WB01343_.gif (599 bytes)    Cyanophyta   WB01345_.gif (616 bytes)
Link to Dr. Smith's Web Page on Cyanophyta WB01436_.gif (236 bytes)

Link to "Seaweeds of Hawaii" by William H. Magruder and Jeffrey W. HuntWB01436_2.gif (236 bytes)

Color = Blue Green - Rust - Black

OscillatoriaHand.jpg (29244 bytes)
Oscillatoria au natural

Unknown Cyanobacterium from a warm spring near Mt. Lassen (CA)

PigmentationLyngbib200Lab.jpg (53960 bytes)

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 theCyanoMiCrust.jpg (91063 bytes)
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,NostocNodules240.jpg (23852 bytes)
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.

CyanoHyellaStella300Crop.jpg (30898 bytes)
Hyella stella: a Cyanobacterium that lives in Marine Limestone

ScytonemaEndolithicum.jpg (83799 bytes)
Scytonema endolithicum

Organisms like this can live in Coral Rubble which rolls about as "sand". Consequently, they inhabit a reef zone that would otherwise be unavailable to them, due to the absence of a stable substrate and the presence of vigorous wave action which would be unfavorable for Planktonic algae. These areas are often turbid due to wave & wind action and from soil runoff.

OoidGrain300Lab.jpg (38886 bytes) "Ooid" sand grain containing Endolithic Cyanobacterium (CB).

The presence of these organisms in limestone & coral reefs can lead to erosion. This is caused by grazing animals who eat the Cyanobacteria and consequently decrease the amount of limestone present. The effect of this is illustrated below.

IntertidalNotchHue400Lab.jpg (125092 bytes)
Erosion caused by Endolithic Cyanobacteria
Biokarst.jpg (33814 bytes)
This can sometimes produce fantastic "Biocarst" shapes.

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.

YellowstoneBGA.jpg (99488 bytes) YellowSyong.jpg (155888 bytes)
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) StromatolyteThinSection.jpg (19511 bytes)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.

CyanoFossil.jpg (19447 bytes)
Fossil Cyanobacterium

Stromatoliths400.jpg (75126 bytes)
Illustration showing Fossil Stromatolites
FossilBGA400.jpg (146811 bytes)
Limestone deposit from ancient Cyanobacteria

(Glacier Park, Montana)
StromatolytesBeach.jpg (92174 bytes)
Living Stromatolites on the Beach
ThromboheadsUnWater.jpg (32764 bytes)
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.

AkineteHeterocystLMHigh300Lab.jpg (15127 bytes)
Heterocyst flanked bt two Akinetes: Both have Thick Cell Walls
Heterocyst-3M600Lab.jpg (103891 bytes)
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.

BlueLichen.jpg (189449 bytes)
Lichen with a Blue Color
lichenCB.jpg (32934 bytes)
Lichen showing Cyanobacteria inside

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

AnthoGametoSporoOver500.jpg (257907 bytes)
Hornwort (Anthocerophyta)
Hornwortdendrocerosnostoc.jpg (73080 bytes)
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.

Azolla.jpg (118912 bytes)
Azolla Plants

(Dorsal View)
AzollaApexDorsal300Lab.jpg (95014 bytes)
Individual Azolla Plant showing its Main Axis and Branches
AzollaDorsalAnthApexLab.jpg (125424 bytes)

Azolla Shoot Apex

AzollaIsoLeafLab.jpg (56162 bytes)
Isolated Azolla Leaf: The dark area is due to the presence of Anabaena inside a pouch formed by the Leaf.

Section through an Azolla Leaf showing the chamber in which the N-fixing Cyanobacteria reside.
Heterocysts-2-LMagLab.jpg (96683 bytes)
Anabaena isolated from an Azolla Leaf: note the Heterocysts and Photosynthetic Cells.

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.

CycafMegaStrobOver300.jpg (136958 bytes)
Cycas
CoralloidRootsLargeNature.jpg (75052 bytes)
Upward Growing Root Nodules
CycadNoduleX-S300.jpg (70625 bytes)
Cross Section through a root nodule showing the dark zone that contains Cyanobacteria.
CycadNodLSMacro400.jpg (83941 bytes)
Long Section through a Root Nodule: The dark areas contain Cyanobacteria.
NodAZCycasXSCommercLowMagLab.jpg (152965 bytes)
Commercial Cross Section showing the "Algal Zone" which contains CBs
AZLMSectTolBlLab.jpg (136109 bytes)
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.

GunneraLeavesPeople.jpg (285570 bytes) GunneraPetioles.jpg (244125 bytes)
Gunnera growing in Hawaii

Growth Forms

Single Cells -> Colonies -> Filaments -> Branched Filaments

Synechococcus.jpg (42395 bytes)
Synechococcus
ChrococcusNihonCrop.jpg (23977 bytes)
Crococcus
Anacystis.jpg (44283 bytes)
Cells of Anacystis
Anacystir.jpg (53631 bytes)
Anacystis Colony
NostocNodulesCropHueLab.jpg (33078 bytes)
Nostoc Ball (Colony)
NostoColony.jpg (184307 bytes)
Microscopic View of a Nostoc Colony
NostocNihon500.jpg (64936 bytes)
Nostoc Filaments seen with Phase Microscopy
NostocFilamentsPhase240.jpg (80012 bytes)
Individual Noctoc Filaments from a large Colony
(Phase Microscopy)
AnabaenaBest.jpg (30865 bytes)
Anabaena
OscilatoriaLongNihon400Ctrast.jpg (30106 bytes)
Oscillatoria
OscillatoriaColony240Lab.jpg (74474 bytes) OscillatoriaEmbos.jpg (71073 bytes)
Oscillatoria
Embossed
Planktothrix.jpg (31457 bytes)
Planktothrix sp.
Lyngbia.jpg (55196 bytes)
Lyngbya sp.
SpirulinaB.jpg (4316 bytes) SpirulinaClose300.jpg (27345 bytes)
Spirlina sp. have a Spiral filament morphology but the
individual cells resemble those of Oscillatoria.

