WB01343_.gif (599 bytes)     Rhodophyta     WB01345_.gif (616 bytes)

Dr. Celia Smith's ReefAlgae Website

Dr. Derek Keats Coraline Algae Website

Diversity   Genera -800     Species - 5200Smithora.gif (15513 bytes)

Color  Red -> Black

Varies with Depth

High Intertidal Zone -> Green -> Black

Lower Intertidal Zone -> Brown - Purple

Greater Depths -> Rosy Red

Depends on Pigment Types & Concentrations

Photosynthetic Pigments

Chlorophyll a, d

Phycobillins - Phycoerythrin & Phycocyanin

Other Accessory PigmentsRedSand120.jpg (22409 bytes)

Carotenoids - Carotene a & b

Xanthophylls - Leutin & Zeaxanthan

Occurrence

Most species are Marine (96%).

In Hawaii 70% of the algal species are in the Rhodophyta.

GelidiumYellowBack.jpg (54612 bytes)
Gelidium - Source of Agar

They have a Diverse Distribution & range from Tropical to Cold Waters. Some Red Algae can be found in Cold Fast Running Freshwater Streams.

They,consequently have the Greatest Range for Macroalgae.

They can be found at depths up to 120m

A few species are Terrestrial

Cell Wall

Inner Cellulose Wall

Random Cellulose Fibrils

Granular Matrix

CoralinAlga240.jpg (18681 bytes)Coralline Rhodophyta

Outer Sheath = Colloidal Substances

Agar

Carrageenan

Others

GeniculateSA.gif (19831 bytes)
Geniculate Coralline Alga

Coralline Algae

These precipitate Calcium Carbonate in their Outer Wall Layers. This forms a rigid extracellular matrix that contributes to the formation of Coral Reefs. There are two general types based on their morphology. These are called Geniculate and Non-Geniculate.GeniclateBWDiagLabHue.jpg (88321 bytes)

Geniculate -> Branched & Flexible due to the presence of Genicula between rigid Intergeniculate segments.

Non-Geniculate -> These are encrusting and are generally prostrate. Howecer some produce small

NonGeniculate-1.gif (37168 bytes)

Nongeniculate Coralline Algae

foliose branches, while others are nodular.

 

 

 

Organelles

Nucleus

Halydictyon250.jpg (26376 bytes)

Halydictyon sp.- (A Coenocyte) Its Nuclei are Stained with DAPI

Some simple organisms are Uninucleate.

Complex Organisms are Multinucleate (Coenocytic).

There can be  100 to  1000 Nuclei per cell!

They are Eukaryotes and have True Chromosomes. N = 2 - 90

 

Cell Division & New Wall Organization

The New Cell Wall exhibits Annular Growth.PitConnectionLab.jpg (55232 bytes)New Wall formation starts from the Outside and proceeds centripetally (towards the center).This is similar to closing an Iris Diaphragm like the Field Diaphragm of a microscope or the diaphragm in a camera lens or in your eye.

A large opening remains between the two cells. A Protein Plug completely blocks cytoplasmic continuity.

This is called the "Primary Pit Connection" and is unique to Rhodophyta.

Chloroplasts

There can be from one to many chloroplasts per cell.

ChlPlastPorph300Lab.jpg (64887 bytes)

Poryphyridium Cell Showing the Chloroplast & Nucleus

Simple Forms have 1 to several.

Larger More Complex Forms have many Small chloroplasts.

Ultrastructure

Chloroplasts have a Double Limiting Membrane with Solitary Thylakoids that are Widely Spaced. This is partly due to the presence of Phycobilisomes which protrude above the surface of the Thylakoids.

PhycobilisomeDraw.jpg (28474 bytes)

Pyrenoids may be present. These have high Concentrations of RUBISCO, the enzyme involved in the Fixation of CO2. Pyrenoids may allow carbon fixation at lower CO2 levels.

ChlpLaurencia300Lab.jpg (88190 bytes)

Chloroplast of Laurencia

Stroma (matrix) is Homogenous & DNA is present.

Starch (Floridean Starch) is stored in the Cytoplasm which also contains, Mitochondria, Dictyosomes, Endoplasmic Reticulum & Ribosomes.

 

Life Modes

Morphology-1-240.jpg (13485 bytes) FilM-BranchedFilm-240.jpg (27338 bytes)
Growth Forms Bangiophyceae

Most species are Autotrophs.Some are Epiphytes while a few are Endophytes with Diatoms.Some are Symbionts and some are Parasites

Class Bangiophyceae

Mostly Marine with a few feshwater speceis.

They conatin Unicellular and Filamentous species.

The latter range from Uniseriate to   Multiseriate filaments.

