4 THE ALGAE

some groups, e.g. Chrysophyceae, Bacillariophyceae and Xantho-

phyceae, and between these as a whole and the Phaeophyceae; also
between the Dinophyceae and Cryptophyceae, and the Rhodo-
phyceae and Myxophyceae. These relationships are, however,
discussed more fully later (see pp. 315, 321).

It should be noted that chlorophyll a, one of the components of

the green pigment, is the most abundant chlorophyll in all types of
algae, that jS carotene is equally widely distributed, that starch is
not a widely distributed food reserve outside the Chlorophyceae
and that cellulose is not the universal wall constituent. There are a
large number of xanthophylls and each algal class possesses its own
characteristic members. The Siphonales, an order of the Chloro-
phyceae, is treated separately because of the carotinoid or yellow
pigment differences. The phycobilins of the Rhodophyceae and
Myxophyceae are not absolutely identical and hence are differen-
tiated by the prefix r- and c- respectively. In several classes of
algae, e.g. diatoms, Cryptophyceae, Chlorophyceae, colourless
forms are known. Such organisms have to lead a saprophytic

existence.

Using a combination of the pigment and biochemical differences,

together with morphological differences, the algae as a group can
be classified as follows :

Euphycophyta:

Myxophycophyta :
Chrysophycophyta :

Charophyceae
Chlorophyceae
Phaeophyceae
Rhodophyceae

Myxophyceae

Chrysophyceae
Xanthophyceae
Bacillariophyceae

Pyrrophycophyta :

Cryptophyceae
Dinophyceae

Some workers consider that the Chlorophyceae, Phaeophyceae

and Rhodophyceae should have independent rank (e.g. Chloro-

CLASSIFICATION 5

phycophyta, Phaeophycophyta, Rhodophycophyta), and a few

would also treat the Charophyceae in the same way. The present
writer considers that phylogenetically these four classes are dis-
tantly related (see p. 309) and therefore groups them together.
Ultimately the classification adopted must depend upon the view
taken of the inter-relationships of these groups. A class of wholly
fossil organisms, the Nematophyceae, is also included among the
algae, but this is somewhat tentative as we have no means of know-
ing whether the plants, which only existed in the very earliest
times, really were algae.

A brief summary of the principal characteristics of the different

classes follows.

EUPHYCOPHYTA

(i) Chlorophyceae

This group used to comprise four great subclasses, the Isokontae

(equal cilia), Stephanokontae (ringed cilia), Akontae (no cilia) and
Heterokontae (imUke cilia). These subdivisions, as their names
imply, were based upon the organs of locomotion of the motile re-
productive bodies. The last subdivision has since been renamed
the Xanthophyceae and, because of its relationships with the
Bacillariophyceae and Chrysophyceae, has been transferred to the
Chrysophycophyta (see below). The structure of the Chloro-
phyceae covers a great range from simple unicells to multicellular
plants that are quite complex, though not as complex as those of
the Phaeophyceae and Rhodophyceae. In the Chlorophyceae the
chloroplasts vary considerably in shape and size whilst the final
product of photosynthesis is mostly starch together with fat.
Structures known as pyrenoids are often present and a starch
sheath can generally be demonstrated around them. The motile
cells of most members of the class are very similar and commonly
possess either two or four equal fiagellae, but in the Oedogoniales
(the former Stephanokontae) there is a ring of cilia whilst in the
Conjugales (the former Akontae) organs of locomotion are absent.
Sexual reproduction is of common occurrence and ranges from iso-
gamy to anisogamy and oogamy. Asexual reproduction normally
takes place by means of motile zoospores, but a variety of non-
motile spores (p. 19) may also be produced. The colour of the
cells is usually grass green because the pigments are the same as

6 THE ALGAE
those present in the higher plants and, furthermore, they are
present in much the same proportions.

The Characeae comprise one predominantly fresh-water order,

the Charales, which differ from other green algae (Chlorophyceae)
because of their remarkable morphological and reproductive fea-
tures. They are consistently uniform in structure with characteris-
tic nodes and internodes and whorls of branches arising at the
nodes. There is no asexual reproduction and the sexual reproduc-
tive organs present an elaborate oogamy. There is some evidence
for beheving that the plants may have a common ancestry with the
Chaetophorales (seep. 115).

