Link to
"Seaweeds of Hawaii" by William H. Magruder and Jeffrey W. Hunt
Color = Blue Green - Rust - Black

Oscillatoria au natural |

Unknown Cyanobacterium from a warm spring near Mt. Lassen (CA) |
Pigmentation
Chlorophyll (a)
Carotenoids
Phycobillins
Diversity = Species 200 - 7500
Occurrence
Fresh Water -> Greatest Abundance
& Diversity
Salt Water
Planktonic: There are only a few species but they are an
important source for
fixed Nitrogen.
Reef
Lagoon: Lyngbya majuscula is the most common
Marine Species in Hawaii.
Rubble - Intertidal Zone in
"Ooid Sand Grains"
Terrestrial
Surface Crusts which appear
lifeless, are revived following significant rains. These dry to a
brittle crust as the
substrate looses water.
These can be ecologically significant because they add Biomass and Nitrogen to barren areas.
This is a long-term process,
however. These crusts can be seen in places like Arches
National Park (Utah) where it is very dry & there is
little ground cover or litter.
These can also be seen locally on abandoned,
dry lots. Heavy rains allow desiccated Cyanobacteria to revive. They form undulating
to round hydrated masses. These gradually shrink and crack as the substrate dries.
Some
Cyanobacteria (CBs) are also found in "green snow" which appears in the springtime on semi-permanent snowfields and glaciers.
Endolithic (Inside Rocks)
Cyanobacteria
have recently found in the most barren area of Antarctica where no other life has been found. Similar observations have been made
in the high Arctic. They can
live just below the surface of rocks. This sounds unbelievable but it is true. They may
also survive in the cold dry soil
of Antarctica. They become active when melt water appears.
Cyanobacteria
can also inhabit carbonatic
substrates like Limestone.
(http://bio.bu.edu/~golubic/marine-cyano.html).
Some
species inhabit the Calcium Carbonate secreted by Coralline Algae.
Cyanobacteria that live
in hot springs can deposit limestone therein. This accounts for
some of the various cascading "sculptures" that can be seen in areas of
Yellowstone Park. Some Marine species precipitate Calcium Carbonate (Limestone). Consequently, they become a part of the reef building biota.
 |
 |
| Hot
Spring at Yellowstone Park: The dark color is due to the presence of Cyanobacteria. |
Limestone
deposit at Yellowstone Park. The localized areas of green are due to the
presence of Cyanobacteria |
Stromatolites (Stromatoliths)
are large
columnar Calcium
Carbonate structures produced by Cyanobacteria. They accumulate slowly over immense time-spans. These were abundant in primeval times and constitute some of the oldest fossils. They can still be found
today, particularly in Australian Tidal Flats. They are multilayered deposits that form in shallow pools where the climate is warm. The
oldest Stromatolites are from the Precambrian and are 2.7
Billion Years-old!
Cyanobacteria are thought to be the oldest Oxygen
producing organisms. They profoundly changed the earth's atmosphere and allowed for the
evolution of organisms that can use oxygen for respiration.
Cyanobacteria use water (H20)
as an electron source and produce Oxygen. Other
photosynthetic bacteria use Hydrogen
Sulfide H2S as an electron donor
and produce Sulphur.

Fossil Cyanobacterium

Illustration showing Fossil Stromatolites |

Limestone deposit from ancient Cyanobacteria
(Glacier Park, Montana) |

Living Stromatolites on the Beach |

Stromatolites under water. |
Life
Modes

Most
species are Autotrophic
Some are Epiphytic or Epizooic (Polar
Bear)
Some are Symbiotic
Cyanobacteria are involved in several symbiotic
associations.
CBs can fix atmospheric N which
becomes available for its symbiotic partner.
Heterocysts are the cells
which fix nitrogen for most
Cyanobacteria.

Heterocyst flanked bt two Akinetes: Both have Thick Cell Walls |

Heterocysts of Anabaena isolated from Azolla. |
Lichens: Cyanobacteria
are frequently the Photosynthetic partners of Fungi that
compose Lichens. Lichens are
important pioneer organisms
which inhabit extremely difficult sites that may be dry, hot, cold, windy or all of the
above. The fixed nitrogen
supplied by Cyanobacteria allow
lichens to inhabit substrates that are nitrogen deficient.

