Textbook An Introduction To Phytoplanktons Diversity and Ecology 1St Edition Ruma Pal Ebook All Chapter PDF
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Ruma Pal · Avik Kumar Choudhury
An Introduction to
Phytoplanktons:
Diversity and Ecology
An Introduction to Phytoplanktons:
Diversity and Ecology
Ruma Pal • Avik Kumar Choudhury
An Introduction
to Phytoplanktons:
Diversity and Ecology
Ruma Pal
Avik Kumar Choudhury
Department of Botany
University of Calcutta
Kolkata, West Bengal, India
v
vi Preface
The development of this book was made possible by the financial support
from University Grants Commission (UGC), Govt. of India, for two research
projects on phytoplankton dynamics of Eastern Indian coast and other
ecological niche. Dr. Avik Choudhury and Ms. Anindita Singha Roy worked
as Project Fellows, and their research findings are represented as Case
Study in Chap. 5.
Financial support from Council of Scientific and Industrial Research
(CSIR), Govt. of India, is also acknowledged for survey work at Indian
Sunderbans area under NIMTLI program. Mr. Nirupam Barman and Sri Gour
Gopal Satpati surveyed different parts of Sunderbans, and their work is also
included in this book.
vii
Contents
ix
x Contents
Dr. Ruma Pal did her M.Sc. and Ph.D. from University of Calcutta. Presently,
she is Associate Professor in Botany, Department of Botany, University of
Calcutta, India. Prior to this assignment she had served in Presidency College,
Calcutta and Nara Sigh, Dutt College Howrah. She has more than 30 years of
research experience in Phycology. Her research interest is related to various
fields of algal biotechnology, like, Phycoremediation, aquaculture, biofuel
production, nanotechnology, etc. Algal diversity study, phytoplankton
dynamics and ecological modelling are also the areas of her interest. She has
already conducted more than 15 research projects in the field of algal applica-
tion and has published more than 50 papers in refereed journals.
xi
xii About the Author
part of the twentieth century, Professor P.S. In early times, the classification was mainly
Welch wrote the first American textbook on lim- based on the morphotaxonomy. But very
nology. Similar books were also written like recently the situation is marked by the quest of
Fundamentals of Limnology by Franz Ruttner a compromise between the conventional (artifi-
(1940) and Professor G.E. Hutchinson published cial) system and the phylogenetic system
the book entitled Treatise on Limnology (1957, together with the molecular genetics. The classi-
1967) which is considered as standard reference cal approaches using morphological characters
work throughout the world. do not reflect the phylogenetic relationships,
In the nineteenth century, it was felt to publish e.g. molecular data revealed that among the
the journals on limnology that would pull together green algae, the genera with spherical ball-type
the increasing volume of limnological informa- thallus evolved independently in different
tions that were developing every day. Thus, on lineages of algae. On the other hand, highly
January 1, 1936, the Limnological Society of diverse morphotypes can belong to one and the
America was established, which was later same phylogenetic lineage.
(1948) named as the ‘American Society of In algal systematics nowadays, a polyphasic
Limnology and Oceanography’ and used to pub- approach is widely suggested. Therefore, phylo-
lish till date the most well-circulated and popular geneticists suggest the following criteria for
journal titled Limnology and Oceanography. The proper identification and phylogenetic place-
year 1948 is also marked for the formation of the ment of any algal groups:
‘Freshwater Biological Association’ in Britain. 1. Conventional morphological and ecophy-
This association maintains a continuous record of siological study of algae under field
physical, chemical and biological informations condition
of 17 lakes in the Lake District of northwest 2. Isolation of unialgal culture for morphological,
England. Similarly ‘Istituto Italiano di ontogenetic, biochemical and physiological
Idrobiologia’ in Italy carried out intensive studies studies under laboratory conditions
on northern Italian lakes, and the science of lim- 3. Ultrastructural studies of different cell organ-
nology in Europe flourished. Similar other elles of algal cell, especially the chloroplast
renowned institutes developed around this time ultrastructure
in Europe, for example, at Plön, Germany; 4. Use of molecular markers (conserved sequences)
Uppsala and Lund, Sweden; Copenhagen, for molecular phylogenetic analysis
Denmark; and on Lake Constance, Germany. The most commonly used sequences are the
16S rRNA or the small subunit (SSU) of ribo-
somal RNA (rRNA) for prokaryotic cells and 18S
1.3 Algal Classification rRNA for eukaryotic cells. Besides some other
markers are also used for phylogenetic analysis,
Algae are considered as the most primitive e.g. rbcL and tufA gene.