PolystromLab.gif (21527 bytes)

Polystromatic - Prostrate Form

They may be Unbranched or Branched, Prostrate or Upright

They may be Monostromatic (One Layer Thick) or Polystromatic (Many Layers)

The Most Advanced forms are called Diffuse Foliose and their Cells Divide in 2 - 3  Planes

 

Attachments

They are attached to the substrate by a

Holdfast Cell or a Group of Rhizoidal Cells

 

Porphyra.gif (31861 bytes)

Porphyra

Life Cycle Porphyra

Thallus
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Vegetative Cell
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Monospore
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Relase
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Spores Germinate 
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New Organism

LifeCycleDirectLab.jpg (92320 bytes)
Porphyra Direct Life cycle

Direct with Aplanospores

Thallus
WB01569_.gif (193 bytes)
Clevage of a Cells
WB01569_.gif (193 bytes)
Several Aplanospores (NonFlagellated)
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Each Spore -> New Thallus

PorphyraLifeCycle2(Aplano)300Lab.jpg (103968 bytes)

Porphyra Life Cycle with Aplanospores

Type 3 with a Conchocelis

Gametophyte Thallus (N)
WB01569_.gif (193 bytes)
Spermatangium->Spermatia & Carpogonium->Functional Egg
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Syngamy -> 2N
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Carposporangium
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Carpospores
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Germinate
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Conchocelis Stage (2N) = Sporophyte
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Unicellular -> Filamentous
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Monosporangia -> Meiosis
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Conchospores
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Thallose  Gametophyte

PorphyraLifeCycle-3-300Lab.jpg (193325 bytes)

Porphyra Life Cycle with Conchocelis

GelidiumYellowBack240.jpg (20286 bytes)

Gelidium

Florideophyceae

Larger Class

Genera & Species

Marine

Freshwater -> 3%

Substrates -> Solid

Pit Connections

Primary Pit Connections = Always

Delessaria.jpg (16558 bytes)

Delessaria sp.

Secondary Pit Connections

Found with Larger Forms

Unique

Growth Forms

No Unicells

Simplest= Uniseriiate - Branched Filaments

Complex Forms

Filaments Latetrally Fused

"Pseudoparenchymatous" Thallus

"Cortex" Small Cells

"Medula" Loose Filamentous Cells

Apical Cells -> Growth

 

Growth Forms

Uniaxial -> Single Row Large Cells

Multiaxial-> Rows of Parallel Filaments

 

Growth Potential

Main Axis -> "Unlimited" Growth

Lateral Axes -> Determinate Growth

Gross Morphology

Highly Variable

Size Microscopic -> 1m -> 4m (Gracilaria)

 

Nemalion150Red.jpg (13919 bytes)

NemalionX-S240.jpg (18710 bytes)

Nemalion Plants Nemalion X-S

Generalized Life Cycle

GeneralRedAlgalLifeCycle.gif (19144 bytes)
Generalized Sexual Reproductive Cycle for a Red Alga  Florideophyceae

 

Sexual Reproduction Overview

Reproduction = Oogamous

Female = Large & Nonmotile

Male = Small (with OR without Flagellum)

Gametes = NONMotile

Gametangia

Spermatangium -> Spermatia

Carpogonium ->   "Egg"

Trichogyne = Receptive " Hair" of   Female Gametangium

Trichogyne300Lab.jpg (44213 bytes)

Female Gamete

Retained on Gametophyte

Fertilization in situ

Diploid Sporophyte -> Carposporophyte

Retained within the Carpogonium

Carposporangia -> Carpospores ->

Tetrasporophyte -> Meiosis (Tetrasporangium)

Tetraspores (4/sporangium) -> Gametophytes

Monoecious (Male & Female = Same Plant)

Diecious (Male vs Female Plants)

Polysiphonia Parts

PolysiphoniaVeg240.jpg (24231 bytes)

PolysiphoniaMicroTip240.jpg (22319 bytes)

Polysiphonia Gametophyte

Polysiphonia - Branch Tips

 

PolysiphoniaAnther240.jpg (15799 bytes) PolysiphoniaCarpoSporangiaClosed240.jpg (24526 bytes)
Polysiphonia   Spermatangium Polysiphonia Carpogonium

 

PolysiphoniaCarpoOpen240Lab.jpg (84810 bytes)

Polysiphonia - Open Carpogonium

 

PolysiphoniaTetrasporophyte.240.jpg (22489 bytes)

Polysiphonia - Tetrasporophyte

 

Tropical Reefsridgcorl.gif (12402 bytes)

Coralline Algae  Link to Derek Keys Web Site WB01436_.gif (236 bytes)

Nongeniculate Forms -> Reef Stability

Most Abundant Organisms

Most Important for Reef Stability

Cement Coarse & Fine Grain Sediments

Principal Reef Builders

Withstand Desiccation at Full Sunlight

Maxium Abundance occurs in Shallow, Physically Disturbed Sites.

