Algae_1 - a pdf on the structure of the simpler algae.
Seaweeds - more complex algae.
Algae - building bodies from balls, chains, sheets and tubes
Chlamydomonas detailed
Volvox - ball-shaped colony
Building bodies - filaments

Many algae are filaments - chains of cells that form when the plane of cell division is fixed resulting in a '1D'
colony. Many bacterial and fungal forms are filamentous and one of the advantages is that it allows organisms to
reach above the substratum (surface) and through the boundary layer (a layer of almost motionless fluid that
forms around all solid objects in a fluid) and obtain a good supply of nutrients in the flowing water and to shed
spores into the faster moving air or water. Some filamentous cyanobacteria (photosynthetic bacteria that are
sometimes called the blue-green algae) live inside slime tubes that they secrete for themselves and they can
glide up an down inside this tube, positioning themselves higher in the water column when needed. See the pdf
book
chapter for more advantages of being filamentous to bacteria. An advantage of being multicellular is that
different cells can specialise to perform different functions.
Building bodies - spheres

Algae demonstrate well some of the different approaches to building multicellular bodies - there are a
number of ways to construct a 3D body from cells. Cells can be joined together to form chains (essentially
1D organisms) or more precisely remain joined together when they reproduce by cell division. If all the
cells divide in the same plane, then a chain will result. By dividing sometimes in a second plane at right
angles to the first, a 2D sheet of cells may be produced (see
Ulva and Porphyra below). Volvox is a green
alga that consists of a hollow ball of cells, which is essentially a 2D colony or sheet folded around and
joined together. Each ball, or
coenobium, consists of a single layer of superficial cells - it is a colony of
cells. Each cell is surrounded by a thick mucilaginous wall, forming an enclosing cell. In some species,
these mucilaginous walls may extend toward the centre of the ball and almost completely fill it. Each cell
sits on the surface of the sphere and bears two
flagella which protrude into the surrounding water and
beat to propel the whole colony through the water. Each cell has a red eyespot (stigma) which forms part
of a photosensor.
Volvox, like other algae, is photosynthetic and so it has to keep itself illuminated and so
it will swim toward the light (or away from very bright lights that may damage its chlorophyll). This
immediately poses a problem - if each cell beats its flagella independently of the others then the colony
will move nowhere, rather they must coordinate themselves so as to beat in unison. To achieve this
coordination, all the cells are connected to their nearest neighbours by
protoplasmic bridges, such that
all the cell cytoplasms are continuous with one another through these bridges. This allows waves of
electric charge to travel throughout the colony, triggering flagella motion in an organised and controlled
manner.

The number of cells present in
Volvox and related genera (which form a group called the Volvocales);
Gonium forms flat plates with 4 or 16 cells, Pandorina forms a sphere of 16 cells. The number of cells in
the larger
Volvox is always a power of two, and may number several thousand, a result of coordinated cell
division with all cells dividing and doubling in unison. The
Volvox ball has a preferred front-end and cells
in this part of the sphere have larger eyespots than the rest.
Chlamydomonas is an example of an alga
comprising a single cell that resembles a cell from a
Volvox colony.

Reproduction in Volvox

Volvox
can reproduce asexually by forming daughter colonies. Daughter colonies form as hollow balls
of cells inside the parental colony. The daughter colonies form from enlarged cells in the surface of the
parent colony, called
gonidia, at the posterior end of the colony. Each gonidium divides repeatedly in a
plane at right-angles to the surface of the parent coenobium, forming a cup-shaped plate of cells.
Divisions continue and the plate forms a small spherical daughter colony inside the parent colony and
suspended from its surface. The daughter colonies are originally formed inside-out, with their flagella
pointing inwards, and invaginate or invert, often at around the time that they escape from the parent
colony which ruptures and dies, releasing the daughters which are miniature versions of the parent and
grow by cell division. Daughter colonies may contain small granddaughter colonies upon hatching.

Some species are monoecious (hermaphroditic) whilst others are dioecious (with two separate sexes). In
dioecious forms, sexual reproduction begins when some male colonies appear and secrete pheromones
that induce the gonidia of other colonies to undergo sexual rather than asexual reproduction. The gonidia
of colonies of female clones (descended or cloned by asexual reproduction from a mother colony) are
stimulated to develop into female daughter colonies at the posterior of the parent colony. In colonies of
male lineages (descended or cloned by asexual reproduction from a father colony) the gonidia are
stimulated to develop into male colonies. Female colonies produce specialised
egg cells - enlarged
superficial cells at the posterior of the colony and which lack flagella and remain attached to neighbouring
cells by the protoplasmic bridges. Male colonies produce packets of
spermatozoids, also at the
posterior of the colony. These form much like daughter colonies in asexual reproduction - a superficial
cell divides to produce a hollow ball of cells hanging down from the wall into the interior of the coenobium.
These cells are naked (lack cell walls) and flagellated and are originally inside-out but invert so that their
flagella point outwards. These cells detach from the parent colony and swim towards egg cells, which are
immotile and remain bound to the female colonies.  This is oogamy - large immotile egg cells are formed,
toward which the sperm swim. The gametes are produced by mitosis, so the parent coenobia are haploid.
When a spermatozoid fertilises an egg cell, a thick-walled
hypnozygote is formed. A hypnozygote is a
diploid cell formed by fertilisation (zygotes) which enters a mandatory dormant stage. It has a thick spiny
wall to protect the zygote within.

