Mucilages and Sheaths

These are general terms for some sort of outer gelatinous covering present in both
prokaryotic and
eukaryotic algae. Mucilages are always present and we can observe a degree of
development of
a sheath that is associated with the type of the substrate the cells contact (Figure 2.2). All
cyanobacteria
secrete a gelatinous material, which, in most species, tends to accumulate around the
cells or trichome in the form of an envelope or sheath. Coccoid species are thus held
together to
form colonies; in some filamentous species, the sheath may function in a similar manner, as
in
the formation of Nostoc balls, or in development of the firm, gelatinous emispherical domes
of
the marine Phormidium crosbyanum. Most commonly, the sheath material in filamentous
species
forms a thick coating or tube through which motile trichomes move readily. Sheath
production is a
continuous process in cyanobacteria, and variation in this investment may reflect different
physiological
stages or levels of adaptation to the environment. Under some environmental conditions the
sheath may become pigmented, although it is ordinarily colorless and transparent. Ferric
hydroxide
or other iron or metallic salts may accumulate in the sheath, as well as pigments originating
within
the cell. Only a few cyanobacterial exopolysaccharides have been defined structurally; the
sheath of
Nostoc commune contains cellulose-like glucan fibrils cross-linked with minor
monosaccharides,
and that of Mycrocystis flos-aquae consists mainly of galacturonic acid, with a composition
similar to that of pectin. Cyanobacterial sheaths appear as a major component of soil crusts
found throughout the world, from hot desert to polar regions, protecting soil from erosion,
favoring water retention and nutrient bio-mobilization, and affecting chemical weathering of
the
environment they colonize. In the Cyanophyta the cell wall lies between the plasma
membrane and the mucilaginous
sheath; the fine structure of the cell wall is of Gram-negative type. The innermost layer, the
electron-opaque layer or peptidoglycan layer, overlays the plasma membrane, and in most
cyanobacteria
its width varies between 1 and 10 nm, but can reach 200 nm in some Oscillatoria species.
Regularly arranged discontinuities are present in the peptidoglycan layer of many
cyanobacteria;
pores are located in single rows on either side of every cross wall, and are also uniformly
distributed
over the cell surface. The outer membrane of the cell wall appears as a double track
structure tightly
connected with the peptidoglycan layer; this membrane exhibits a number of evaginations
representing
sites of extrusion of material from the cytoplasm through the wall into the slime.
Cyanophyta and Prochlorophyta
The photosynthetic apparatus of these algae is localized on intracytoplasmic membranes
termed
thylakoids. The thylakoids membranes show considerable variations in structure and
arrangements
depending on the species. Also the amount of thylakoid membranes per cell is variable due
to
growth conditions and taxonomic specificity.
In most species the thylakoids are arranged peripherally in three to six layers running
parallel to
the cell membrane, forming an anastomosing network of concentric shells. This peripheral
region
has been called the chromatoplast, which is separated from the inner nucleoplasmic region
called
the centroplasm. This type of thylakoid arrangement is characteristic of many unicellular
and
filamentous cyanobacteria, such as Synechococcus planctibus and Anabaena sp. In some
other
organisms such as Oscillatoria and Arthrospira the thylakoids are orientated perpendicular
to
the longitudinal cell wall. Radial arrangement is present in Phormidium retzi. Thylakoids are
not always restricted to the cell periphery but can be found scattered throughout the cell as
in
Gloeotrichia sp. However, the arrangement of thylakoids can change from cell to cell and
cell
types (vegetative cells, heterocyst, akinete) within the same culture from parallel to a
convoluted
appearance. Thylakoids can fuse with each other and form an anastomosing network; unlike
the
chloroplasts of eukaryotic algae, the thylakoids of cyanobacteria form stacking regions only
to a
very limited extent. During cell division the thylakoids have to be separated and divided into
the
daughter cells; generally this division is an active process. Cell division starts by a
centripetal
growth of cytoplasmic membrane and peptidoglycan layer, to form the septum; the
thylakoid cylinder
is narrowed at the level of the cross wall by invagination before the septum is formed.
Proteins,
lipids, carotenoids, and chlorophyll a are major components of thylakoids. On the outer
surface of
the thylakoids regular rows of electron dense granular structures are closely attached.
These granules
termed phycobilisomes contain the light harvesting phycobiliproteins, that is,
allophycocyanin,
phycocyanin, and phycoerythrin (Figure 2.75). The phycobilisome structure consists of a
threecylinder
core of four stacked molecules of allophycocyanin, closest to the thylakoid membrane,
on which converge rod-shaped assemblies of coaxially stacked hexameric molecules of only
phycocyanin
or both phycocyanin and phycoerythrin (Figure 2.76). Phycobiliproteins are accessory
pigments for the operation of photosystem II also in Glaucophyta, Cryptophyta, and
Rhodophyta.

