Seaweeds: biology and diversity 1.

Introduction to seaweed diversity; cyanobacteria

Tijs Ketelaar, CLB

Part 1 Life forms, diversity and Part 2 Abiotic stress reproduction
reproduction of algae
CO2 (HCO3-)
O2

(c) Masao Takahashi

Part 4 Production & Farming

Part 3 Photosynthesis

Part 5 Environment, global distribution

(c) Mission blue

Overview of this part of the course

 General introduction
● Algal evolution
● Algal diversity

 Characteristics of different phyla

● Body plan
● Reproduction

3
Suggested literature

 Algae (Third Edition)

Linda E. Graham, James M.
Graham, Lee W. Wilcox, and
Martha E. Cook
ISBN 978-0-9863935-3-2
 http://www.ljlmpress.com/algae.html

 $40,-; eBook only

4
How to define algae?

 Heterogeneous assemblage of organisms from single cells to

giant seaweeds

 Belong to diverse evolutionary lineages

 Photosynthetic species that produce oxygen and live in aquatic
habitats

 Lack body and reproductive features of land plants

 This concept includes eukaryotic photosynthetic protists and
the prokaryotic cyanobacteria

 Anything that has plastids and is not a land plant

5
 Photosynthesis

 Oxygenic photosynthesis
● Oxygen producing, common type of
photosynthesis

● Anoxygenic photosynthesis occurs in

some bacteria and archae
(prokaryotes). Strictly anaerobic,
often sulfur producing

 The essential light absorbing pigment

is chlorophyll a
 Accessory pigments extend
wavelength range of light absorption
compared to chlorophyll a – important
in water!
Much more about this in the 6
photosynthesis part of the course!
Optimizing photosynthesis efficiency: phototaxis

 Sensing light quality/quantity/directionality and responding to

these cues
 Sensing:
● Light quality/quantity sensing proteins: phototrophin,
phytochrome, rhodopsin

● Light directionality sensing: eyespot/stigma that shades light

detection machinery depending on the orientation of the cell

 Responding:
● Positioning in the water column: often active; e.g. osmotic
activity, gas vacuoles, flagellar movement

7
 Algal body types

 Microalgae (microscope needed for observation)

● Unicells: cocoid or flagellate
phytoplankton
● Colonies/coenobia (fixed size)
● (Branched) filaments: periphyton

 Macroalgae (can be observed with the naked eye)

● Coenocytic (=siphonous): single, multinucleate cell
● Parenchymatous: 3-D tissue (often plasmodesmata)
● Pseudoparenchymatous: appear 3-D tissue but is composed of
adjoined filaments

8
Chlorococcum: Microcystis: colony of Pediastrum: Platydorina:
coccoid microalga coccoid cyanobacteria coenobial colony flagellate colony

Mougeotia: Stigeoclonium: Vaucheria: Macrocystis: 9

unbranched filament branched filament coenocytic body parenchymatous

Algal reproduction - asexual

 Unicells:
● Zoospores (flagellate)
● Aplanospores or autospores (non-flagellate; environment or
genetic)

 Autocolonies: produced by coenobia colonies

 Fragmentation of filaments
 Akinetes: develop from growing cells under unfavourable
conditions for survival
● Accumulate storage materials
● Develop thick cell wall

10
Bisection (regular mitosis of unicellular organisms
Autospore formation

Zoospore formation

Aplanospore formation
Fragmentation

Akinete formation
Autocolony formation 11
Sexual reproduction: concepts

 Not in bacteria; they exhibit other transfer mechanisms

 Production and fusion of gametes (n), production and
development of zygote (2n)
 Reproduction may be:
● Isogamous: male and female gametes have the same size
● Anisogamous: one gamete is larger and/or behaves differently
● Oogamous: male gametes fuse with larger, immobile female
egg cells

