10 Algae:

Photosynthetic Protists
Phycology or algology is the study of algae. The word phycology is derived from
the Greek phykos, meaning seaweed. The term algae [singular, alga] was originally
used to define simple “aquatic plants.” As noted above, it no longer has any formal
significance in classification schemes. Instead the algae can be described as
eukaryotic organisms that have chlorophyll a and carry out oxygen-producing
photosynthesis. They differ from other photosynthetic eukaryotes in lacking a well-
organized vascular conducting system and in having very simple reproductive
structures. In sexual reproduction the whole organism may serve as a gamete;
unicellular structures (gametangia) may produce gametes; or gametes can be
formed by multicellular gametangia in which every cell is fertile. Unlike the case with
plants, algal gametangia do not have nonfertile cells.

10.1 Distribution of Algae

Algae most commonly occur in water (fresh, marine, or brackish) in which they may
be suspended (planktonic) or attached and living on the bottom (benthic). A few
algae live at the water-atmosphere interface and are termed neustonic. Plankton
[Greek plankos, wandering] consists of free-floating, mostly microscopic aquatic
organisms. Phytoplankton is made up of algae and small plants, whereas
zooplankton consists of animals and nonphotosynthetic protists. Some algae grow
on moist rocks, wood, trees, and on the surface of moist soil. Algae also live as
endosymbionts in various protozoa, mollusks, worms, and corals. Several algae grow
as endosymbionts within plants, some are attached to the surface of various
structures, and a few lead a parasitic existence. Algae also associate with fungi to
form lichens.

10.2 Classification of Algae

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According to the five-kingdom system of Whittaker, the algae belong to seven
divisions distributed between two different kingdoms (Table 26.1). This classical
classification is based on cellular, not organismal, properties. Some more important
properties include: (1) cell wall (if present) chemistry and morphology; (2) form in
which food or assimilatory products of photosynthesis are stored; (3) chlorophyll
molecules and accessory pigments that contribute to photosynthesis; (4) flagella
number and the location of their insertion in motile cells; (5) morphology of the cells
and/or body (thallus); (6) habitat; (7) reproductive structures; and (8) life history
patterns. Based on these properties the algae are arranged by divisions in Table
10.1.

Table 10.1: Classical Classification of Algaea

___________________________________________________________________
Division (Common Name) Kingdom
___________________________________________________________________
Chrysophyta (yellow-green and Protista (single cell or colonial; eukaryotic)
golden-brown algae; diatoms)
Euglenophyta (photosynthetic Protista
euglenoid flagellates)
Pyrrhophyta (dinoflagellates) Protista
Charophyta (stoneworts) Protista
Chlorophyta (green algae) Protista
Phaeophyta (brown algae) Plantae (multicellular; eukaryotic)
Rhodophyta (red algae) Plantae
___________________________________________________________________
a
Five-kingdom system.

10.3 Ultrastructure of the Algal Cell

The eukaryotic algal cell (Figure 10.1) is surrounded by a thin, rigid cell wall. Some
algae have an outer matrix lying outside the cell wall. This usually is flexible and
gelatinous, similar to bacterial capsules. When present, the flagella are the locomotor
organelles. The nucleus has a typical nuclear envelope with pores; within the nucleus
are a nucleolus, chromatin, and karyolymph. The chloroplasts have membrane-
bound sacs called thylakoids that carry out the light reactions of photosynthesis.
These organelles are embedded in the stroma where the dark reactions of carbon
dioxide fixation take place. A dense proteinaceous area, the pyrenoid that is
associated with synthesis and storage of starch may be present in the chloroplasts.

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Mitochondrial structure varies greatly in the algae. Some algae (euglenoids) have
discoid cristae; some, lamellar cristae (green and red algae); and the remaining,
(golden-brown and yellow-green, brown, and diatoms) have tubular cristae.

Figure 10.1 Algal Morphology. Schematic drawing of a typical eucaryotic algal cell showing
some of its organelles and other structures.

10.4 Algal Nutrition

Algae can be either autotrophic or heterotrophic. Most are photoautotrophic; they
require only light and CO2 as their principal source of energy and carbon.
Chemoheterotrophic algae require external organic compounds as carbon and energy
sources.