Filaments may be Branched, Unbranched or Clustered

Calothrx.jpg (34830 bytes)
Calothrix
GleotrichiaHue.jpg (51065 bytes)Gleotrichia Individuals
GleotrichiaColony240.jpg (15496 bytes)

Gleotrichia Colony


GloetricLab.jpg (44961 bytes)
Tolypothrix.jpg (27979 bytes)
Tolypthrix

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.

CellInvaginationLab.jpg (47388 bytes)

CBDivideTEM.jpg (97978 bytes)

Electron Microscope Images of Dividing Cyanobacterial Cells

Intercellular Communication

Microplasmodesmata connect adjacent cells inMicplasmLab.jpg (24624 bytes) 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

Typbgaem.jpg (60326 bytes)     

CBDivideThylakoidsTEM.jpg (102893 bytes) CBTEMThylakoidsLab.jpg (82662 bytes)

They have Photosynthetic Thylakoids which contain Chlorophyll a and the other photosynthetic pigments.

Thylakoids are Invaginations of the Plasmalemma.AnabaenaBest300Lab.jpg (66267 bytes)

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.

Heterocyst-1-600L.jpg (116469 bytes)
Heterocyst & Photosynthetic cells from Anabaena seen with Standard Optics.

Heterocyst-1-600EmbCropLab.jpg (83569 bytes)
Embossed image of Anabaena showing the Thick Walls and Pores of a Heterocyst
NostocNihonCropHet.jpg (20294 bytes)
Heterocyst as seen with Phase Optics
Heterocysts seen with different types of Light Microscopy
Heterocyst.jpg (60034 bytes)
Heterocyst seen with an Electron Microscope

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) RivulariaSEMConnectHeteroVeg.jpg (27676 bytes)

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.

AnabaenaHetero400MLab.jpg (45316 bytes)
Locate the Pores in this Photo!!!!!

AkineteHeterocystLMHigh300Lab.jpg (15127 bytes) AkineteEM400Lab.jpg (22711 bytes)
TEM of an Akinete
AkineteEMBlue400.jpg (133744 bytes)
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

AnabaenaSheathHCystSheathLab.jpg (39461 bytes)
Anabaena with a nearly translucent Sheath: Identify the Round Smooth-looking Cell.
SolentiaAchromaticaSheathLab250.jpg (55584 bytes)

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?

Wall-1-300Lab.jpg (46161 bytes)

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

ThylakoidsThylakoidsAnabaena300Lab.jpg (42378 bytes)

Invaginations of the Plasmalemma

Number - Responds to Light Intensity

Low light -> Many

High Light -> FewThylakoids.jpg (46140 bytes)

Oscillatoria Thylakoids = 20% Dry Weight

Principally Found - Peripheral Area of Cell

Various Appearances

Series of LayersThylakoidsNostocLab.jpg (29208 bytes)

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 Phycobilisomes300Lab.jpg (31193 bytes)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.

 

 

phycobilisome.jpg (28474 bytes)

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

SpectrumLab500.jpg (106148 bytes)

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!

SpectrumLab500Combined.jpg (86828 bytes)

Good Coverage except at 500nm

Absorption of Light by Water

SpectrumWaterDepth450Lab.jpg (57961 bytes)

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 BacteriaGasVacuoleCell240Lab.jpg (93521 bytes)

Cyanobacteria

Common in Nature but Lost in Culture.

Composition

Membrane Bound

Proteinaceous Walls (10% Cells' Protein)

Long CylindersGasVesicleLab.jpg (66229 bytes)

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

BloomMicrocystsiAir.jpg (34951 bytes)
Aerial View of a Large Cyanobacterial Bloom
CyanophytaBloomHue.jpg (102493 bytes)
Cyanophyte Bloom
BloomMicrocystisB.jpg (18001 bytes)
Microcystis Bloom
MicrocystisLab.jpg (147108 bytes)

Nitrogen Fixation

Nostocaceae with Heterocysts fix NitrogenAnAZ100XFZHetSolo400Lab.jpg (68437 bytes)

Marine Species with Heterocysts

Calothrix & Scytonema

Heterocysts

These have enlarged Thick Walls

The outer envelope is Bilayered.Heterocyst240.jpg (28953 bytes)

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.

ReproductionAkineteHeterocystLMHigh300Lab.jpg (15127 bytes)

Fragmentation

Hormogonia = Gliding Fragments

 

Akinetes -> AkineteEM400Lab.jpg (9321 bytes)

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.CalothrixPairCropHue200.jpg (65118 bytes)

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.