12
Generic life cycle of eukaryotes

Fertilisation
Gametes

n 2n First cell: zygote

Meiosis

13
3 types of life cycles: 1. zygotic meiosis

 The diploid (2n) zygote undergoes meiosis

● Consequence: all vegetative tissue is haploid (n)

14
3 types of life cycles: 2. gametic meiosis

 The diploid (2n) zygote divides by mitosis, meiosis occurs

during gamete formation
● Consequence: all vegetative tissue is diploid (2n)

15
3 types of life cycles: 3. sporic meiosis

 The zygote (2n) divides by mitosis. Meiosis results in spores

that form a multicellular haploid (n) organism
● Consequence: two multicellular organisms are formed, a
gamete producing gametophyte (n) and a spore producing
sporophyte (2n)

16
 Summary: the timing of meiosis determines
algal ploidy

 Zygotic meiosis
● Haploid organism
 Gametic meiosis
● Diploid organism
 Sporic meiosis
● Independent, multicellular haploid and diploid organism
● Alternation of generations
● Isomorphic: haploid and diploid generation look alike
● Heteromorphic: haploid and diploid generation do not look alike

17
Diversity of evolutionary origin of algae

18
Cyanobacteria
 A survey of algal diversity

 Cyanobacteria
● Well-defined phylum within Bacteria
● Occur in marine, fresh water and terrestrial ecosystems
● Unicellular, colonial, filamentous; sometimes branched
● No gametes, zygotes and meiosis
● Often nitrogen fixation capacity
● Can cause harmful blooms and often produce toxins
 Glaucophytes
● Unicellular and colonial sweet water algae
● Eukaryote with plastids that most resemble cyanobacteria

19
Diversity of evolutionary origin of algae

Cyanobacteria

Merismopedia 20

Glaucocystis
A survey of algal diversity

 Chlorarachniophytes
● small group of unicellular marine species
● Fall within the eukaryotic supergroup Rhizaria
● Plastids resemble those of green algae
● Of interest for study of plastid endosymbiosis
 Euglenoids
● Many fresh water, some marine species
● Fall within the eukaryotic supergroup Discoba
● Some species are heterotrophic, some lack plastids

21
 Diversity of evolutionary origin of algae

Gymnochlora

22
A survey of algal diversity

 Cryptomonads
● Marine and fresh water unicellular flagellates
● Some species lack plastids or plastids are colourless
● Plastid colors vary: red, blue-green, olive, brown
● Nutritious food source for aquatic animals
 Haptophytes
● Unicellular flagellates and non-flagellate unicells and colonies
● Primarily marine habitats
● Yellow or golden plastids
● Subgroup coccolithophorids have carbonate scales (coccoliths)
– carbonate and petroleum deposits

23
Diversity of evolutionary origin of algae
coccolith

Emiliania

24
A survey of algal diversity

 Dinoflagellates
● Unicellular flagellates with two flagella of different types
● Mostly marine organisms
● Form the eukaryotic supergroup Alveolata with heterotrophic
ciliates, parasitic apicomplexans (e.g. Plasmodium – malaria!)
and other protists

● Alveolata possess alveoli: membranous sacs at the periphery of

the cells.

● Alveoli of Dinoflagellates often contain cellulose

● Golden plastids
● Source of marine toxins
● Undergo symbiosis with marine invertebrates
25
Diversity of evolutionary origin of algae

26
A survey of algal diversity

 Photosynthetic stramenopiles
● Wide range of algae displaying diverse body types
● Golden-brown coloured plastids (fucoxanthin)
● Comprise at least 12 classes of algae, e.g.
● Silica-walled unicellular diatoms
● Giant kelp, brown macro-algae

27
Diversity of evolutionary origin of algae

Cymbella

A kelp forest

28
A survey of algal diversity

 Rhodophyta – red algae

● Occur as unicells, filaments, aggregations of filaments, and
sheets of cells

● Most abundant in marine environments

● Complete lack of flagella – weird sexual reproduction
● Source of industrial products and food
● Choralline red algae: carbonate producers, choral reefs