plants with well diversifications. They all have a Therefore, for convenience we can consider
chlorophyll-bearing thalloid plant body and two phases of algal systematics. In the first
their reproductive structures are without sterile phase, starting from placement of algae in
jackets. They have variations in morphology plant kingdom by Eichler (1886), followed by
and cellular biochemistry together with repro- algal classification by Fritsch (1935) and Smith
ductive behaviour and life cycle patterns. On the (1950), up to Bold and Wynne’s (1985) system,
basis of these characters, algologists classified only morphological characters were consid-
the entire algal kingdom into different divisions ered for taxonomic purpose. Therefore, these
and classes from time to time. classifications can be considered under ‘old
The systematics or classification of algae classical taxonomy’. In another approach or in
has been changing dramatically through ages. the second phase, other parameters including
1.3 Algal Classification 3
ultrastructural, biochemical and molecular between algae and fungi was recognized as the
characters are also considered – reflecting the presence or absence of chlorophyll.
evolution and phylogeny of and can be desig- This system of classification is considered as
nated as ‘modern system of classification’. incorrect by most of the scientists now. Indeed,
we think it is still needed to be taught to a bota-
nist to understand the phylogenetic position of
1.3.1 Old Classical Taxonomy algae and other plant groups which also indicate
the evolutionary tendency among different plant
Eichler (1886) mentioned the phylogenetic groups in a broader sense.
position of algae in relation to other plant In the old classical system of algal classifica-
groups. He divided the entire plant kingdom tion, famous algologists like Smith, Fritsch,
into two groups: Cryptogamia (spore-producing Prescott, Bold and Wynne and Chapman and
plants) and Phanerogamia (seed plants). These Chapman have classified the algal kingdom on
two groups are again divided into three divi- the basis of variation in cell structure (prokary-
sions and four classes. Algae are placed in Class otic and eukaryotic), flagellar position, number
I of division Thallophyta under Cryptogamia in and structure, pigment composition, reserve food
close association with fungi. The only difference matters, mode of reproduction, etc.
Plant kingdom
A. Cryptogamae B. Phenerogamae
(Spore producing plants) (Seed plant)
-Division I –Thallophyta
(Lower plant with a thallus) Division I- Gymnospermae Division II- Angiospermae
(Plants with naked seeds) (Plants with enclosed seeds)
Class I- Algae Class II- Fungi Class I- Monocotyledonae Class II- Dicotyledonae
(Plants with 1 cotyledon) (Plants with 2 cotyledons)
algae. The termination ‘phyceae’ has been 3. Chrysophyceae – Members are with brown-
adopted wherever the class includes forms with or orange-coloured chromatophores contain-
an algal organization, while for flagellates the old ing one or more accessory pigments. Starch
designation is retained. Therefore, he designated is absent, but naked pyrenoid-like bodies are
the different groups of algae as ‘classes’ – rather occasionally present. Fat and a compound
than ‘divisions’. leucosin are found in the form of rounded
The 11 classes were characterized as follows: whitish opaque lumps as the food storage.