The possession of dormant stages is particularly important to freshwater organisms that live in ephemeral
ponds. If a pond dries or freezes, then the dormant stages can survive until better conditions for growth
return. After dormancy, the
Volvox hypnozygote undergoes meiosis and germinates to produce a haploid
zoid which undergoes mitosis to form a new coenobium. The zygote is the only diploid stage, making
Volvox an example of what is called a haplont.
Above: Chlamydomonas with a detailed version on the right and simpler versions thumbnailed on the left. A,
mitochondrion; B, cell wall; C, thylakoid membrane of chloroplast; D, cell-surface membrane; E, flagellum x 2; F,
vesicle/vacuole or storage granule; G, nucleus; H, Golgi apparatus; I, (cup-shaped) chloroplast, and J, starch
grains. One of the morphologically simplest single-celled algae is
Chlorella, which grows on walls and tree bark.
Bacteria - study more about bacteria, including cyanobacteria. Filamentous bacteria are covered in more
detail
here.
Spirogyra
Spirogyra, illustrated above in a 3D mathematical computer model (Pov-Ray) is a green alga which forms filamentous chains of
cells, which float freely in ponds and other bodies of still water. Each cell has a distinctive and characteristic spiral green
chloroplast. Many cells will typically be joined in long filamentous chains.
Spirogyra lives in freshwater ponds and wet drainage
ditches and the like.

Spirogyra, like other algae, needs light for photosynthesis. Simply relying on passive means of reaching the light (like flotation)
can be problematic and many algae can also move toward the light.
Spirogyra has no flagella and cannot swim quickly as can
Chlamydomonas, but it is nevertheless capable of limited movement. Blue light will trigger positive phototaxis or movement
toward the light.  First the filaments align, pointing toward the light source and then they bind together into bundles (this
bundling is essential for the following movements). First the anterior filaments in the bundles, nearest the light, curve toward
the light and then those at the rear also roll up toward the light forming open hoops and then the filaments stretch out again.
Repeated rolling and stretching allows the mat of filaments to move toward the light in mass, albeit quite slowly at about 1
millimetre per minute. Filaments are also capable of slowly gliding along, possibly as a result of mucilage secretion, and will
move up the sides of glass containers, generally moving faster in the dark. It really is amazing teh tyricks that apparently simple
organisms have evolved to enhance their survival!

The end cells on each filament of
Spirogyra are also capable of adhesion. Blue or UV light will rapidly stimulate these anchor
cells to adhere reversibly to glass, probably by secretion of cementing mucoproteins. An increase in temperature and shaking
the culture in the dark can also trigger this rapid and reversible adhesion. Red light will also trigger a slow adhesion (takes
about 1 hour to develop) which is apparently irreversible.

Filaments of
Spirogyra grow by division (binary fission) of cells throughout the filament (except perhaps the end cells) and
asexual reproduction comes about when filaments fragment and each sub-filament continues growing. This fragmentation is
not necessarily passive - there are changes in the joining walls that weaken the filament at certain breakage points and then
one cell may swell more than its neighbour as it takes up water and stretches under turgor pressure pushing against the
weakened join and breaking it.