29
 Diversity of evolutionary origin of algae

Nitophyllum

30
A survey of algal diversity

 Chlorophyta – green algae

● Diverse algal body types
● Diverse marine, fresh water and land
● Considered to be the ancestors of land plants
● Streptophytes most closely related

31
Diversity of evolutionary origin of algae

32
The role of algae in biogeochemistry: huge!

 Biogeochemistry: the study of chemical interactions between

atmosphere, hydrosphere (aquatic systems), lithosphere
(crustal minerals), and biosphere (living organisms)
 Ancient cyanobacteria generated earth’s first oxygen-
containing atmosphere
 Ancient algae have formed carbonate rock formations and
important fossil fuel deposits
● Store carbon for geological time periods and reducing
atmospheric CO2 levels

 Modern algae produce ~ half of the atmosphere’s oxygen, land

plant photosynthesis produces the other half

33
Oxygenic photosynthesis has increased
atmospheric O2 levels 105 fold!

 Evolution of cyanobacteria probably ~ 3 billion years ago

 Oxygenic photosynthesis created an atmospheric chemistry
suitable for other life forms
● Oxygen-rich atmosphere allowed energy-efficient aerobic
respiration

● An ozone shield against ultraviolet radiation from the sun was

formed – allowed for life to emerge from oceans

34
 The great oxidation event

 Linked with increased diversification of

the cyanobacteria
● including multicellular forms
 First eukaryotes evolved ~ 1.5 billion
years ago
● Exact timing hard to determine
● 1.2 billion yrs old fossils similar to
modern filamentous red algae

● 1 billion yrs old remains like modern

stramenophile algae

● 750 million yrs old branched

filamentous green alga

● ~ 575 million yrs old macrofossils

having shapes similar to modern 35

macroalgae
Evolution of eukaryotic algae

 Uptake of cyanobacterial cells by heterotrophic eukaryotes

 Cyanobacteria persisted in a stable symbiotic relationship;
endosymbionts
 Endosymbionts eventually evolved into plastids
 Plastids of modern-day protists and plants likely originate from
evolutionary processes that started with cyanobacterial
endosymbionts
 Oxygen levels reached modern levels (~21%) 550 million
years ago
● Correlates with diversification of multicellular marine animals

36
How did oxygen accumulate in the
atmosphere?

 Carbon sequestration
● 1. production of carbonic compounds resistant to microbial
breakdown – are buried in anoxic sediments

● 2. generation of sedimentary carbonate minerals (e.g. calcium

carbonate)

 CO2 concentration low in atmosphere and water

● Algae have developed CO2 concentrating mechanisms for CO2
uptake 37
Carbon sequestration (refractory carbon)

38
 Carbon sequestration: some examples

 Cyanobacterium with a sheath containing

the indole-alkaloid scytonemin
● Degradation resistant compound
● Often present in extracellular mucilage
or sheaths of cyanobacteria

 Desmodesmus (green alga) with

degradation resistant algaenan

39
Carbon sequestration: atmospheric CO2 levels
have decreased

 CO2 uptake required for

photosynthesis
 CO2 diffuses 104 x more
readily in air as in water
 CO2 concentrating
mechanisms required
● Rubisco
● Photorespiration

40
CO2 concentrating mechanisms in
cyanobacteria

 HCO3- Uptake transporters

(symport and pumps)
 Carbonic anhydrase (CA):

 CA can transform up to a million

molecules of CO2 per second

 Carboxysomes - polygonal protein

structures coated with a protein
shell contain CA

41
CO2 concentrating mechanisms in eukaryotic
algae

 Carbonic anhydrase
 Membrane transporters
 Pyrenoid: CO2 concentrating
structure in plastids
● May be homologous to
cyanobacterial carboxysomes

● Does not occur in all algae

 Proton excretion: conversion of
carbonate to CO2
● Often associated with
calcification
42
CO2 concentrating mechanisms in eukaryotic
algae

 Pyrenoids (arrows) in the  Calcification of the red alga

plastid of the green alga Bossiella sp.
Mycrosterias sp.