They Require Wave Shock & Grazing to prevent Overgrowth by other Fleshy Algae.

Some Corallines have the ability to Shed Surface Cells. This Removes excessive amounts of Epiphytic organisms.

This SEM image shows the process of  Epithallial Shedding which removes excess Epiphytes ShedSheet.gif (32356 bytes)

Encrusting coralline algae flourish where waves break on the reef, forming an elevated bank known as the algal ridge. As water from the waves returns seaward, its load of suspended sand and gravel scours a series of narrow ridges. The alternating ridges and valleys are known as spurs and grooves.

Corals growing on top of the spurs are stoutly formed and compressed in shape to withstand the extreme force of breaking waves.

Being algae, the plant body in corlallines is referred to as a thallus (pl. thalli). Most coralline algae have an internal anatomy of the thallus that is based on the aggregation of branching filaments, each of which is formed through the action of its own single meristematic cell. This type of thallus anatomy is said to be pseudoparenchymatous

Most nongeniculate corallines show some degree of dorsiventral arrangement, that is, an arrangement in which a distinct upper and lower surface can be distinguished. There are two different types of organization in dorsiventral thalli:

In dimerous thalli, there are two distinct groups of filaments oriented more-or-less at right angles to one another, with a ventral layer of filaments (called basal filaments) that is uually one cell thick, and an erect layer in which filaments (called erect filaments) vary from one to many cells in length and comprise most of the thallus thickness. Dimerous thalli arise when a spore germinates to produce a unistratose layer of radiating, repeatedly branched filaments, that grow via meristematic cells that are located at the apices of the branches. The growth of this layer of basal filaments contributes to the areal expansion of the thallus. Erect filaments are formed in most dimerous species when cells of the basal filaments divide periclinally to form erect filaments. The first cell that is cut off remains meristematic and forms new cells of the erect filament ventrally, and epithallial cells dorsally. Since it is usually overlain by one or more epithallial cells, the meristematic cell is usually referred to as a subepithallial initial. Dimerous thalli occur in Lithophyllum, Titanoderma, Exilicrusta, Melobesia, and some species of Hydrolithon and Pneophyllum.

In monomerous thalli, there is but a single pseudoparenchymatous system of repeatedly branched filaments in which some derivatives contribute to a medulla which runs more-or-less parallel to the thallus surface (or substratum in dorsiventral, attached species) and some derivatives curve outwards and collectively form a cortex. In most dorsiventral thalli, downwards-curving filaments are suppressed, and only occur vestigially. The cortical filaments terminate at the upper surface in one or more epithallial cells, that are subtended by a subepithallial initial. Continued division of the subepithallial initial increases the thallus girth, and may give rise to surface protuberances which may themselves branch. Monomerous thalli occur in Clathromorphum, Leptophytum, Lithothamnion, Phymatolithon, Mesophyllum, and many species of Hydrolithon, Spongites, and Pneophyllum. In some species, the primary thallus is dimerous, but monomerous regions may develop secondarily from the cells of erect filaments (e.g. Lithophyllum incrustans).

In both monomerous and dimerous forms, the cells of adjacent filaments below the merismetic cells often fuse together, or form pit connections, allowing for cellular interchange throughout the plant in spite of the calcified cell walls. Cell fusions and secondary pit connections are used for taxonomic purposes at the sub-family as well as species level.

In dimerous thalli, there are basal and erect filaments. There is a primary terminal
initial (meristmatic cell) located at the margin of the thallus, and from which the thallus
expands laterally.

The pseudoparenchymatous nature
of the thallus is evident in dimerous
thalli. This is incorporated into the
terminology used to describe the
regions of the thallus: basal and erect
filaments.
Dimerous thalli occur in Lithophyllum, Titanoderma, Exilicrusta, Melobesia, and some species of Hydrolithon and Pneophyllum

 

In monomerous thalli, there is a single system of repeatedly branching filaments. In dorsi-
ventrally oriented species, there is usually a layer if filaments located near the substratum, from
which branches repeatedly arise and curve towards the dorsal surface.

 

Under the traditional terminology that was applied to corallines, the lower filaments were referred to as
a the hypothallus, while their dorsally curving derrivatives were referred to as the perithallus.

 

This terminology suggests that corallines are somehow different from other red algae
with respect to the origin of these filaments. However, monomerous thalli are
homologous with multiaxial thalli in many erect red algae, in which the inner filaments
are referred to as the medulla and their outward derrivatives are called the cortex.