1. Chlorophyceae (Isokontae) – Members with A large proportion of the members are flag-
grass-green-coloured chromatophores and ellate and devoid of a special cell membrane.
contain the same pigment compositions and The motile cells possess one or two (rarely
approximately in the same proportions as in three) flagella attached at the front end. The
higher plants. Starch is the customary form of cells typically contained one or two parietal
storage products of photosynthesis, often chromatophores. The most advanced habit is
(especially in resting stages) accompanied by that of a branched filament. Sexual reproduc-
oil, and pyrenoids commonly surrounded by tion is extremely rare and not yet quite
a starch sheath are frequently present inside clearly established in any one case; the exist-
the chloroplasts. Cells are surrounded by a ing records point only to isogamy. The class
cell wall in which cellulose is the constitu- is widely distributed in freshwaters, but a
ent. The motile cells are with equal whiplash few are marine. An example is Chromulina.
type of flagella (commonly two or four) 4. Bacillariophyceae (diatoms) – Unicellular
which arise from the front end of swarmers. members with yellow or golden-brown
In many members the cells contain only one chromatophores containing accessory brown
or few chromatophores. The members of pigments. Pyrenoid-like bodies are often
Chlorophyceae exhibit sexual reproduction present and the products of photosynthesis
– ranging from isogamy to advanced oogamy. are fat and volutin. All the members are
Most of the taxa are haploid with zygote rep- unicellular or colonial. A cell wall is always
resenting the only diploid phase, but some present and is composed of mainly silica and
exhibit a regular alternation of generation partly of pectic substances. The cell consists
with similar haploid and diploid individuals. of two halves, each composed of two or more
The class is more widely represented by pieces, and is commonly richly ornamented.
freshwater members than in salt water, and One set of forms (Centrales) is radially and
there is a marked terrestrial tendency. Examples the other (Pinales) bilaterally symmetrical.
are Chlamydomonas and Spirogyra. The diatoms produce a special type of spore –
2. Xanthophyceae (Heterokontae) – The mem- the auxospore. The Pinales show a special
bers are with yellow-green chromatophores type of sexual fusion between the protoplasts
due to presence of xanthophylls as major of the ordinary individuals. The members of
pigment. Starch is absent and storage product this class are probably diploid. Diatoms are
is oil. The algal members have a cell wall very widely distributed in the sea and in all
which is often rich in pectic compounds. The kinds of freshwaters, as well as in the soil
motile cells possess two very unequal flagella and in other terrestrial habitat. Examples are
(or sometimes only one) arising from the Navicula and Chaetoceros.
front end. As a general rule the cells contain a 5. Cryptophyceae – The members are of flagellate
number of discoid chromatophores. Sexual organization and with oogamous type of repro-
reproduction is always isogamous. The most duction. The cells are with usually two large
advanced forms have a simple filamentous parietal chromatophores showing very diverse
habit. All are probably haploid. The class is pigmentation. Pyrenoid-like bodies occur, but
more widely distributed in freshwater than in appear often to be independent of the chromato-
the sea. An example is Vaucheria. phores; the products of photosynthesis are
1.3 Algal Classification 5
nor flagellate members are known. Evident The divisions were characterized by the fol-
protoplasmic connections are the rule lowing basic characteristics:
between the cells of the majority of forms. 1. Chlorophyta – The grass-green algae with
Most of the Rhodophyceae are marine. All the major pigments chlorophyll a and b
exhibit sexual reproduction of an advanced together with carotenes and xanthophylls.
oogamous type, the female organ having a Photosynthetic reserves are usually stored in
long receptive neck and the antheridium the form of starch. It is always in association
producing but a single motionless male cell. with pyrenoid. Motile stages have flagella of
As a result of fertilization, special spores equal length with a few exceptions. The zoo-
(carpospores) are produced from bunches of spores and motile gametes have two or four
threadlike structures that arise from the flagella. Sexual reproduction is a phenomenon
female organ after fertilization. The of wide occurrence within the group and in
Rhodophyceae are either haploid or exhibit various orders it ranges all the way from isog-
a regular alternation of similar haploid and amy to oogamy. This group also shows a wide
diploid individuals, the latter bearing char- range in vegetative structures (Chlorella).
acteristic sporangia (tetrasporangia), each 2. Euglenophyta – All the members are unicel-
producing four spores. An example is lular and most of them are naked free-
Polysiphonia. swimming cells with one, two or three flagella.