43
Limitations in algal growth conditions

 Limitation in one or more

nutrients
 Limitations in CO2
 Limitations in light quantity or
quality

 Oligotropic aquatic systems

● Low nutrient concentrations
● Clear water
● Large species diversity

44
Limitations in algal growth conditions

Algal growth under limiting Algal growth under high

CO2 conditions levels of CO2

45
 Lack of limitations: eutrophic or
hypereutrophic aquatic systems

 Eutrophic aquatic systems

● Relatively high concentrations of
nutrients

● Large populations, low species

diversity

● Nuisance level growths: e.g.

algal blooms

 Hypereutrophic aquatic systems

● Arise by human actions
● Repeated algal blooms

46
Over-fertilization of aquatic systems: dead
zones

 Foster the development of anoxia (lack of oxygen)

● Caused by oxygen uptake by decay organisms that degrade
algal blooms

● Currently more than 40 coastal zones with anoxya have been

reported: dead zones

● Dead zones result from the input of large amounts of N and P

from inadequately treated wastewaters and run-off from
agricultural operations

Mississippi watershed in
the Gulf of Mexico has
created a large dead
zone
(b) Displays oxygen
levels 47
Biotic associations of algae

 Food webs
 Defence against predators
 Symbiotic interactions

48
Food webs

 A food web is a model that illustrates the feeding interactions

among diverse types of organisms in a particular habitat
 Predator – food size relationship is important
 Plankton: all small organisms that swim/float in the water
● Picoplankton - 0.2 to 2.0 μm: most bacteria, including some
cyanobacteria and very small eukaryotes
● Nanoplankton - 2.0 to 20 μm, many algal species that occur as
single cells, colonies or filaments
● Microplankton - 20 to 200 μm, many algal species that occur as
single cells, colonies or filaments
● Mesoplankton – 0.2 to 2 mm, some types of floating or swimming
algae
● Macroplankton – 2 to20 mm, some types of floating or swimming
algae
49
Food webs

1. Phagotrophs: protists that

consume by phagocytosis
● Collectively named
microzooplankton

● Typically lack rigid cell

coverings
● Include flagellates,
ciliates and amoebae

50
A ciliate phagotroph
 Food webs

2. Zooplankton
● Small animals that occupy open water
habitats

● Include rotifers (c) and microscopic

crustaceans (d and e)

3. Mesograzers
● Somewhat larger animal herbivores
● Include oligochaete worms, freshwater
dipteran larvae, marine amphipods
and pteropods, limpets (f), mussels
(g) crabs, sea urchins (h), insects (i)
and fish (j)

51
Food webs

 Complex relationships between different organisms

● Only recently (over the last decades) the complexity of marine
food webs has been recognised

DOM: dissolved organic

matter
POM: particulate organic
matter

52
 Feeding on algae – some examples

Median view

Cortical view

The ciliate Nassula with ingested An amoeba that is actively feeding

diatom cells on diverse algae

53
Feeding on algae – some examples

A sessile (attached) vorticellid ciliate that feeds on algae and

other particles by using undulating membranes and cilia to
generate water currents

54
 Feeding on algae – some examples

A rotifer that has consumed algae Copepods (Crustaceae (Nl. Kreeftachtigen)) are
Rotifer prey ranges from 1 – 200 µm in important herbifores in marine food webs
diameter

55
 Food webs – algal food quality

 Algal species can

● Have features that decrease ingestibility
or digestibility

● Lack content of mineral nutrients and

essential organic compounds
● Contain toxins
● Prevent ingestions by herbivores
 These algae are defined as low quality
food