Sexual reproduction occurs by
conjugation (not to be confused with conjugation in bacteria which is a different process that
serves a very different function). The parent (vegetative) filaments are haploid (n). Two filaments will line up side-by-side
(solitary filaments are apparently non-motile) and then projections will grow from the sides of the cells in one filament toward its
neighbour. The neighbouring filament responds in like fashion and the protuberances of each filament contact one-another
(and push the filaments apart slightly as they develop) and an open bridge is formed. The protoplast of the male cell will round
up and crawl across the bridge into the female cell. The two protoplasts fuse and a thick-walled diploid (2n)
zygospore is
formed. The zygospore can survive harsh conditions, such as the pond drying-up but will eventually divide by meiosis to
produce 4 haploid cells(n) only one of which will survive to produce a new vegetative filament on germination.
Spirogyra
Scenedesmus
Scenedesmus
Scenedesmus
Scenedesmus is a green alga that forms small colonies equipped with
spines for flotation. The morphology varies with conditions and
species, but this arrangement of 4 cells with 4 spines is quite
common. Each cell is about 10 micrometres long. [3D Pov-Ray model].
The spines are flotation devices, keeping the
colony floating high in the water column near the
source of light needed for photosynthesis. This
may seem surprising, but at this microscopic scale
water behaves much like treacle (low Reynold's
number) and spines are efficient flotation devices.
Scenedesmus is a beautiful organism to observe under your microscope! The large pyrenoids (for starch synthesis) inside
the large green chloroplasts are clearly visible. Flotation helps keep the cells at the surface near the valuable sunlight, and
flotation will also help disperse the organism and the spines probably also provide some protection against predators by
making the organism larger. Indeed,
Scenedesmus often grows as single-cells in culture (each cell often possessing 4
spines) but chemicals released by predators are detected by
Scenedesmus (which are too large for many predators) and
this induces the formation of longer chains of cells and larger colonies, an effect which has been empirically shown to
reduce losses to predation. However, the larger colonies sink more easily and so this defense comes at a cost, which is why
in the absence of predators single cells dominate. As a further aid to dispersion,
Scenedesmus produces wall-less
biflagellate
zoospores or gametes (smaller cells, about 5 micrometres long with a chloroplast but no pyrenoid) that swim in
the water and if two compatible zoospores meet then they fuse in fertilisation to form a zygote and a new
Scenedesmus. In
addition to this mode of sexual reproduction,
Scenedesmus reproduces asexually by producing a miniature autocolony by
division of each protoplast inside the parent cell wall. The wall of the parent cell then splits and the autocolony emerges. In
addition to the spines, which are often confined to the end cells, the middle cells may bear long fine bristles. These bristles
possibly assist in flotation.
Cyanobacteria link
Ulva external features
Ulva
Ulva anatomy
Young Ulva
Ulva life cycle
Spirogyra_diagram_labeled
Sea-Lettuce - Ulva lactuca
Sea-lettuce, Ulva lactuca is a seaweed of rocky
shores (upper, middle and lower shores). Its thallus
or frond is a thin bright-green translucent sheet of
varied shape anchored by a holdfast and
sometimes a short stalk (stipe) though in some
locations this seaweed survives as a free-floating
weed. Size varies from 5 to 5o cm, sometimes over
one metre in length. The thallus is only two cells
thick.
Left: a section through an Ulva frond.
Two sheets of closely spaced
photosynthetic cells make up the frond
(
Ulva is said to be distromatic). Each
cell has a single cup-shaped
chloroplast which is parietal (lies in the
margin of the cell) and during the day is
positioned facing the surface, to
intercept the sunlight for
photosynthesis. At night the
chloroplasts move, typically sitting at
one of the long edges of the cell. Some
of the cells, especially those in the
stalk, give out multinucleate hyphal-like
appendages called
rhizoids, which
strengthen the stalk.

Growth is
intercallary (occurring at
areas throughout the length of the
frond rather than the base or apex only)
as any of the cells can divide. The
plane of division is always at right
angles to the surface of the thallus, so
that there are always two sheets of cells.

All the vegetative cells along the margin
of the thallus can reproduce by spore
formation. There are two genetic forms
of
Ulva - a haploid generation with only
one complete set of chromosomes (n)
called the
gametophyte and a diploid
sporophyte generation with two
complete sets of chromosoems (2n).
these generations alternate throughout
the life-cycle (alternation of
generations).

The gametophytes produces haploid
wall-less (naked) spores by mitosis, and
these spores are the gametes. Female
weeds produce female gametes and
male weeds male gametes (
Ulva is
dioecious - the sexes are separate).
The gametes are biflagellate (they have
two flagella). A male and female gamete
fuse in a process of
anisogamous
fertilisation. The gametes are produced
inside the vegatative cell walls and
escape into the sea via pores - each
vegetative parent cell forming a pore
connecting it to the outside surface.
Isogamy is the production of identical
male and female gametes, and
although
Ulva is often described as
isogamous, the female gametes are
actually considerably larger and
Ulva is
more correctly described as
anisogamous (producing distinctly
different male and female gametes).

The zygoge germinates to produce a
diploid sporophyte, which is identical in
appearance to the gametophytes. The
sporophyte, however, produces
haploid zoospores by meiosis, each
zoospore is quadriflagellate (has 4
flagella) and germinates into a
gametophyte thallus.