CULTURE PARAMETERS
A culture has three distinct components: a culture medium contained in a suitable vessel;
the algal
cells growing in the medium; air, to allow exchange of carbon dioxide between medium and
atmosphere.
For an entirely autotrophic alga, all that is needed for growth is light, CO 2, water, nutrients,
and
trace elements. By means of photosynthesis the alga will be able to synthesize all the
biochemical
compounds necessary for growth. Only a minority of algae is, however, entirely autotrophic;
many
are unable to synthesize certain biochemical compounds (certain vitamins, e.g.) and will
require
these to be present in the medium (obligate mixotropy condition, see Chapter 1).
The most important parameters regulating algal growth are nutrient quantity and quality,
light,
pH, turbulence, salinity, and temperature. The most optimal parameters as well as the
tolerated
ranges are species specific and the various factors may be interdependent and a parameter
that is
optimal for one set of conditions is not necessarily optimal for another.

Temperatures lower than 16ºC will slow down growth, whereas those higher than 35ºC are
lethal for a number of species.

BG11 Medium Composition

Reagents Per Liter
NaNO3 a 1.5 g
K2HPO4 * 3H2O 0.004 g
MgSO4 * 7H2O 0.075 g
CaCl2 * 2H2O 0.027 g
Citric acid (C6H8O7) 0.006 g
Ammonium ferric citrate (C6H8O7 * nFe * nNH3) 0.006 g
EDTANa2Mg 0.001 g
Na2CO3 0.02 g
Microelement stock solution 1 ml
Microelement Stock Solution
H3BO3 2.860 g
MnCl2 * 4H2O 1.810 g
ZnSO4 * 7H2O 0.222 g
Na2MoO4 * 2H2O 0.390 g
CuSO4 * 5H2O 0.079 g
Co(NO3)2 * 6H2O 0.0494 g
pH ј 7.4
aTo be omitted for N2-fixing cyanobacteria

Beijerinck Medium Composition

Reagents Per Liter
Stock I 100 ml
Stock II 40 ml
Stock III 60 ml
Micronutrients 1 ml
Stock I
NH4NO3 1.5 g
K2HPO4 0.2 g
MgSO4 * 7H2O 0.2 g
CaCl2 * 2H2O 0.1 g
Stock II
KH2PO4 9.07 g
Stock III
K2HPO4 11.61 g

Micronutrients
H3BO3 10 g
MnCl2 * 4H2O 5g
EDTA 50 g
CuSO4 * 5H2O 1.5 g
ZnSO4 * H2O 22 g
CoCl2 * 6H2O 1.5 g
FeSO4 * 7H2O 5g
(NH4)6Mo7O24 * 4H2O 1g
pH ј 6.8

Micronutrients
H3BO3 10 g
MnCl2 * 4H2O 5g
EDTA 50 g
CuSO4 * 5H2O 1.5 g
ZnSO4 * H2O 22 g
CoCl2 * 6H2O 1.5 g
FeSO4 * 7H2O 5g
(NH4)6Mo7O24 * 4H2O 1g
pH ј 6.8

(Pasteur Culture Collection of Cyanobacteria http://www.pasteur.fr/recherche/banques/PCC/)