10.5 Structure of the Algal Thallus

(Vegetative Form)
The vegetative body of algae is called the thallus [plural, thalli]. It varies from the
relative simplicity of a single cell to the more striking complexity of multicellular
forms, such as the giant kelps. Single-celled algae may be as small as bacteria,
whereas kelp can attain a size over 75 m in length. Algae are unicellular (Figure
10.2a,b,g), colonial (Figure 10.2c), filamentous (Figure 10.2d), membranous and
bladelike (Figure 10.2e), or tubular (Figure 10.2f).

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Figure 10.2: Diagrammatic Algal Bodies: (a) unicellular, motile, Cryptomonas; (b) unicellular,
nonmotile, Palmellopsis; (c) colonial, Gonium; (d) filamentous, Spirotaenia; (e) bladelike kelp,
Monostroma; (f) leafy tubular axis, branched tufts or plumes, Stigeoclonium; (g) unicellular,
nonmotile, Chrysocapsa.

10.6 Algal Reproduction

Some unicellular algae reproduce asexually. In this kind of reproduction, gametes do
not fuse to form a zygote. There are three basic types of asexual reproduction:
fragmentation, spores, and binary fission. In fragmentation the thallus breaks up
and each fragmented part grows to form a new thallus. Spores can be formed in
ordinary vegetative cells or in specialized structures termed sporangia [singular,
sporangium; Greek spora, seed, and angeion, vessel]. Flagellated motile spores are

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called zoospores. Nonmotile spores produced by sporangia are termed
aplanospores. In some unicellular algae binary fission occurs (nuclear division
followed by division of the cytoplasm).
Other algae reproduce sexually. Eggs are formed within relatively unmodified
vegetative cells called oogonia [singular, oogonium] that function as female
structures. Sperm are produced in special male reproductive structures called
antheridia [singular, antheridium]. In sexual reproduction these gametes fuse to
produce a diploid zygote.

10.7 Characteristics of the Algal Divisions

10.7.1 Chlorophyta (Green Algae)
The Chlorophyta or green algae [Greek chloros, green] are an extremely varied
division. They grow in fresh and salt water, in soil, on other organisms, and within
other organisms. The Chlorophyta have chlorophylls a and b along with specific
carotenoids, and they store carbohydrates as starch. Many have cell walls of
cellulose. They exhibit a wide diversity of body forms, ranging from unicellular to
colonial, filamentous, membranous or sheet-like, and tubular types (Figure 10.3).
Some species have a holdfast structure that anchors them to the substratum. Both
asexual and sexual reproductions occur in green algae.

Figure 10.3: Chlorophyta (Green Algae); Light Micrographs. (a) Chlorella, a unicellular
nonmotile green alga (x160). (b) Volvox, a typical green algal colony (x450). (c) Spirogyra, a

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filamentous green alga (x100). Four filaments are shown. Note the ribbon-like, spiral
chloroplasts within each filament. (d) Ulva, commonly called sea lettuce, has a leafy
appearance. (e) Acetabularia, the mermaid’s wine goblet. (f) Micrasterias, a large desmid
(x150).
10.7.2 Euglenophyta (Euglenoids)
The euglenoids share with the Chlorophyta and Charophyta the presence of
chlorophylls a and b in their chloroplasts. The primary storage product is paramylon
(a polysaccharide composed of -1,3 linked glucose molecules), which is unique to
euglenoids. They occur in fresh, brackish, and marine waters and on moist soils;
they often form water blooms in ponds and cattle water tanks. The representative
genus is Euglena (Figure 10.4).

Figure 10.4: Euglena. A diagram illustrating the principal structures found in this euglenoid.
Notice that a short second flagellum does not emerge from the anterior invagination. In some
euglenoids both flagella are emergent.

10.7.3 Chrysophyta (Golden-Brown and Yellow-Green Algae;

Diatoms)
The division Chrysophyta is quite diversified with respect to pigment composition,
cell wall, and type of flagellated cells. The division is divided into three major
classes: golden-brown algae [Greek chrysos, gold], yellow-green algae, and diatoms.
The major photosynthetic pigments are usually chlorophylls a and c1/c2, and the
carotenoid fucoxanthin. When fucoxanthin is the dominant pigment, the cells have a
golden-brown color. The major carbohydrate reserve in the Chrysophyta is
chrysolaminarin (a polysaccharide storage product composed principally of -1,3
linked glucose residues). Some Chrysophyta lack cell walls; others have intricately
patterned coverings external to the plasma membrane, such as scales (Figure
10.5a), walls, and plates. Diatoms have a distinctive two-piece wall of silica, called a
frustule. Two anteriorly attached flagella of unequal length are common among

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Chrysophyta (Figure 10.5b), but some species have no flagella, and others have
either one flagellum or two that are of equal length. Most Chrysophyta are unicellular
or colonial.