The cyanobacterium Mirocystis

 Defense mechanisms prevents ingestion by forming

large gelatinous colonies and
produces toxin
56
Algal defences against herbivory

 Small cell size and fast division rates

● Not all cells are found/eaten and populations can rapidly
regrow

● Examples of this strategy can be found in all algal groups

 Grow in a (cold) season when predator number is low
 Some flagellate algal species can flee from herbivores
 Specific herbivore resistant life stages or structures
 Production of toxins or light flashes
Dinoflagellate bioluminescence
deters copepod feeding by
startling the herbivores and by
making copepods that have
fed on glowing dinoflagellates
more visible to their own
predators 57
 Structural defences – decreasing the chance
of ingestion

Green alga
Desmidium –
mucilage sheets

Dinoflagellate
Ceratium
carolinianum
has horns
Green alga Pediastrum –
multicellular colonies

58
Structural defences – surviving ingestion

Colonial green alga Sphaerocystiscan survive

ingestion by zooplankton by producing copious
mucilage and a tough cell wall

59
Chemical deterrents and toxins

 Production of chemicals that

● Deter grazers (e.g. fatty acids, terpenoids, acetogenins,and
polyphenols such as phlorotannins)

● Inhibit digestive enzymes

● Stun predators: predator is easier to prey on
 Production of toxins (compounds that poison and/or kill other
organisms)
● Occurs dominantly in algae that occur as phytoplankton during
and after algal blooms

● Neurotoxins (different groups), hepatotoxins (inhibit protein

phosphatases type 1 and 2A), cytotoxins

60
Algae in symbiotic associations

 Parasites
● Obtain nutrition from living organisms (hosts), without
ingesting them

● Parasites with disease symptoms: pathogens

● Some algae function as pathogens
 Epibionts
● Grow on surface of symbiont without causing obvious harm
 Mutualistic symbionts
● Beneficial for both partners

61
Parasites and pathogens of algae

 Microorganisms that attack algae include viruses, bacteria,

various protists, and fungi
 A typical milliliter of water contains ~ 103 fungal cells, 106
bacteria, and 107 viruses (Rheinheimer 1991)
 Algae avoid infection by using structural or chemical defenses
 Pathogenic microorganisms are important in controlling algal
populations in nature
 Pathogenic microorganisms can be problematic in algal
monocultures for technological applications

62
Viral pathogens

 Lytic viruses – result: cell lysis

● Affect members of all major classes of algae
● Major cause of phytoplankton mortality (2-10% of populations
per day have been estimated)
● Can result in the decline of phytoplankton blooms
● Also affect macroalgae, e.g. by causing sterility

63
Bacterial pathogens

 Attach to cells and kill by degrading the cell surface and/or

secreting toxic compounds
 Common genera: Cytophaga (also known as Cellulophaga),
Saprospira (phylum Bacteroidetes), Alteromonas and
Pseudoalteromonas

64
Protistan parasites

 Some Stramenopile and Alveolate protists are heterotrophic

parasites of algae

The dinoflagellate
Stylodinium has amoeboid
stages that consume the
cytoplasm of algal cells
(Oedogonium in this case)

65
Fungal parasites

 Chytrids consist of multiple lineages

of aquatic fungi
 They have the potential to greatly
reduce algal populations
 Great host specificity: attack one or
few related species
 Fungal epidemics might end or
prevent algal blooms
 Control the timing and pattern of
phytoplankton succession
Chytrid infection of the green
 Chytrids reproduce by means of alga Spyrogyra
flagellate spores that locate
preferred host algae through
chemical cues
66
Algal defences against pathogens

 Structural barriers
● Resistant cell wall polymers
 Antibiotic compounds
● Many types of antibacterial and antiviral compounds that
provide resistance against a wide variety of microbes

● May explain why obviously infected seaweeds are rare

67
Algae as parasites or pathogens

 Parasites and pathogens are found in diverse algal lineages

 Many red algae parasites
● ~15% of red algal species is parasitic and infects other red
algae

● In these interactions the nucleus is transferred to a host cell

● Transformed host cells produce reproductive cells that include
the parasitic genome