11. Myxophyceae (Cyanophyceae) – Members Many of the genera have grass-green, discoid
with a simple type of cell, containing at the band-shaped or stellate chloroplasts, with or
best only a very rudimentary nucleus (central without pyrenoids. The chloroplasts contain
body) and without a proper chromatophore, the same chlorophylls as Chlorophyceae,
the photosynthetic pigments being diffused beta-carotene and xanthophylls unlike
through the peripheral cytoplasm. The pig- Chlorophyta. Food reserve is paramylum;
ments present are chlorophyll, carotene, nutrition may be holophytic, holozoic or sap-
phycocyanin and phycoerythrin, the last two rophytic. There are one or two contractile
being in varying proportions and the colour vacuoles at the anterior end of the cell, which
of the cells being very commonly blue green. are connected with a reservoir, which in turn
The products of photosynthesis are sugars is connected with the cell’s exterior by a
and glycogen. No motile stages are known narrow gullet (Euglena).
and all the members have a membrane 3. Chrysophyta – The members have yellowish-
around the cell. There is no sexual reproduc- green to golden-brown pigment because of
tion. The members of this class are of simple the predominance of carotenes and xantho-
organization and many propagate entirely by phylls. The food reserves include a complex
simple division or by vegetative means. Most carbohydrate leucosin and oils. Some mem-
types are filamentous, many of them with a bers are with cell wall made up of silica and
peculiar ‘false’ branching. They occur very are composed of two overlapping halves.
abundantly in freshwaters and in terrestrial Cells may be flagellated or non-flagellated,
habitats and are not common in the sea. An solitary or united in colonies of definite or
example is Spirulina. indefinite shape. Sexual reproduction is usu-
But in 1950, Smith proposed seven divisions ally isogamous by a union of flagellated and
of algae in his system of classification as follows: non-flagellated gametes, e.g. members of
1. Chlorophyta present-day Xanthophyta (Vaucheria) and
2. Euglenophyta Bacillariophyta (Nitzschia).
3. Chrysophyta 4. Phaeophyta – The Phaeophyta or brown algae
4. Phaeophyta have many celled complex type of plant body
5. Pyrrophyta that is usually of macroscopic size and distinc-
6. Cyanophyta tive shape. The chromatophore or pigment-
7. Rhodophyta bearing organelles are yellowish brown in
1.3 Algal Classification 7
colour due to the presence of xanthophylls in cystocarp. Pigments are localized in chro-
greater amount than that of chlorophyll and matophores. In addition to chlorophylls,
carotenes. The two principle reserve foods are carotene, xanthophyll and R-phycoerythrin
laminarin, a polysaccharide, and mannitol. and R-phycocyanin are present.
Zoospores or gametes are pyriform with two Prescott (1984) first classified the algal king-
laterally inserted flagella of unequal length. dom into seven phyla like other groups of bio-
Reproductive organs are of two kinds – the one logical kingdom. Each phylum is again divided
celled or unilocular reproductive organ is into different classes and orders as follows:
always a sporangium and born on a diploid thal- I. Phylum Chlorophyta
lus. The other kind of reproductive organ is (i) Class – Chlorophyceae (17 orders)
many celled and with each cell containing a (ii) Class – Charophyceae (1 order)
single gamete or single zoospore. Most of the II. Phylum Euglenophyta
members of Phaeophyta have a life cycle in III. Phylum Chrysophyta
which there is alternation of two independent (i) Chrysophyceae
multicellular generations, haploid and the other (ii) Bacillariophyceae
is diploid. The two generations may be identical (iii) Heterokontae
in size and structure, in others they are dissimi- IV. Phylum Pyrrophyta
lar in both size and structure (Ectocarpus). (i) Desmokontae
5. Pyrrophyta – Members of this division are (ii) Dinokontae
greenish to golden brown. The pigments are V. Phylum Phaeophyta
Chl a, Chl c, beta-carotene and four xantho- (i) Isogeneratae
phylls. Photosynthetic compounds are reserved (ii) Heterogeneratae
as starch and also oil. The nucleus is distinc- VI. Phylum Rhodophyta
tive in which chromatin lies in numerous bead- Subphylum – Bangioideae (4 orders)
like structures on thread. Cell wall when Subphylum – Florideae (6 orders)
present contains cellulose. VII. Phylum Cyanophyta
6. Cyanophyta – The Cyanophyta or blue-green Subphylum – Coccogoneae
algae are a distinctive group sharply delim- Subphylum – Hormogoneae
ited from other algae (prokaryotic algae). Among the old classical algal taxonomy,
Their pigments are not localized on chro- Bold and Wynne’s (1985) system of classification
matophores; rather they are present in the is the most well-accepted one. He divided the
peripheral portion of the protoplast and algal kingdom into nine divisions and introduced
include chlorophyll a, carotenes and distinc- a new division Charophyta, which is considered
tive xanthophylls. In addition, there is a blue as progenitor of land plants.