The diatoms (Figure 10.5c,d) are photosynthetic, circular or oblong chrysophyte

cells with frustules composed of two halves or thecae that overlap like a petri dish
[therefore their name is from the Greek diatomsos, cut in two]. The larger half is the
epitheca, and the smaller half is the hypotheca. Diatoms grow in freshwater, salt
water, and moist soil and comprise a large part of the phytoplankton. Diatom
frustules are composed of crystallized silica [Si(OH) 4] with very fine markings
(Figure 10.5c,d). They have distinctive, and often exceptionally beautiful, patterns
that are different for each species. Frustule morphology is very useful in diatom
identification.

Figure 10.5: Chrysophyta (Yellow-Green and Golden-Brown Algae; Diatoms). (a) Scanning
electron micrograph of Mallomonas, a chrysophyte, showing its silica scales. The scales are
embedded in the pectin wall but synthesized within the Golgi apparatus and transported to the
cell surface in vesicles (x9,000). (b) Ochromonas, a unicellular chrysophyte. Diagram showing

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typical cell structure. (c) Scanning electron micrograph of a diatom, Cyclotella meneghiniana
(x750). (d) Assorted diatoms as arranged by a light microscopist (x900).
10.7.4 Phaeophyta (Brown Algae)
The Phaeophyta or brown algae [Greek phaeo, brown] consist of multicellular
organisms that occur almost exclusively in the sea. Some species have the largest
linear dimensions (length) known in the eukaryotic world. The simplest brown algae
consist of small openly branched filaments; the larger, more advanced species have
a complex arrangement. Some large kelps are conspicuously differentiated into
flattened blades, stalks, and holdfast organs that anchor them to rocks (Figure
10.6). Some, such as Sargassum, form huge floating masses that dominate the
Sargasso Sea. The color of these algae reflects the presence of the brown pigment
fucoxanthin, in addition to chlorophylls a and c, -carotene, and violaxanthin. The
main storage product is laminarin, which is quite similar in structure to
chrysolaminarin.

Figure 10.6: Phaeophyta (Brown Algae). Diagram of the parts of the brown alga, Nereocystis.
Due to the holdfast organ, the heaviest tidal action and surf seldom dislodge brown algae from
their substratum. The stipe is a stalk that varies in length; the bladder is a gas-filled float.

10.7.5 Rhodophyta (Red Algae)

The division Rhodophyta, the red algae [Greek rhodon, rose], includes most of the
seaweeds (Figure 26.8). A few reds are unicellular but most are filamentous and
multicellular. Some red algae are up to 1 m long. The stored food is the
carbohydrate called floridean starch (composed of -1,4 and -1,6 linked glucose
residues). The red algae contain the red pigment phycoerythrin, one of the two types
of phycobilins that they possess. The other accessory pigment is the blue pigment
phycocyanin. The presence of these pigments explains how the red algae can live at

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depths of 100 m or more. The wavelengths of light (green, violet, and blue) that
penetrate these depths are not absorbed by chlorophyll a but instead by these
phycobilins. Not surprisingly the concentration of these pigments often increases
with depth as light intensity decreases. The phycobilins, after absorbing the light
energy, pass it on to chlorophyll a.

The cell walls of most red algae include a rigid inner part composed of microfibrils
and a mucilaginous matrix. The matrix is composed of sulfated polymers of galactose
called agar, funori, porphysan, and carrageenan. These four polymers give the red
algae their flexible, slippery texture. Agar is used extensively in the laboratory as a
culture medium component. Many red algae also deposit calcium carbonate in their
cell walls and play an important role in building coral reefs.

Figure 10.7: Rhodophyta (Red Algae). These algae (e.g., Gelidium latifolium) are much
smaller and more delicate than the brown algae.

10.7.6 Pyrrhophyta (Dinoflagellates)

The Pyrrhophyta or dinoflagellates are unicellular, photosynthetic algae. Most
dinoflagellates are marine, but some live in fresh water. Along with the chrysophytes
and diatoms, the dinoflagellates make up a large part of freshwater and marine
plankton and are at the base of many food chains. Species of Noctiluca, Pyrodinium,
Gonyaulax, and other genera can produce light and are responsible for much of the
luminescence (phosphorescence) seen in ocean waters at night. Sometimes
dinoflagellate populations reach such high levels that poisonous red tides result.