Lichen with a Blue Color |

Lichen showing Cyanobacteria
inside |
Hornworts: (Anthocerophyta) have Endophytic (inside-plant) Cyanobacteria which reside in mucilage
filled chambers within
the thallus.

Hornwort (Anthocerophyta) |

Hornwort with Endophytic Cyanobacteria |
Hornworts are colonizers and inhabit wet, unstable sites. The ability to get Nitrogen from the CBs is
extremely advantageous because Nitrogen is usually a limiting element in terrestrial environments.
As they decay Hornworts release
their Nitrogen which becomes available for more complex plants. Species in
the Anthocerophyta can be seen near eroded and trampled areas in the local mountains.

Azolla: The aquatic
fern Azolla (Pterophyta) forms a symbiosis with Anabaena
species. These inhabit a cup-shaped area formed by the ventral leaves of the fern. The CBs fix nitrogen and release nitrogen-rich metabolites into the leaf cavity. These are absorbed by the Azolla which releases carbohydrates that are absorbed by the CBs.
It is almost impossible to separate the
symbiotic partners once they
become established.
Azolla can
form vegetative reproductive structure called Turions
which superficially resemble seeds. They form during drought or other unfavorable
conditions and consist of sclerotic leaves and a shoot apical meristem. They contain Anabaena. Consequently, the symbiosis is quickly established with new leaves that
develop when growing conditions improve.
This relationship has been used for centuries in rice cultivation because it provides a cheap, renewable and pollution-free source of fertilizer.

Cycads: (Cycadophyta) form an interesting symbiosis
with Nostoc. The CBs
live in a circular zone that
develops in upward growing root nodules. The nodules
develop a special tissue layer
that undergoes mitosis and autolysis in the presence of the CBs. The CBs occupy
this zone and fix Nitrogen.
Cycads occupy poor habitats and are almost extinct. Their symbiotic relationship
with CBs is one reason why they still can be found in nature.

Cycas |

Upward Growing Root Nodules |

Cross Section through a root nodule showing the dark zone that contains Cyanobacteria. |

Long Section through a Root Nodule: The dark areas contain Cyanobacteria. |

Commercial Cross Section showing the "Algal Zone" which contains CBs |

Thin section showing the Cyanobacteria in the "Algal Zone". |
Gunnera: (Anthophyta) is a genus of flowering
plants. These produce papillose outgrowths near the base of their enormous leaves. CBs are able
to colonize these areas and even
penetrate the cells in this
structure. At least one Gunnera species grows in Hawaii. One of Dr. Lamoureux's former students worked on these plants.
 |
 |
| Gunnera growing in Hawaii |
Growth
Forms
Single Cells
-> Colonies -> Filaments -> Branched Filaments

Synechococcus |

Crococcus |

Cells of Anacystis |

Anacystis Colony |

Nostoc Ball (Colony) |

Microscopic View of a Nostoc Colony |

Nostoc Filaments seen with Phase Microscopy |

Individual Noctoc Filaments from a large Colony (Phase Microscopy) |

Anabaena |

Oscillatoria |
 |

Oscillatoria Embossed |

Planktothrix sp. |

Lyngbya sp. |
 |
 |
Spirlina
sp. have a Spiral filament morphology but the
individual cells resemble those of Oscillatoria. |
Filaments may be
Branched, Unbranched or Clustered
Cell Division is not like that of Plants
Plasma Membrane & Inner 2 Wall Layers Invaginate
& form a Septum which grows Inward
and Separates the Cells.

|
 |
Electron Microscope Images of Dividing Cyanobacterial Cells
|
Intercellular Communication
Microplasmodesmata connect adjacent cells in
the filament.
Cell Shape Uniform -> Tapering -> Narrow Apical
Growth = General (No Apical Meristems)
Cell Types
Cyanobacteria are Prokaryotes.They have No Nuclear Envelope &
No
Membrane-Bound Organelles
Vegetative
Cells -> Photosynthesis
They have Photosynthetic Thylakoids which contain Chlorophyll a and the other photosynthetic pigments.
Thylakoids are Invaginations
of the Plasmalemma.
Cyanophycin Granules (CPG) contain storage Amino Acids.
Heterocysts have a Round
Shape & Thick Cell Walls. They often appear to be devoid of contents at the light microscope
level.
Heterocysts are the sites for the fixation of atmospheric Nitrogen
.