 

Under the traditional terminology that was applied to corallines, the lower filaments were referred to as
a the hypothallus, while their dorsally curving derrivatives were referred to as the perithallus.

 

This terminology suggests that corallines are somehow different from other red algae
with respect to the origin of these filaments. However, monomerous thalli are
homologous with multiaxial thalli in many erect red algae, in which the inner filaments
are referred to as the medulla and their outward derrivatives are called the cortex.

 

To emphasize this commonality with other multiaxial red algae, the ventral
layer of filaments are referred to here as the medulla, and their peripheral
derrivatives are referred to as the cortex. We can also speak of medullary
filaments
and cortical filaments, to refer to the filaments which make up
the medulla and cortex respectively

. Monomerous thalli occur in Clathromorphum, Leptophytum, Lithothamnion, Phymatolithon, Mesophyllum, and many species of Hydrolithon, Spongites, and Pneophyllum. In some species, the primary thallus is dimerous, but monomerous regions may develop secondarily from the cells of erect filaments (e.g. Lithophyllum incrustans).

In both monomerous and dimerous forms, the cells of adjacent filaments below the merismetic cells often fuse together, or form pit connections, allowing for cellular interchange throughout the plant in spite of the calcified cell walls. Cell fusions and secondary pit connections are used for taxonomic purposes at the sub-family as well as species level.

 

 

Reproduction

Reproduction in the coralline algae is as kinky as it is in most of the other red algae. Three distinct phases occur in the life cycle, two of which are free-living, and one of which is attached to and dependent on the female plant. Both of the free-living stages look almost identical, differing only in their reproductive organs, which are usually borne inside small flask-shaped or elliptical cavities or depressions called conceptacles.

The stage which produces gametes is called the gametophyte. After fertilization, the zygote nucleus undergoes a special type of development to give rise to a tiny, uncalcified plant called the carposporophyte. The carposporophyte has twice the chromosomes number of the gametophyte generation, and develops inside the conceptacles of the female gametophyte. The carposporophyte produces spores, called carpospores, which are released through a pore in the female conceptacle. These spores, which still have twice the chromosome number as the gametophytes, germinate and grow into a third generation called the tetrasporophyte.

 

Female reproductive structures consist of
a carpogonium borne on a hypogynous
cell
. The carpogonium is extended into
a receptive process (received spermatium
during fertilization) called a trichogyne.
Sometimes a sterile cell is also present.

 

The carposporophyte develops from the zygote nucleus following fertilization.
It is thus diploid. It remains within the female conceptacle, which enlarges to
accomodate it. In the diagram below, the carposporophyte is seen to consist of
the gonimoblast filaments, each of which terminates in a carposporangium
which produces a single carpospore. The filaments illustrated in pink are haploid
filaments of the gametophyte which surround the casposporophyte and enclose it
within the conceptacle. The conceptacle and its contents are sometimes referred to
as a cystocarpic conceptacle.

 

Diploid: having twice the haploid chromosome number. Following fertilization, a
diploid carposporophyte develops within the female conceptacle. The diploid
carposporophyte produces diploid carpospores by mitosis, and these are released and
grow into diploid tetrasporophyte

 

Meiosis is the process of reduction division, whereby the chromosome number in the nucleus is halved. The
fusion of gametes restores the chromosome number to the full complement. Following meiosis cells are said to
be haploid. Following the fusion of gametes, cells are said to be diploid. In coralline algae, meiosis takes
place in diploid tetrasporangia and results in haploid tetraspores. See also tetrasporophyte

 

Haploid: having half the diploid chromosome number. In coralline algae meiosis takes place
in the tetrasporophyte to gice rise to haploid tetraspores. The haploid tetraspores grow into
haploid gametophytes. The tetraspores and the gametophytes are the only haploid stages
in the coralline algae

 

Halydictyon arachnoideum from Victoria, Australia, a multinucleate red alga with giant cells. Stained with DAPI, a fluorochrome specific to DNA.

 

Laurencia spectabilis

Named after the French naturalist, de la Laurencie. This alga has very distinct branching, on a thick, almost rubbery thallis, as well as an apical pit (look very hard). For you chemists, Laurencia produces secondary metabolites that contain iodine, bromine, and chlorinated compounds--hence tasting yucky to local herbivores.

 

Smithora naiadum

The monostromatic marvel. Its usually found epiphytic on eel and surf grass in these here parts

 

Porphyra nereocystis

is epiphytic (real close neighbor to) on the Bullwhip Kelp, genus name Nereocystis. See Nereocystis! Porphyra as most know it, is commonly called nori. If you eat sushi, then you have probably seen Porphyra. Its the seaweed that your uncooked fishy is wrapped in at the sushi bars.

Link to Algal Ridge (http://www.cyberlearn.com/ridge.htm)