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Most dinoflagellates have two flagella. The longitudinal flagellum (undilipodium)
extends posteriorly as a rudder; the flattened, ribbon-like transverse flagellum
propels the cell forward while causing it to spin (Figure 10.8). Due to this spinning,
the dinoflagellates received their name from the Greek dinein, to whirl. Most
dinoflagellates have chlorophylls a and c, in addition to carotenoids and xanthophylls.

Figure 10.8: Pyrrhophyta (Dinoflagellates). Diagram of the parts of the brown alga,
dinoflagellate, Ceratium.

10.8 Ecological Importance of Algae

10.8.1 Benefits of Algae
10.8.1.1 Primary Produces
Algae are primary produces, which play an important part of any aquatic food chain
because they fix carbon dioxide into organic molecules that can be consumed by
chemohetrotrophs. Algae account for 50-70% of global photosynthesis.
Phytoplanktons form the base of the ocean food web; especially important are the
diatoms and dinoflagellates.

10.8.1.2 Underwater Forests

Seaweeds are not only food, but shelters for aquatic organisms - especially
important are the kelps, which form underwater forests such as Sargasso Sea
community developed due to growth of a brown alga, Sargassum.

10.8.1.3 Reef Builders

Many of the coralline red algae, which secrete calcium carbonate and play a major
role in building coral reefs in tropical seas.

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10.8.1.4 Symbiosis
Lichens are algae and fungi symbiosis.

10.8.2 Harmful Algae

10.8.2.1 Water Quality
A bad taste to water occurs when certain algae are present in large numbers.

10.8.2.2 Clogging
When water is polluted with nutrients such as fertilizer or sewage, algae grow in
excess and cause clogging of waterways, streams, and filters.

10.8.2.3 Blooms
Freshwater quality strongly influenced by presence of algae. Seasonal changes in
nutrients, light, and temperature cause fluctuation in algal population; periodic
increase in number of planktonic algae are called blooms.

10.8.2.4 Red Tides

Red tides are toxic algal blooms caused by certain species of dinoflagellates. Large
concentrations of Gonyaulax give the ocean a deep red colour, from which the name
red tide originates.

10.8.2.5 Poisoning
Algae can be harmful to humans. Some algae produce toxins that become
concentrated in shellfish and finfish. Although they have little effect on shellfish and
finfish, the toxins accumulate in the seafood flesh, making it unsafe or poisonous for
human consumption. The dinoflagellates (class Dinophyceae) are the most notorious
producers of toxins. For example, paralytic shellfish poisoning is caused by saxitoxin
or any of at least 12 related compounds. Saxitoxin is probably the most toxic
compound known and is 100,000 times more toxic than cocaine. Saxitoxin and
saxitoxin-like compounds are nerve toxins that interfere with neuromuscular
junctions. Alexandrium tamarense and Gymnodinium catenatum are the two species
most often associated with paralytic shellfish poisoning.

10.8.2.6 Human Diseases

Algae can cause human diseases by directly attacking human tissues, although the
frequency is rare. Protothecosis is caused by a chloroplast-lacking green alga,
Prototheca. The alga infects skin lesions, grows subcutaneously, and can eventually

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spread to the lymph glands. Similar infections, in humans and cattle, are caused by
chloroplast-bearing species of Chlorella (a member of Chlorophyta).

10.9 Commercial Importance of Algae

10.9.1 Human Foods
Some red algae, brown algae and green algae, are eaten by humans. Approximately
500 species are eaten by humans, and some 160 are commercially important.
(1) The red alga Porphyra is the most important commercial food alga.
(2) Palmaria palmata, another red alga, is eaten in the North Atlantic region.
(3) Laminaria species (brown algae) is eaten with meat or fish and in soups.
(4) The green algae Monostroma and Ulva look like lettuce leaves (their common
name is sea lettuce), and they are eaten as salads or in soups, relishes, and
meat or fish dishes.
(5) The microscopic, freshwater green alga Chlorella is cultivated and eaten in
Taiwan, Japan, Malaysia, and the Philippines. It has a high protein content (53–
65%) and has been considered as a possible food source during extended space
travel.

