Chapter 1

The Algae and their General Characteristics

Susan E. Douglas*
Institute for Marine Biosclences, National Research Council, Halifax,
Nova Scotia B34 3Z1, Canada

John A. Raven
School of Life Sciences, University of Dundee, Dundee 001 4HN, U:«.

Anthony W. D. Larkum
School of Biological Sciences, University of Sydney, NSW 2006, Australia

Summary 1
I. Introduction 2
II. The Algae: Their Origins and Diversity 2
III. The Green, Red and Brown Algae 4
IV. The Chromophytes 5
A. The Heterokonts 5
1. The Phaeophytes 5
2. The Bacillariophytes (Diatoms) 5
B. The Haptophytes 5
C. The Cryptomonads 6
D. The Dinoflagellates 7
V. The Chlorarachniophytes 7
VI. The Euglenophytes 7
VII. Algal Genomes 7
VIII. Algae as Sources of Natural Products 8
IX. Concluding Remarks 8
Acknowledgements 9
References 9

Summary

In contrast to the land plants, algae have very diverse mechanisms of photosynthesis, and especially of light-
harvesting pigments and assemblages. This diversity is inherited from a great diversity of plastid types with
different evolutionary histories, not withstanding the fact that all plastids appear to be derived by endosymbiosis
from Cyanobacteria or their forebears. The major groups of algae are therefore related to the type ofprotist host
and the type of plastid, and these are described. In most groups of algae it appears that the plastid has been
derived by at least two serial endosymbioses. A single endosymbiosis appears to have oecurred in the green
algae, the red algae and the glaucocystophytes. The rich variety of types and mechanisms has given rise to many
biochemical products which today form the basis of a growing biotechnology industry. Algae are important
economically in many other ways. From a photosynthetic point of view the algae will be a rich source of ideas
for many years to come.
*Author for correspondence, email: [email protected]

Anthony W. Larkum, Susan E. Douglas and John A. Raven (eds): Photosynthesis in Algae, pp. ].··10.
© 2003 Kluwer Academic Publishers.
2 Susan Douglas, John Raven and Anthony Larkum

I. Introduction II. The Algae: Their Origins and Diversity

Land plants have inherited from their green algal The algae, all of which carry out oxygenic
ancestors a remarkably uniform strategy for photosynthesis by means of a cytoplasmic organelle,
harvesting and converting light energy into chemical the plastid, constitute a diverse grouping of protists
energy. This is based on processes using the mag- scattered in the crown group of protists (Fig. 1).
nesium-tetrapyrroles chlorophyll a (Chi a) and Within the different algal groups there arc non-
chlorophyll b (Chi b), together with a limited number photosynthetic sub-groups, but through close affin-
of carotenoids. In eukaryotic organisms photosyn- ities with their photosynthetic cousins, they can be
thesis is carried out in cellular organelles known as judged to have lost plastids secondarily. A moot
plastids. Only in terms ofcarbon fixation has evolution point is found with the apicomplexans which are
brought about any changes at a cellular and plastid non-photosynthetic parasitic protists (e.g. Plasmo-
level; in the way in which carbon is fixed as C J or C4 dium, which causes malaria) in which there exists a
acids and the presence or absence of photo systems I relic plastid. These organisms are not currently placed
and II in a limited number of plastid types. Using a in the algae, although their algal roots are acknowl-
limited range ofpigments, harvesting oflight energy edged (Chapter 2, Larkum and Vcsk).
involves the same basic mechanism throughout the The origin of algae occurred about 1-1.5 Ga ago
land plants. In sharp contrast, the diverse assemblage by the capture of free-living oxygenic phototrophic
of organisms grouped together under the general eubacteria (most likely ancestors of present-day
classification of algae, or photosynthetic protists, Cyanobacteria) by host cells that were early protists.
have evolved a great assortment of pigments and Most workers in the field consider this endosymbiotic
many elaborate strategies for light-harvesting and origin so well-supported that it is taken to be axio-
energy conversion. These 'variations on a theme' matic. These host cells already possessed a mito-
have been selected during evolution, resulting in the chondrion (or mitochondrion-like organelle), whose
current diversity of form and function. However, origin was also certainly by endosymbiosis through
because the algae have played a rather small direct the acquisition of a free-living e-proteobacterium.
role in the economies of the world up to the present Thus the origin ofalgae and ofmitochondriate protists
time, they have received scant attention. Fortunately was through the formation ofa chimaera. And, indeed
this situation is changing rapidly and is likely to the origin ofthe first eukarya may also be chimaeric:
change even more rapidly in the future. Biotechnology from eubacterial and archaebacterial origins (Gupta
already offers a number of ways to harness products and Golding, 1996). While Cyanobacteria, as pro-
from algae and in the future this will no doubt karyotes, are not, strictly speaking, algae, they are
increase. Global climate change has also provided an conventionally dealt with to varying extents in texts
impetus to research on algae as solar collectors and on algae and many of the chapters in this volume
converters of clean energy. In Japan, for example, consider Cyanobacteria as well as eukaryotes.
several large laboratories working on algae have Both the origin of the plastid and the origin of the
been granted funds based on carbon credits according host in the chimaera which gave rise to the first algal
to the Convention on Global Climate Change. cells are controversial. Using Ockham's razor one
Nevertheless there is still a long way to go before we would seek a single origin for both. Hypothetically, a
can say that we really have the basic knowledge single cyanobacterium-like eubacterium would enter
needed to understand the scope and breadth of algal into symbiosis with a single kind ofmitochondriate
photosynthesis. We hope that this book will be timely protist and from this first photosynthetic eukaryote
in mapping out the past work and putting forward a all the other algae would arise. The majority of
logical framework for future research in the area. workers in the field broadly support such a single
event (monophyly) as the origin ofall primary plastids
surrounded by two membranes. However, there are
difficulties with this simple version especially in
terms of plastid characters (Chapter 2, Larkum and
Abbreviations: Chi - chlorophyll; EST -- expressed sequence
tags; kbp - kilobasc pairs; NCB I - National Centre for
Vesk). There are also difficulties that arise from the
Biotechnology Information, http://www.ncbi.nlm.nih.gov/; rather haphazard distribution ofalgae on phylogenetic
rRNA - ribosomal ribonucleic acid trees based on small subunit rRNA or other genes. It