Both the immature gametophytes and
sporophytes develop first into a
uniseriate filamentous juvenile stage
with rhizoids (shown above) by
repeated mitosis - uniseriate means
that the filament is comprised of a
single column of cells.
Later on the cells divide in a second plane and a multiseriate filament (made of several columns of cells is produced) followed by a
hollow tubular stage. This then expands and flattens to form the sheet-like mature thallus.
Enteromorpha is a closely related form that
consists of hollow tubes, each made from a single sheet of cells, apparently
Enteromorpha passes through a young sheet-like, Ulva-like
stage but then the two cell sheets separate and curve around to form cylinders.
Ulvaria, another closely related form, passes through an
Enteromorpha-like tubular form which splits open at the upper end to form a sheet that is only one cell thick (monostromatic). These
species are closely related and hybrid-forms are known. These species all develop from a young filamentous stage, which suggests that
they evolved from such forms (often, though not always, the development of an organism resembles the evolutionary stages that it went
through as evolutionary progress can occur when genetic changes modify the developmental process, and one such modification is to
extend the development by adding additional stages on to the end of it). Thus, we have seen how single-cells that fails to divide
completely can evolve into multicellular filaments, like
Spirogyra, and how adding in an additional plane of division can produce sheets
and cylinders. The most complex seaweeds are types of brown algae (Phaeophyceae) not green algae like
Chlamydomonas, Volvox,
Spirogyra, Scenedesmus and Ulva.
Brown seaweeds - the most
complex algae.
A myriad of colours

There are many types of algae, but three of the main divisions are the green-algae or Chlorophyta (like Volvox, Chlamydomonas,
Spirogyra, Scenedesmus and Ulva), the brown algae and the red algae. All contain a type of chlorophyll called chlorophyll-a to trap
sunlight for photosynthesis, but additional pigments give the algae there various colours and expand the range of wavelengths or
colours of light that can be used for photosynthesis. Green algae also contain chlorophyll-b and carotenes, especially beta-carotene
(whose orange colour is masked by the green chlorophyll). Brown algae contain traces of chlorophyll c and a brown pigment called
fucoxanthin. Red algae contain traces of chlorophyll-d, beta-carotene and a red pigment called phcoerythrin.

The blue-green algae or cyanobacteria, like
Anabaena, have chlorophyll-a, beta-carotene and the blue pigment phycocyanin. there is
a group called the golden or golden-brown algae, which contain chlorophyll-a and beta-carotene.

Siphonaceous Algae

So far we have looked at unicellular algae, simple colonial and filamentous algae and sheet-like seaweeds (and more complex brown
seaweeds) that are beginning to resemble plants in complexity and design. One major body-type that has not yet been considered are
the siphonaceous algae. These consist of single large multinucleate cells, often with tubular form and with peripheral cytoplasm and
large central vacuoles.
Caulerpa is one of the most plant-like of these forms; it consists of a long rhizome tube from which branch
root-like rhizoids and leaf-like, sometimes feather-like, fronds. The whole structure can be several metres long and each frond several
centimetres in height and yet it is a single cell. Struts project from the cell wall into the cytoplasm to give it additional mechanical
strength and in rougher waters these trabeculae increase in development. This suggests the reason why most complex algae have
adopted multicellularity - it gives tissues mechanical strength. Both the formation of an enlarged multinucleate cell by nuclear division
without cytoplasmic division and multicellularity result from modifications to cell division. In the former, there is nuclear division without
cytoplasmic division and in the latter by a failure of the cells to separate completely after cell division.
The theme of this article is the algae or protoplants. In addition to explaining general and some specific
aspects of algal biology, this page concentrates on the theme of building bodies. Algae illustrate many
different ways in which single cells can evolve into more complex and larger multicellular forms by certain
key modifications. The diversity of algae is astonishing and only a few examples can be covered here.
Acetabularia
Acetabularia
Above: A 3D Pov-Ray computer model of the reproductive stage of Acetabularia, without and with terminal hairs.
Acetabularia is also called the Mermaid's Wineglass or the Mermaid's Cup and is essentially a single-celled alga that can
grow to about 5 cm in height. The vegetative stage consists of the stalk and anchoring rhizoids with whorls of branched hairs
along the stalk. These hairs generally drop-off by the time the reproductive cap or umbel forms. A single giant nucleus is
situated at the base of the stalk and this produces and sends out smaller nuclei that migrate to the developing cap to
become incorporated into the cap rays where they develop into spores. There is a large central vacuole throughout the cell.
When mature, the cap becomes divided from the stem and then constitutes a separate cellular system. Different species can
be identified by the number of fleshy rays and the degree to which the rays are free, fused together or fused at their bases
only. The terminal hairs are not always visible in photographs and may not be present in all species (I have been unable to
find information on this).
Spiyrogyra_conjugating
Spirogyra
Right: micrograph of Spirogyra conjugating (fixed permanent
preparation). Far right: A
Spirogyra cell (fixed permanent prep.) -
click to enlarge.
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