pigment (C-phycocyanin) and a red pigment
(C-phycoerythrin). The unique feature of Divisions
Cyanophyta is the presence of primitive type I. Cyanophyta and Prochlorophyta
of nucleus within the cell (central body) II. Chlorophyta (16 orders)
which lacks a nucleolus and a nuclear mem- III. Charophyta
brane. They lack any flagellated structure and IV. Euglenophyta
devoid of sexual reproduction (Oscillatoria). V. Phaeophyta (13 orders)
7. Rhodophyta – The Rhodophyta or red algae VI. Chrysophyta (6 class)
have multicellular thalli of microscopic or Chrysophyceae
macroscopic size and often of distinctive Prymnesiophyceae
shape. Red algae differ from all other algae Xanthophyceae
in structure of their sexual organs, in mode Eustigmatophyceae
of fertilization followed by formation of a Raphidophyceae
spore-producing structure, the so-called Bacillariophyceae
8 1 A Brief Introduction to Phytoplanktons
from other primitive eukaryotes like algae and ultrastructural and biochemical and more recently
protozoans. the molecular data are also considered for algal
4. The five-kingdom classification gives a clear classification together with the morphological
indication of cellular organization and modes characters of algal thallus. Since then the science
of nutrition. of phycology has sustained major conceptual
However, the five-kingdom classification has changes. The increased resolution of the electron
some demerits also, particularly with reference to microscope revealed the presence and structure
the lower forms of life: of flagella, flagellar hairs, flagellar roots, eyespots,
1. The kingdoms Monera and Protista include chloroplast, endoplasmic reticulum, etc., which
both photosynthetic (autotrophic) as well as were found to be important in basic systematics
non-photosynthetic (heterotrophic) organisms of algae especially the green algae. Important
and organisms which have cells with cell wall observations were made by Pickett-Heaps (1967,
as well as without cell wall. 1969 , 1972a, b , 1975), which revealed that
2. The three higher kingdoms or multicellular different types of microtubular arrangements are
lines have originated from Protista several involved in cytokinesis of green algae. In some
times (polyphyletic). orders of Chlorophyta, cell divisions are charac-
3. Unicellular or colonial green algae like terized by the collapse of the interzonal spindle
Chlamydomonas and Volvox have not been apparatus after mitosis, which give rise to a
included under Protista because of their ‘phycoplast’ with the microtubules oriented in
resemblance to other green algae. the plane of cell division, viz. Volvocales,
4. Slime moulds are different from other members Tetrasporales, Chlorococcales, Oedogoniales
of Protista. and Ulotrichales. On the other hand, some
5. Viruses have not been given proper place in members together with land plants have a persis-
this system of classification. tent spindle and develop a cleavage and furrow
Nevertheless, the five-kingdom classification (Klebsormidiales) or a ‘phragmoplast’ in which
has found a wide acceptance with biologists all the microtubules are oriented perpendicular to
over the world. the plane of cytoplasmic division (Coleochaetales,
Charales and Conjugales). At the same time, bio-
Status of Viruses chemical analysis also clarified the presence and
The position of viruses in the biological kingdom structure of algal pigments, storage products and
is one of the unsolved mysteries. Due to the cell wall constituents. Considering all these
absence of a cellular organization, viruses cannot parameters, Stewart and Mattox (1975) classified
be placed with either prokaryotes or eukaryotes. the green algal division Chlorophyta into five
They are considered as intermediate between classes, considering comparative cytology, viz.
living and nonliving systems. Viruses are active Chlorokybales, Zygnematales, Klebsormidiales,
and show reproduction only inside the host cell. Coleochaetales and Charales.