 The non-photosynthetic green algae Prototheca wickerhamii

and P. zopfii can cause infections of humans and cattle

68
Prototheca
Algae as epibionts

 Another organism serves as substrate

● Algae can be epibionts or serve as epibiontic
surface

 Substrates can be bacterial biofilms, other

algae, protozoa, plants, zooplankton, and
larger animals, such as crabs, fish, turtles,
whales, tree sloths, and polar bears
 Ship fouling is caused by epiphytic algae on
bacterial biofilms
A golden-brown algal
epiphyte (Peroniella sp.)
growing on the green
alga Desmidium majus
69
Algae in mutualistic symbioses

 Wide variety, two species and also consortia (multiple species)

 Bacterial associations
● Surface attachment
● Exchange of nutrients and/or vitamins
 Ulva (sea lettuce) grows abnormally in the absence of bacteria

Epibacteria growing on the surface of the

branched green alga Cladophora glomerata

Epibacteria (blue) growing within the

mucilaginous sheath of the green alga 70

Desmidium grevillii
Fungal associations

 Lichens
● Alga or cyanobacteria are termed phycobiont or photobiont
● Over 17,000 lichens
● ~20% of all fungi can form lichens
● 85% contains green algae, 10% cyanobacteria and 4% both
● Algae fix carbon – fungus fixes nitrogen
● Morphological complexity differs greatly (organisation in layers)

71
Animal associations

 Many marine animals maintain cyanobacterial or dinoflagellate

endosymbionts in their bodies (zooxantallae)
● The animals obtain organic food and oxygen, while the algae
benefit from animal wastes (minerals and CO2)

● Extracellular associations are also common

72
Plant associations

 Liverwort, hornwort,
moss, fern, cycad, and
angiosperm species are
closely associated with
nitrogen-fixing
cyanobacteria
 Associations may have
been important to
obtain Nitrogen during
colonization of land
 Azolla - Anabaena
The water fern Azolla contains cavities with
Anabaena cyanobacteria that fix Nitrogen
73
Applications of algae

 Research
● Evolutionary research
● Environmental monitors - bioindicators
● Paleoecological indicators
 Food
Haematococcus –
● Microalgae for food additives (e.g. astaxanthin) astaxanthin
production
● Porphyra (nori)
● Gelling agents: alginates, carrageenan, agar, agarose
 Pharmaceuticals
 Biofuels
● Not only cellulosic biomass, also H2 production
 Waste water treatment 74
Diversity of evolutionary origin of algae

75
Cyanobacteria
 Most supergroups include algae phyla in
addition to organisms not regarded as algae
Rhizaria, Alveolates and Stramenopiles: SAR
Land plants and algal relatives contain
More related to each other than to other groups
the red and green algae

Cryptomonads: unsure,
probably linked to Land
plants and algal relatives
Haptophytes: unsure,
probably somewhere
with SAR

Discoba contains diverse,

mostly heterotrophic,
protists
76

Amoebozoa and Opisthokonta (animals and fungi) supergroups do not include algae
Sizes of different groups algae

 Cyanobacteria: ~3,300 recognised species

 Euglenoids: >1,100 species
 Red algae: >6,000 species
 Green algae: >8,000 species
 Dinoflagellates: >2,000 species
 Fotosynthetic stramenopiles: >11,000 species
 Haptophytes: ~500 species
 Cryptomonads: ~200 species
 Glaucophytes:~14 species

77
1. Cyanobacteria

 Cyanobacteria form a phylum

within the domain bacteria
 Prokaryotes:
● absence of membrane-bound
organelles (e.g. nuclei and
mitochondria)

● Lack other distinctive features

of eukaryotic cells

78
Cyanobacteria are the only photosynthetic
bacteria

 Contain chlorophyll a and produce oxygen: oxygenic

photosythesis
● Referred to as blue-green algae (Nl. Blauwwieren)
 Body plans: unicells, colonies, filaments
● Coated with mucilage sheaths
● Specialisation in filaments: formation heterocysts that fix
Nitrogen