(2) Phaeophyceae

This group comprises the common brown algae of the seashore,

and it is worth noting that the majority are wholly marine, pre-
dominantly of cold waters. There are a few, rare fresh- water
species. The brown colour is due to the fucoxanthin pigment
which masks chlorophyll a (see Table i). The principal product of
photosynthesis is a sugar alcohol, known as mannitol, together with
the polysaccharide laminarin, whilst the cell walls not only contain
cellulose but also specific materials such as algin and fucoidin (see
p. 452). In some of the larger and more advanced brown algae these
compounds are present in sufficient quantity to be of commercial
importance (see p. 446). The class is further characterized by the
fact that the sugar residues tend to have a i : 3 linkage instead of
the more normal i : 4 linkage between the carbon atoms. The
simplest forms are filamentous, and there are all stages of develop-
ment and increasing differentiation up to the large seaweeds of the
Pacific and Arctic shores with their great size and complex internal
and external differentiation. There is also a number of greatly
reduced parasitic and epiphytic forms. The pyriform, motile repro-
ductive cells, which possess two flagella, one directed forwards
and the other backwards, are commonly produced in special organs
or sporangia that are either uni- or plurilocular. Some members
possess non-motile, asexual reproductive spores (monospores and
tetraspores). Sexual reproduction ranges from isogamy to oogamy,
but in the latter case the ovum is liberated before fertiUzation. The
life cycles are extremely diverse, and although alternation of
generations does occur, it is an irregular rather than a regular
process.

CLASSIFICATION 7

(3) Rhodophyceae

The members of this class form the red seaweeds, and although
most of them are marine, nevertheless a few occur in fresh water.
Their colour, red or bluish, is produced by the phycobihn pig-
ments r-phycoerythrin and r-phycocyanin, whilst the products of
photosynthesis are floridean starch, floridoside and a material
known as mannoglycerate. This class, like the preceding one, differs
from the other algal classes in that the sugar residues have a
tendency to form i : 3 Hnkages. There is also a tendency to form
polysaccharide sulphate esters (cf. p. 448). Reproductive stages with
locomotor appendages are not known, even in the case of the male
body or spermatium. Whilst most simple members are filamentous,
all stages up to a complex thallus can be found, although there is
not quite the same degree of complexity as in the Phaeophyceae.
Despite variations in form there are only two basic types of thallus
construction. Protoplasmic connections exist between the cells of
all forms except most of those that comprise a small group, the
Proto-fiorideae (cf. p. 223). Sexual reproduction is oogamous, the
ovtmi being retained within the parent plant, and although the
subsequent development of the zygote varies to a certain extent, it
usually gives rise to filaments that bear special reproductive bodies
known as carpospores; these latter are normally responsible for the
production of a tetraspore-bearing diploid individual. There is a
great imiformity in reproduction throughout the class, and most
members also exhibit a regular alternation of generations.

Myxophycophyta

(4) Myxophyceae

The plants in this group show very little evidence of differentia-

tion and the cells contain only a simple form of nuclear material.
There is no true chromatophore, the pigments being distributed
throughout the entire peripheral portion of the cytoplasm, nor are
there any organs of locomotion, even among the reproductive
bodies. The products of photosynthesis are a specialized poly-
saccharide, myxophycean starch, and a proteinaceous material,
cyanophycin, present in granular form. The cell walls contain both
pectin and cellulose, the former sometimes forming a gelatinous
sheath. The colour of the cells is commonly blue-green, sometimes
olive green, the colour being due to the varying proportions of

8 THE ALGAE

chlorophyll a, c-phycoerythrin and c-phycocyanin. One interesting

feature is the complete lack, so far as is known, of sterols in this
class. There is no known sexual reproduction, propagation taking
place by simple division, by non-motile gonidia, by spores or else
by vegetative fragmentation. Members of the group are widely
distributed in marine and fresh waters and also occur terrestrially.
Some have the capacity to fix nitrogen from the atmosphere and
others have the capacity to live in quite hot water.