In the free state, they are totally inactive. They
may even be purified and crystallized like chemi- 1.3.2.1 Algal Chloroplast
cal substances. Viruses have a genetic material in Classification
represented by either DNA or RNA, surrounded Mereschkowski (1905) first studied about the
by a protein sheath. Viruses reproduce by using nature and origin of chloroplast in eukaryotic
the metabolic machinery and raw materials of the algae and the evolutionary pattern among them.
host cell. Because of these peculiarities, viruses In 1905, he published the most extensive paper
do not fit into any of the five kingdoms of life. on the origin of chloroplast based on the idea
From 1975 onwards, with the advent of elec- that eukaryotic algal cell originated by the pro-
tron microscopy and other sophisticated instru- cess of endosymbiotic events between the cya-
ments like high-resolution SEM, TEM, HPLC nobacterial cell and the primitive eukaryotic
and HPTLC, fine characters of algal cells like phagocytotic protozoa. He was with the opinion,
10 1 A Brief Introduction to Phytoplanktons
structure thylakoids embedded in it, are present in The starch is similar to that of higher plant and
three divisions of algae, viz. Glaucophyta, is composed of amylose and amylopectin.
Rhodophyta and Chlorophyta (group II). These A pyrenoid is a differentiated region within the
algae evolved through primary endosymbiosis. chloroplast that is composed of polypeptides
Among these three algal divisions, in chloro- with enzymatic properties of ribulose-1,5-
plast of Glaucophyta, thylakoids are arranged bisphosphate carboxylase that are capable of
equidistantly at peripheral region of chloroplast fixing carbon dioxide. Storage products are fre-
(like cyanobacterial cell). There are similarities quently associated with pyrenoids. The pyrenoid
between chloroplast of Glaucophyta and cyano- is denser than the surrounding stroma and may or
bacterial cell as follows: may not be traversed by thylakoids.
• They are about the same size. The chloroplast of group III algal divisions,
• They evolve oxygen in photosynthesis. viz. Euglenophyta, Dinophyta and Apicomplexa,
• They have 70S ribosomes. is surrounded by two membranes of the chloro-
• They have circular prokaryotic DNA without plast envelope and one membrane of chloroplast
basic proteins. endoplasmic reticulum, which is not continuous
• They have peptidoglycan wall surrounding the with nuclear membrane. The thylakoids are
chloroplast envelope. grouped in bands of three. Major pigments
For this reason chloroplast of Glaucophyta is present are Chl a and b (Euglenophyta); Chl a
termed as ‘cyanelles’ or the incipient chloroplast. and c2 with peridinin as major xanthophylls
The chloroplast of Rhodophyta (rhodoplast) is (Dinophyta); and rudimentary chloroplast
also surrounded by two chloroplastic membranes (Apicomplexa).
with no chloroplast ER. Inside the chloroplast, In addition, some accessory chloroplasts are
thylakoids occur singly and the DNA molecule present in Dinophyta (originated from further
occurs as microfibrils and the phycobilin endosymbiotic process between dinoflagellate
pigments are localized into phycobilisomes on cell and Cyanophyta or Bacillariophyta or
the surface of the thylakoids (similar to Rhodophyta) giving characteristic colour.