● Akinetes are specialised resting cells that develop within

filaments during unfavourable conditions

79
 Variety in body plan of cyanobacteria

Unbranched filament - Lyngbya

Colony - Aphanothece

Unicells - Synechocystis

True-branched
form - Stigonema

Unbranched filament
with heterocytes and
akinetes - Anabaena

Aggregation of multiple filaments

in a common sheath (trichome) - False-branched 80

Microcoleus filament - Scytonema

 Some cyanobacteria meet major criteria to define
multicellularity:
● Cells attached to each other
● Specialized cells
● Intercellular communication
● Involvement of
controlled cell death
in development

81
Mucilage secretion by cyanobacteria
commonly occurs

 Mucilage outside the cell wall may have functions in:

● Colony formation
● Attachment to substrates
● Capturing micronutrients from the surrounding water
● Aid buoyancy
● Deter herbivores
● Absorption of UV radiation – allows living in exposed habitats

82
Cyanobacterial cell wall

 Composed of peptidoglycan: carbohydrates linked by pectin

● Enclosed by a lipopolysaccharide envelope
● Cell wall is like any other Gram-negative bacterium, but often
thicker

 Unique for some cyanobacteria:

● S-layer for surface mineral deposits, such as calcium carbonate
● Protein fibrils in some species for gliding movement

83
 Cytoplasmic features
 Thylakoids
● Photosynthetic membranes with
chlorophyll a and other
photosynthetic components

 Carboxysomes
● Rubisco aggregations for carbon
fixation

 Cyanophycin
● Aspartic acid and arginine polymers
– function as nitrogen reserves

 Cyanophycean starch – in cytosol!

● α-1,4-linked polyglucans
 Gas vesicles or gas vacuoles
● For regulating vertical position in the 84

water column
Gas vesicles or gas vacuoles

 Unique for cyanobacteria

 Misleading name: not surrounded by membrane
 Hollow cylinders surrounded by 2nm protein wall
 Induced by low light levels: upward movement
 Counteracted by photosynthesis
● More carbohydrate production -> higher turgor pressure -> gas
compression

 Vertical movements of up to 70 m (!) have been reported

● This movement allows cyanobacteria to outcompete eukaryotic
algae under eutrophic conditions

85
Gas vesicles or gas vacuoles

Gas vacuoles sequentially appear

and disappear depending on the
osmotic condition in the cell

TEM image of gas vacuoles (yellow

arrows) 86
Light harvesting systems in cyanobacteria

 Cyanobacteria absorb light

energy with chlorophyll a
● Some species that largely lack
chlorophyll a possess
chlorophyll d as light
harvesting molecule

 Cyanobacterial accessory
pigments extend the wavelength Much more about light
range of light from which energy harvesting and
can be harvested photosystems in the
● In cyanobacteria: phycobilins, photosynthesis lectures
carotenoids, in some
cases,chlorophyll b

87
 Reproduction of cyanobacteria

 Division by binary fission

 Filamentous cyanobacteria form hormogonia
● Short motile filaments
● Result from controlled death of certain cells: separation disks
or necridia

 No sexual reproduction (meiosis and gamete formation)

 Viral gene transduction by
bacteriophages
● Has been shown to be relevant
for adaptation to habitat variation

Hormogonium (a)
and separation disks 88

(arrows in (b)
Ecological importance of (cyano)bacteria:
Nitrogen fixation

 Bacteria are the only known organisms that can convert N2

gas into ammonia
● Essential for protein production!
 Inducible process, regulated by environmental levels of
ammonium (NH4+) or nitrate (NO3–)

 Nitrogenase is the key enzyme for nitrogen fixation

● Nitrogenase is inhibited by environmental oxygen
 Heterocyte adaptations protect nitrogenase from oxygen
● No active PhotoSystem 2: no oxygen production
● PhotoSystem 1 is active: ATP production
● Heterocytes receive carbohydrates from neighbouring cells for
respiration: oxygen consumption 89
 Cyanobacteria occur in extreme habitats

