, Swinging on a Spider Web,

Swimming in a Jelly . . ............................................................................................. 314

Some Like It Cold . . . ...............................................................................................320

Some Like It Hot . . .................................................................................................. 322

Some Like It Dry . . . ................................................................................................324

Selected Reading...................................................................................................... 325

xiii

© 2010 Taylor & Francis Group, LLC

Preface

In the seven years since the first edition of this book was published, we have built up a large amount

of new material and data in the field of algology, based on our own experiences in reading, writing,

and reviewing. With the aid of all this information, we have completely revised the book, introduc￾ing the following changes and additions:

• We have added 27 new figures for a total of 205 figures, many of them in color

• All the 38 tables have been revised and rewritten

• We have updated the literature in all chapters

• We wrote an entirely new chapter on how odd algae can be

• We have rewritten Chapter 1, updating the classification of algae and modifying the section

on the endosymbiosis and origin of eukaryotic photosynthesis

• We have expanded Chapter 2, adding new types of root systems and algal swimming pat￾terns and modifying the section on photoreception and photoreceptors

• We have updated Chapter 3, adding absorption spectra measured on samples from all algal

divisions together with their decomposition in pigments. We have also added the absorp￾tion spectra of all the chlorophylls and the accessory pigments

• We have modified notation and wording of Chapters 4 and 5

• We have expanded section on collection storage and preservation in Chapter 6, adding new

information on automatic algae recognition and classification

• We have rewritten Chapter 7, updating the section on algal toxins and algal bioactive

molecules

• We have, of course, corrected the numerous errors present in the first edition (we do apolo￾gize for them), doing our best to avoid errors in this new edition

Like the previous edition, this book is written and designed for undergraduate and postgraduate

students with a general scientific background, having their first academic experience with the world

of algae, as well as researchers, teachers, and professionals in the field of phycology and applied

phycology. Our major commitment is still the same, challenging and stimulating both students and

teachers to move beyond the limit of the written page to further explore not only the topics high￾lighted in the book, but also all the new ideas that can spring to mind (we hope!) after reading each

chapter.

Though updated, the bibliography is still by no means exhaustive; we have not attempted to be

comprehensive and many excellent papers will be missing. Our intention was to put in only enough

to lead the readers into the right part of the primary literature in a fairly directed manner and to

provide a sort of orienteering compass in the “mare magnum” of scientific literature.

We are deeply grateful to the staff at CRC Press, Boca Raton, FL, particularly our patient and

comprehensive editor John Sulzycki for trusting us enough to ask for a second edition and to the

senior project coordinator Jill Jurgensen, who had to cope with all our e-mail.

Again, our sincere gratitude and a special thanks to Valter Evangelista for his skillful assistance

and ability in preparing the final form of all the drawings and illustrations, and for his careful atten￾tion in preparing all the technical drawings of this second edition. We appreciate his efforts to keep

pace with us both and to cope with our ever-changing demands and corrections and second thoughts

without getting too upset. We know we have driven him crazy.

And we will always be grateful to Vincenzo Passarelli, who took care of the lab, making our

work lighter and smoother. Next February he will retire, leaving our group after more than 30 years;

xiv Preface

© 2010 Taylor & Francis Group, LLC

we have grown old together and we already know we will miss his smile, and his special trumpet￾like whistling.

For the new illustrations present in the book, we are indebted to Luca Barsanti, brother of Laura

and Maria Antonietta, who succeeded in realizing most of the drawing of our book before dying in

February 2005. He made the drawing work in a wonderful way, confirming his artistic skill. Though

almost eight years have passed by, and some snow has also fallen on his roof, he is still the same

light-hearted and amusing company who delighted us during the preparation of the first edition. We

will be always grateful to him.