Cyanophyceae). Chl a and d are present inside The membranes of the fourth evolutionary
the thylakoids. Among the carotenoids zeaxan- group Heterokontophyta contain chloroplast
thin is found in the greatest quantities. having four membranes (two chloroplast enve-
Phycobiliproteins include R-phycocyanin, allo- lopes and two chloroplastic endoplasmic
phycocyanin and three forms of phycoerythrin reticula).
in maximum amounts. The chloroplast of Phaeophyta is known as
In Chlorophyta, chloroplasts are highly phaeoplast. Phaeoplast is a four-membrane-
evolved like higher-plant chloroplast, with two bound structure having three thylakoids per band.
membranes of chloroplast envelope; thylakoids Membrane-bound structures are also present
are stacked to form grana which are embedded in between chloroplast envelope and chloroplast
the matrix called stroma. Chloroplast pigments ER. The chloroplast contains Chl a, c1 and c2
are also similar to higher plants. Chlorophylls a with major carotenoid, fucoxanthin. Pyrenoids of
and b are present and the main carotenoid is Phaeophyceae are stalklike structures which set
lutein. Extraplastidic carotenoids are sometimes off from the main body of chloroplast containing
present (haematochrome). granular substances not traversed by thylakoids.
The chloroplastic DNA is partially looped Surrounding the pyrenoid is a saclike structure
and ribosome is of 70S type. Starch is formed in containing the reserve food matters, generally
the chloroplast in association with pyrenoid. present (laminarin as major component).
12 1 A Brief Introduction to Phytoplanktons
1.3 Algal Classification 13
1.3.2.2 Outline of Lee’s (2008) band and no chloroplast ER, floridean starch
Classification with Basic grain synthesized in the cytoplasm, no flagella,
Characters of Different Groups pit connections between the cells. Reproductive
In this system of algal classification, the entire algal unit spermatia and carpogonia. Post fertilization
kingdom is classified into four distinct groups: change prominent.
I. Prokaryotes Single class – Rhodophyceae
II. Eukaryotic algae with chloroplast with two Chlorophyta: Members have chlorophylls a and b
membranes and starch is the reserve food matter formed
III. Eukaryotic algae with chloroplast with one within the chloroplast. Chloroplast with two
chloroplast ER (total three membranes) membranes only, thylakoids form the grana.
IV. Eukaryotic algae with chloroplast with two Both freshwater and marine in habitat with wide
chloroplast ER (total four membranes) range in morphology. Flagella isokont type with
Group I: fine hairs if present. Flagellar root cruciate type
1. Cyanobacteria or unilateral type. Eyespot present.
Group II: Four classes: Prasinophyceae, Charophyceae,
1. Glaucophyta Ulvophyceae and Chlorophyceae
2. Rhodophyta Euglenophyta: Euglenoid flagellate, unicellular,
3. Chlorophyta surrounded by pellicle. Chloroplast with two
Group III: chloroplastic membranes and one chloroplastic
1. Euglenophyta endoplasmic reticulum containing Chl a and b.
2. Dinophyta Marine or freshwater in habitat. Cytosome or
3. Apicomplexa gullet-like structures are present at the anterior
Group IV: region of the cell.
1. Cryptophyta Single class: Euglenophyceae
2. Heterokontophyta Dinophyta: Unicellular with two halves, epi-
3. Prymnesiophyta cone and hypocone, made up of thecal
Cyanobacteria: Prokaryotic members containing plates. Chloroplast with two chloroplastic
chlorophyll a and phycobiliproteins. Some membranes and one chloroplastic endoplas-
members contain chlorophyll b and are known mic reticulum with Chl a and c2 and the
as green cyanobacteria (Prochlorophyta). Cell carotenoids peridinin and neoperidinin as
wall is similar to Gram-negative bacteria major pigments. Storage product is starch.
containing peptidoglycan layer outside the Two flagella – one longitudinal and one
cell membrane. Naked circular DNA present transverse.