Vegetative cells are converted into Heterocysts.
An
Oxygen-free Anaerobic Environment
is Required because Oxygen inhibits Nitrogenase which is the
N-fixing enzyme.
The Thick Cell Walls are relatively impervious to Oxygen and this helps to create an anaerobic environment inside the Heterocyst.
The internal Membranes are thylakoids that have
lost Chlorophyll. They provide the sites for Nitrogenase.
Other cell
contents are generally lost. This helps to explain the lack of detail seen with the light Microscope.
| SEM
of Rivularia sp. showing the Pore (arrow) that connects the Heterocyst (H) to a Vegetative Cell (V) |
 |
Heterocysts are connected to the Vegetative Cells through a special pore in their end walls. These are much larger than the Microplasmodesmata. These Pores can be seen at
the light microscope level.

Locate the Pores in this Photo!!!!!
 |

TEM of an Akinete |

Larger Image of an Akinete |
Akinetes
Akinetes are Asexual Propagules.
These are
also derived from Vegetative Cells.
These tend to
be Elliptical in Shape.
They have
very Thick Walls.
Survive unfavorable Conditions
Many thylakoids can be seen with an Electron Microscope.
The Thylakoids are NOT
Organized for Photosynthesis.
They contain Storage Products
Cyanophyphtcin
Granules contain storage Amino Acids & are
noticeably present in resting cells.
Lipid Vacuoles can be seen.
Polyglucan Granules are present.

Cell Structure
Ultrastructure=Prokaryote

Size = Small
vs Eukaryotic Smallest = 1u

Complex Cell
Sheath & Wall

Sheath

Anabaena with a nearly translucent
Sheath: Identify the Round Smooth-looking Cell. |
 |
Some = Thin
Watery
Others -> Thick
& Fibrous
Composition
Acidic
Polysaccharides similar to Pectins
Inner
Layer is called the Glycocalyx
due to the presence of Glycoproteins. These are Proteins that contain Sugars as Sugar
Amides.A Glycocalyx is not present in all cases!
The Sheath Provides a Buffering Microenvironment.
It may absorb damaging Ultraviolet radiation.
Pili
Observed in Some
Species (Synechococus - Nostoc)
Extend from
Wall through Sheath
Tubular -
Protein Composition
60 nm x 1000
nm
Function
Unknown - Secretion?
Surface
Spines (Spinae)
Seen in some
Marine Forms (i.e. Synechococus)
Conical
& Project from Surface
Helical
Construction
Function ? Defense? ? Buoyancy?


Cell
Wall

Gram Negative Bacteria
Four Layers - (100 A each)

Layer
4 = Outer Membrane - Enzymes
Some are Digestive Enzymes.

Layer
3 = Electron Transparent

Layer
2 = Mucopolymer (Glycoproteins) Glycocalyx
Thicker
in Certain Species like Oscillatoria
sp.

Akinetes
-> This layer is Thicker.
Provides Protection from the Environment this aids in
Asexual Reproduction

Layer
1 = Electron Transparent
Immediately Outside Plasmalemma

Pores

Present in Transverse & Lateral Walls
Associated
with Gliding Movements
Function = Secretory?

Absence of Nuclear Envelope
Bacterial
"Chromosome" in the Center of Cell

Photosynthetic
Apparatus

Thylakoids
Invaginations of the Plasmalemma
Number - Responds to Light Intensity
Low light -> Many
High Light -> Few
Oscillatoria Thylakoids
= 20% Dry Weight
Principally Found - Peripheral Area of Cell
Various Appearances
Series of Layers
Undulated
Sac-Like
Tube-Like
Freeze Etch -> Reveals Surface Particles (Fluid Mosaic
Model)

Photosynthetic
Pigments

Principal
Light Harvesting Pigment Chlorophyll a

Accessory
Pigments

Carotenoids
B-Carotene & Zeaxanthin

Thylakoids have a Precise
Spacing
This is partly due to
the Phycobilisomes which are attached to the Surface of
the Thylakoids
Phycobilisomes contain Accessory Pigments for Photosynthesis
These are Water Soluble and are stabilized
by bonds to Proteins.

These include the
following Pigments.

C-Phycocyanins Absorb Green-Yellow Light (615-620A).