10.9.2 Food Additives

The cell walls of many types of seaweed contain phycocolloids that have received
increasing use in prepared foods. The three major phycocolloids are alginates, agars,
and carrageenans. Alginates are extracted primarily from brown seaweeds, and agar
and carrageenan are extracted from red seaweeds. Phycocolloids are safely
consumed by humans and other animals and are therefore used in a wide variety of
prepared foods, such as “ready-mix” cakes, “instant” puddings and pie fillings, and
artificial dairy toppings.
(1) Alginates, or alginic acids, are commercially extracted from brown seaweeds,
especially the kelp Macrocystis, Laminaria, and Ascophyllum. Alginates are used
in ice creams to limit ice crystal formation, thereby producing a smooth texture,
and are also used as emulsifiers and thickeners in syrups and as fillers in candy
bars and salad dressings.
(2) Agars are extracted primarily from species of the red alga Gelidium, but they
are also obtained from other red algae, especially Gracilaria, Pterocladia,
Acanthopeltis, and Ahnfeltia. Agars are used in instant pie fillings, canned
meats or fish, and bakery icings. Agar is also used as a clarifying agent in beer
and wine.

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(3) Carrageenan, from the Irish word “carraigin” (meaning Irish moss), are
extracted from various red algae: Eucheuma in the Philippines, Chondrus
crispus in the United States and the Canadian Maritime Provinces, and Iridaea
in Chile. Carrageenans are used as thickening and stabilizing agents in dairy
products, imitation creams, puddings, syrups, and canned pet foods.

10.9.3 Active Additives in Medical Drugs or Insecticides

Phycocolloids have industrial uses in addition to their important roles in food
products. Because they are relatively inert and have good gelling properties, they are
used as creams and gels for carrying minute amounts of active additives, as in
medical drugs or insecticides.
(1) Agar is used extensively as a bacteriologic culturing substrate in medical and
research facilities and is also used as a substrate for eukaryotic cell and tissue
culture, including the culture of algae themselves.
(2) Carrageenans are used in the manufacture of shampoos, cosmetics, and
medicines.

10.9.4 Diatomaceous Earth or Diatomite

The frustules, or cell walls, of diatoms (Chrysophyta) are deposited in ocean
sediments, and the fossilized remains are called diatomaceous earth or diatomite.
Diatomite contains approximately 50 million diatom frustules per cubic inch.
Diatomite is relatively inert and contains many fine pores. It consists of
approximately 90% silica, and the remainder consists of compounds such as
aluminium and iron oxides. It has a high absorptive capacity, large surface area, low
bulk density, and relatively low abrasion.
(1) The fine pores in the frustules make it an excellent filtering material for
beverages, chemicals, industrial oils, cooking oils, sugars, water supplies,
varnishes, lacquers, jet fuels, and antibiotics.
(2) The low abrasive properties make it suitable for use in toothpaste,
“nonabrasive” cleansers, polishes, and buffing compounds.
(3) Diatomite is widely used as a filler and extender in paint, paper, and rubber and
plastic products; the gloss and sheen of ‘flat’ paints can be controlled using
diatomite.

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(4) Because diatomite can absorb approximately 2.5 times its weight in water, it
also makes an excellent anti-caking carrier for powders used to dust roses or
for cleansers used to clean rugs.
(5) Diatomite is also used in making welding rods, battery boxes, concrete,
explosives, and animal foods.

10.9.5 Fertilizer
Seaweeds have been used as agricultural fertilizers for centuries in many parts of
the world. Coastal farmers cut seaweeds that were spread over the soil. Kelp is now
used to extract macronutrients and micronutrients for specialized plant fertilizers and
animal feed supplements. Dried kelp is almost 50% mineral matter; Ascophyllum
nodosum, for example, contains 55 trace elements.

10.9.6 Industrial Chemicals

Algae are used for production or extraction of some important chemical that have
wide industrial values.
(1) Iodine can be extracted from brown algae.
(2) The green unicellular flagellate Dunaliella is cultivated in saline ponds. The
culture conditions are manipulated so that carotene or glycerol is produced in
large amounts. These compounds are extracted and sold commercially.

10.9.7 Forensic Medicine

Diatoms have been used in forensic medicine. Where death by drowning is
suspected, lung tissue is examined. The presence of silica diatom cell walls can verify
death by drowning; in mysterious cases, the diatom species can be used to pinpoint
the exact location of death because the species are characteristic for a given lake,
bog, or bay.

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References
1. Foundations in Microbiology, Eighth Edition. 2011. Kathleen Park Talaro and Arthur Talaro. The
McGraw−Hill Companies, Inc., New York.
2. Prescott's Microbiology, Eighth Edition. 2010. Joanne Willey, Linda Sherwood and Chris
Woolverton. The McGraw−Hill Companies, New York.

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