at the central portion of the cell. Cyanophycin, Single class: Dinophyceae
the polymer of arginine and aspartic acid; Apicomplexa
carboxysome; polyphosphate bodies; and Unicellular having reduced colourless plastid
polyglucan granules are the other cellular called apicoplast. Apicoplast and dinoflagellate
inclusions. Gas vacuoles, containing a large plastids originated from red algae by a single
number of gas vesicles, are present. endosymbiotic event. The apical complex consists
Glaucophyta: Members unicellular with primitive of a polar ring and a conoid formed of spirally
type of chloroplast, showing many characters coiled microtubules. Apicomplexa are endopara-
similar to cyanobacterial ancestor and are sites that cause some of the most significant
known as cyanelle. Pigments are similar to tropical diseases like malaria or diarrhoea. The
cyanobacteria. Members are unicellular parasite attaches to the host cell with the conoid
flagellates. protruding to produce a stylet that forms a tight
Rhodophyta: Both marine and freshwater in habitat. junction with the host cell. The apicomplexan
Unicellular to huge thallus, especially for marine cell is taken up into the host cell in the parasi-
seaweeds. Chloroplast with one thylakoid per tophorous vacuole.
14 1 A Brief Introduction to Phytoplanktons
important book, Phytoplankton Pigments in groups) from a protistan perspective has been
Oceanography: Guidelines to Modern Methods proposed by Adl et al. (2005), which is mainly
(SOR-UNESCO volume). They listed 12 micro- used in oceanographic literature. According to
algal classes as common members of phyto- them the traditional ‘kingdoms’, such as Metazoa,
planktons, viz. diatoms, dinoflagellates, Fungi and Plantae, are recognized as deriving
haptophytes, chrysophytes, rhodophytes, raphi- from monophyletic protist lineages. The authors
dophytes, cryptomonads, chlorophytes, eugleno- grouped the molecular phytogenies into six clus-
phytes, eustigmatophytes, cyanobacteria and ters as follows:
prochlorophytes. 1. Opisthokonta: animals, fungi, choanoflagellates
The use of advanced HPLC and ultrahigh- and Mesomycetozoea
performance liquid chromatography (UPLC) 2. Amoebozoa: traditional amoebae, slime
helps to identify different phytoplankton taxa in moulds, etc.
addition to it considering culture code and gene 3. Rhizaria: foraminifera, radiolarian, heterotro-
bank references. phic flagellates, etc.
A new system of classification of eukaryotes 4. Archaeplastida: red algae, green algae,
(including only photosynthetic microalgal Glaucophyta and Plantae
— Milloin?
Käsi nousee. Näyttää siltä kuin hän aikoisi tukkia korvansa. Mutta
sitte käsi taas hervahtaa takaisin paareille, joilla hän makaa.
Rakkaus äitiin auttaa häntä. Hän tahtoo unohtaa ja elää. Hänellä on
suuri tehtävä.
Siunaa ja varjele,
Siunaa ja varjele
Sodan tantereilla kärsineitä,
Haavat Sä lääkitse
Ijällä pitkällä
Palkitse, kruunaa heitä!
veisaa kuoro.
Iljan katse kostuu, ja autuas ilo virtaa koko hänen olemuksensa
läpi.
Ilja painuu takaisin vuoteelle. Totta, totta siis kuitenkin se, mitä hän
ei ole tohtinut eikä tahtonut uskoa.
*****
— Oli —?
— Haim Jankel?
*****
— Mitä ihmeessä?
Tällainen oli heidän ystävyytensä alku ollut. Ja sitä jatkui siitä sekä
läpi koulun että yliopistossa.
Tällä kannalla olivat asiat, kun sota puhkesi. Se repi kohta kuin
pohjan heidän jalkojensa alta. Pitkään aikaan he eivät puhuneet siitä
mikä tässä asiassa oli kipeintä heille. Mutta kerran se purkautui kuin
vahingossa.
*****
Hän koetti välttää näitä ajatuksia. Mutta Dimitri näki hänen kerran
ryntäävän ulos tallista paeten kuin henkensä edestä. Ja Dimitri
ymmärsi.
*****
— "Sisar", huudahti Dimitri äkkiä ohikulkevalle
sairaanhoitajattarelle.
— Tulkaa tänne, minulla olisi asiaa.