Chrysophycophyta

(5) Chrysophyceae

These form a primitive group, mostly of unicellular organisms,

in which the brown or orange colour of the chloroplasts is deter-
mined by excess of the carotinoid pigments. Most of the forms have
no cellulose cell wall and hence are *flagellates' in the old sense of
that term, but there are some members which do possess a cellulose
wall and hence are *algal' in the old sense of that term. Leucosin (a
proteinaceous substance) and oils are the usual forms of food
storage, whilst another marked feature is the silicified cysts or
statospores, which generally have a small aperture that is closed by
a special plug. The motile cells possess one, two or, more rarely,
three equal flagella attached at the front end, but in one sub-
section tiie two flagella are unequal in length. The most advanced
habit known is that of a branched algal filament, e.g. Phaeothamnion
(cf. p. 273), whilst the palmelloid types attain to a higher state of
differentiation, e.g. Hydrurus (cf. p. 273), than is commonly en-
coxmtered in either the majority of the palmelloid Chlorophyceae
or the Xanthophyceae. The occurrence of sexual reproduction is
uncertain and such records as there are point to simple isogamy.

(6) Xanthophyceae

The plants in this group are usually of a simple nature, but their
lines of morphological development frequently show an interesting
parallel or homoplasy with those observed in the Chlorophyceae
(cf. p. 325). The chloroplast is yellow-green in colour because of an
excess of /S-carotene, and starch is replaced by oil or leucosin as the
normal food storage material. The cell wall is frequently composed
of two equal or unequal halves overlapping one another. The
motile cells possess two unequal flagella arising from the anterior

CLASSIFICATION 9

end. Sometimes non-motile aplanospores are produced instead of

zoospores. Sexual reproduction is rare and when present is iso-
gamous, though in one genus ( Vaucheria) there is a well-developed
oogamy. This group is commonly regarded as having distinct
afiinities with the preceding group.

(7) Bacillariophyceae {Diatoms)

One of the characteristic features of these plants is their cell wall

which is composed partiy of sihca and partly of pectin. The wall is
always in two halves and frequently ornamented with deHcate
markings, which are so fine that microscope manufacturers make
use of them in order to determine the resolving power of the lenses.
Each cell contains one to many, variously shaped chromatophores,
which are yellow or golden brown, containing accessory xantho-
phylls in addition to the usual chlorophyll a, and /S-carotene. They
are divided into two major groups, the representatives of one
group being radially synmietrical, those of the other being bilater-
ally symmetrical. The presence of flagellate stages is highly prob-
able in the former whilst there is a special type of sexual fusion in
the latter group (cf p. 272). The relationship of this group to the
two preceding groups is not as distinct as it is between the Chryso-
phyceae and Xanthophyceae.

Pyrrophycophyta

(8) Cryptophyceae
Each cell commonly contains two large parietal chromatophores
of diverse colour, though frequendy of a brown shade, whilst the
product of photosynthesis is starch. The motile cells have two un-
equal ciHa and often possess a complex vacuolar system. Nearly
all the members have a *flagellate' organization and there are no
true filamentous algal forms. One type {Tetragonidium) (cf p. 275)
has been described as having a tendency towards the coccoid (non-
motile unicell with a cell wall) habit, but in general the class must
be regarded as the least *algal' like of all. Isogamy has been recorded
for one species.

(9) Dinophyceae

Most of the members of this class are motile unicells, but there
has been an evolutionary tendency towards a sedentary existence

10 THE ALGAE

and the development of short algal filaments, e.g. Dinothrix^ Dino-

clonium (cf. p. 277). Many are surrounded by a cellulose wall bearing
elaborate sculptured plates. Inside the cells there are dark yellow
or brown discoid chromatophores which contain a number of
xanthophylls not so far found in other algal classes. The products
of photosynthesis are starch and fat. The motile cells normally
possess two furrows, one transverse and one longitudinal, although
these may be absent in some of the more primitive members. A
transverse flagellum hes in the former, and the latter is the starting
point for the second flagellum which points backwards. Sexual
reproduction, if it occurs, is isogamous, but it has not been clearly
established in the few cases reported. Characteristic resting cysts
are also produced by many of the forms.

Nematophytales
(10) Nematophyceae

This is a fossil group of which one genus has been known for a
long time (Nematophyton), whilst the other (Nematothallus) has
only been described more recentiy. There is still considerable
doubt as to their true affinities, but it would seem that a place can
best be found for them as a very highly developed type of alga.
Their internal morphology suggests an alhance with more advanced
members of either the Chlorophyceae or Phaeophyceae. The only
reproduction so far recorded is that of spores which were developed
in tetrads.