 Diverse habitats
● Ponds, lakes, rivers, oceans,
temperate soils, geothermal
waters, desert soils and rocks,
polar regions, hypersaline waters

Cyanobacteria in
a hypersaline
salar in the
Atacama desert
of Chile

Thermophile cyanobacteria in
Yellowstone hot springs survive
90
up to 74°C
Evolution of cyanobacteria

 Studied by archeological and phylogenetic (molecular clock)

approaches
● Oldest evidence for oxygenic photosynthesis: 3.4-3 billion
years ago

● Stromatolites: calcareous mounds having layered internal

structure by calcification of sheets of cyanobacteria

● Evidence for colonization of land ~2.6 billion years ago

A fossil stromatolite

91
Evolution of cyanobacteria

 700-800 million years ago: highest number of marine

stromatolites deposited
● Decline afterwards probably caused by evolution of herbivorous
protists, burrowing organisms and competition with eukaryotic
algae

92
 Diversity of cyanobacteria

 Relevant for understanding evolution

of photosynthesis
 Earliest cyanobacteria could probably
not fix Nitrogen and were
thermophyllic
 The earliest diverging lineage of
modern cyanobacteria is Gloeobacter
● Obligate photoautotroph
● Lacks thylakoids, photosynthetic
pigments are associated with the
cell membrane

● Genome has been sequenced

93
Gloeobacter
Unicellular forms lacking specialised cell types

 Synechococcus
● Small cylindrical unicells (1µm in diameter)
● Important primary producer in marine and fresh waters
● Nu mucilaginous sheath
● Cells may exhibit swimming motility
● Cell surface gradually becomes coated with calcium carbonate
and is shed regularly

94
Unicellular forms lacking specialised cell types

 Prochlorococcus
● One of the most numerous components of open-ocean
plankton

● Multiple ecotypes; temperature, irradiance, nutrients

● Natural populations consist of 100-rds of co-existing
populations that have diverged few million years ago

Prochlorococcus
marinus

95
 Colonial forms lacking specialised cell types
 Chroococcus
● Multiple habitats
● Cells adhere after division,
form colonies of 2-32 cells
● Blue-green in colour
 Gloeocapsa
● Cell have distinct sheaths
and multiple cells together
are contained by older
sheaths
96
Colonial forms lacking specialised cell types

 Aphanotheca  Merismopedia
● Diverse, including ● Floating or sedentary
marine, habitats in fresh and marine
waters
● Very large,
macroscopic colonies ● Ovoid cells are
embedded in orderly arranged in
mucilage flat, rectangular
colonies

1 cm

97
Filamentous forms lacking specialised cell types

 Oscillatoria  Trichodesmium
● Diverse, including marine, ● Common in tropical
habitats open-ocean waters
● Can oscillate, rotate and glide ● Sometimes forms
visible blooms
● Disk-shaped cells
● Produces hormogonia by ● Can fix nitrogen but
separation disks lacks heterocytes

98
Spore producing cyanobaceria

 Chamaesiphon  Chroococcidiopsis
● Bud off exospores ● Produces
endospores
● Common inhabitant
of aquatic plant ● Occurs in extreme
surfaces environments (arid,
hot, cold)
● Mostly fresh water
species

Cladophora

 These species are intermediates to 99

cyanobacteria with specialized cell types

Heterocyte and akinete-producing cyanobacteria

 Anabaena
● Primarily planktonic in fresh and marine waters
● Producer of blooms and toxins
● Symbiont of the water fern Azolla anabaena

100
Heterocyte and akinete-producing cyanobacteria

 Most branched and false branched cyanobacteria occur in

fresh water habitats and in peat, rocks, etc.

Tolypothrix; exhibits single false branches

adjacent to heterocytes
Stigonema; true-branches that are
multiseriate (more than one cell in width)
101
Tomorrow

Characteristics and
reproduction of
different groups of
eukaryotic algae

102