Allophycocyanins Absorb Orange-Red (650-670A)

C-Phycoerythrin Absorbs Green Light (495-570)

The
Visible Spectrum

Light Reaching The Surface
of the Earth
Peak approx. 500 nm
Drops Off at Higher & Lower
Wavelengths

Chlorophyll
& Carotenoids

Broad Absorbency
in the Blue
Drops Off in Green
Good Absorbency in Red Light
Poor Absorbency at 550-650 nm

Phycobillin
Pigments

Phycoerythrin
Good Absorbency 500 - 600 nm
Phycocyanin
Good Absorbency 550 - 650 nm
Allophycocyanin
Good Absorbency 600 - 675

If we combine all of these, including Chlorophyll
there is good light absorption across the Visible Spectrum!

Good
Coverage except at 500nm

Absorption
of Light by Water

Red
& Blue -> Preferentially Absorbed
Wavelengths
500 - 600nm Absorbed Least
This region
of the Spectrum corresponds with
Absorption Spectra of Phycobillins.

Consequently, organisms with these pigments are
better adapted for
life in an aquatic environment
& can exploit greater depths in lakes and oceans!

Gas
Vacuoles

These are common in Bacteria
Cyanobacteria
Common in Nature but Lost in Culture.
Composition
Membrane
Bound
Proteinaceous Walls (10% Cells' Protein)
Long Cylinders
Conical Ends
Regulates Physical Density of
individual cells NOT cells/volume
Vesicles can Fill with Gas or Collapse
Controls
Position in Water Column
Regulated by Photosynthesis
Ecological Significance
In Cyanobacterial
Algal Blooms the Gas Vacuoles are NOT Regulated.
CB Float to
Surface -> Massive Reproduction ->Secrete Toxins ->
Kill other Organisms

Aerial View of a Large Cyanobacterial Bloom |

Cyanophyte Bloom |

Microcystis Bloom |
 |
Nitrogen
Fixation

Nostocaceae with Heterocysts
fix Nitrogen
Marine Species with Heterocysts
Heterocysts
These have enlarged Thick Walls
The outer envelope is Bilayered.
Outer Layer contains
mostly Polysaccharide.
Inner Layer is
composed of Glycolipids.
Internal Membranes have
a
concentric to Reticulate Pattern
Nitrogen Fixation = Anaerobic
Nitrogenase is the enzyme
that fixes Nitrogen. It is very
sensitive to Oxygen which inhibits
N-fixation.
Thick Cell
Walls prevent Oxygen from entering Heterocysts.
Heterocysts lack
Photosystem II which is the Oxygen Releasing
Step.
Nitrogen levels in the environment regulate Heterocyst production.
Low N levels stimulate
heterocyst formation.

NonHeterocystic N-Fixation
Oscillatoria
(Planktonic)
Fix
N - Low Oxygen (Paerl &
Bebout 1988)
Associated
with Gliding & Rotating Movement
(10 microns/Second)
Secretion
occurs through Pores.
This produces waves of Mucilage.

Reproduction
Fragmentation
Hormogonia = Gliding Fragments
Akinetes -> 
Some Species
Enlarged
Thick Walls
Resist Heat - Drought
& Cold

Ecological
Roles

Terrestrial
Environments
Pioneering
Organisms contribute fixed nitrogen & some biomass. Nostoc
balls and crusts can grow on bare soil including Volcanic Sites.
They may
be Free Living
They can
be Epiphytic and even
Epizooic.
Their Symbiotic relationships with Lichens is ecologically important,
especially in the Arctic..
Other
symbiotic relationships are significant for the plants which have them.
Azolla can carpet the surface of small
lakes and ponds. This can be a major,
local ecological effect. This can lead to eutrophication!
The
symbiosis with Hornworts (Anthocerophyta) may have ecological
significance in Hawaii because Hornworts colonize disturbed
sites like pig runs and runoff streams.
Fresh
Water
Cyanobacteria
can grow in thermal Hot Springs
where algal photosynthetic organisms can't survive.
They deposit limestone in the process. However,
this is an extremely slow process.
They can grow in low light environments due to their Phycobillin pigments.
They can expolit habitats that Green Algae can not.
They release Carbon & Nitrogen Metabolites
which can stimulate the growth
of other organisms.
They can grow at low Oxygen levels and may enrich the local
Oxygen concentration because the produce O2.