SAR IV:
Kingdom Heterokontae (cont’d)
Kingdom Hacrobiae
Lecture 11
Kingdom Heterokontae (or Stramenopila)
Phyla:
1. Opalinata / Opalinida
2. Oomycota (fungus-like)
3. Xanthophyta (yellow-green algae)
Kingdom Hacrobiae (Eukaryomonadae)
Cryprophyta
Phylum Opalinata
highly unusual protists with large cells, multipleshort
flagella, and anywhere from 2 – 100’s of nuclei
Cell surface is arranged in parallel folds (opalescence)
Taxonomic features: number of nuclei and cell form
(cylindrical vs. flattened)
Opalina (mulitinucleate) and Zelleriella spp. are the most
common
Blastocytis – human enteroparasite
The geographical distribution of opalinids is dependent on
the distribution of frogs because they are so closely
associated with their amphibian hosts,
Opalinids seem not to adversely affect their hosts
Phylum Opalinata
Named after the gem opal
display of colours which is seen when white light reflects from
structures on or in gemstones.
for opals, prismatic colours are seen because spectral colours are
separated due to the folds of the plasma membrane
They are quite large (≤ 3 mm) and have multiple
flagella and two to many nuclei
The surface of an opalinid cell is arranged in parallel
folds.
Light striking the pellicle leads to the beautiful
opalescence/iridescence observed when opalinids are
observed in reflected light
Phylum Opalinata
Microaerophilic heterotrophs
Leaf-like or cylindrical cell in outline
Covered by multiple short flagella borne on a
furrowed or folded pellicle
superficially resembling the ciliates
no alveoli
absence of dimorphic nucleation (cf. Ciliata)
lack a cytostome, nourished by pinocytosis
Phylum Opalinata
Commensals of the hindgut of ectotherms (large
intestine & cloaca) of amphibians
also reportedly found in some fish and insects
Individual cells contain multiple nuclei that are
typically conspicuous (Opalina)
of taxonomic importance along with cell cross-sectional
shape.
The flagellar arise from the falx (sickle-shaped
structure) at the anterior end of the vegetative cell
Flagellar activity also arises from the falx
Phylum Opalinata
Since the anuran absorbs the nutrients from its
food in the small intestine, the opalines are
probably not depriving their hosts of nutrients
It is apparent that the opalines are simply living off
the "left-over" nutrients in the faeces
The nutrition of the opalines might be
supplemented by contributions from the rich
bacterial flora which also reside in the cloacal
region of the alimentary canal
Phylum Opalinata
Anuran hosts containing many thousands of opalines
appear to be completely healthy
no obvious irritation or other pathological signs on their
intestinal or cloacal walls.
These microorganisms play an important role in
research in the fields of:
1. cell biology,
2. physiology,
3. life cycle,
4. mutualisms / host-symbiont interactions, as well as,
5. (co)evolutionary studies
Phylum Opalinata
Asexual reproduction via closed mitosis
Cytokinesis cleaves the cell in the anterior-posterior axis.
Sexual reproduction accomplished through anisogamy
Gametes difficult to distinguish
Cysts are formed upon fertilisation which enables migration from
one host to another through faecal transmission.
life cycle alternates between the mature frog and tadpoles.
Four recognised genera of which Opalina and Zelleriella are the
most commonly occurring forms
Generic taxonomy based on
cell shape of the organism in cross-section
the number of nuclei per cell.
Opalina sp. observed in faeces of Bufo japonicus
http://protist.i.hosei.ac.jp/PDB/Images/Opalinata/Opalina.jpg
An opalinid
Pinocytotic vesicle
Zelleriella sp.
Two nuclei
Anterior proboscis
Numerous, short flagella arising from pellicle
Phylum Oomycota
Includes the
1. water moulds
2. downy mildews
Heterotrophic, filamentous protists which absorb their
food from
1. the surrounding water or soil, and/or
2. other organisms
a. Saprotrophy
b. Parasitism
Phylum Oomycota
Ecologically very important
Disease-causing
1. Plant diseases:
a. Blight of potato, Cocoa pod disease – Phytophthora spp.
b. Damping off of seedlings – Pythium spp.; Grapes – Plasmopara sp.
c. White rusts – Albugo spp.
2. Animal diseases:
− Ulcerative Dermal Necrosis (UDN)
− Skin lesions on fish skin & scales – Saprolegnia spp.
Saprophytes
Downy Mildews that appear on leather products during warm & humid
weather conditions
Phylum Oomycota
Formerly placed in Kingdom Fungi:
Similarities include…
1. Cottony mycelial habit, haustoria
2. Heterotrophs: saprotrophy & parasitism
3. Life cycles
– sporangia, spores, oogonia, antheridia
4. "Oomycota" literally means "egg fungi," a clear reference
to the large round oogonia (female gametes)
Oogonium of Saprolegnia sp.
Saprolegnia mycelia &
reproductive structures
Saprolegnia mycelia & fertilized oogonia
Zoospores being liberated Male mycelial fertile tips
zoosporangium
Egg cells within oogonium,
about to be fertilized
Mycelial strands
Diagrammatic
representation of an
oomycete
Sexual Reproduction
Fish infected with Saprolegnia sp.
Saprolegnia infection
Sea Trout with Saprolegnia infection
https://en.wikipedia.org/wiki/Oomycete#/media/File:Seatrout_UDN_saprolegnia.jpg
Sea Trout with Saprolegnia infection
Phylum Oomycota
Some of these also result in plant diseases
Powdery mildews and discoloured rots of plant parts
Phytophthora spp.
P. infestans on Solanaceae (tomato, Irish potato)
P. capsici on a variety of crops
cucumber, squash, melons, pumpkin, pepper, tomato and eggplant and has
recently been described on snap and lima beans
P. palmivora - Black Pod disease (Cocoa), bud rot on palms (Arecaceae)
P. megakarya - Black Pod of Cocoa
Sclerophthora macrospora on Poaceae
corn – crazy top disease
grass – yellow top disease
Others also are saprotrophs, showing up as “downy mildews”
on leather products
Black Pod of Cocoa
P. Capsici on Capsicum annuum
Phytophthora sp.
Phytophthora sp.
Phylum Oomycota
The ultrastructure, biochemistry, and molecular
sequences of these organisms indicate that they belong
with the Kingdom Heterokontae rather than to the
Kingdom Fungi:
1. The free-swimming spores bear two dissimilar flagella,
one with mastigonemes
2. Storage of food energy in the form of
mycolaminarin
3. Their cell walls are made of cellulose cf. chitin in
fungi
4. Their nuclei are diploid and not haploid as in the
fungi
Phylum Oomycota
Reproduction is both asexual and sexual
Asexual reproduction via closed mitosis:
1. Specialized sporangia develop at the ends of the
filaments.
2. Two types of zoospores with heterokont flagella are
produced:
a. 1° zoospores - both flagella originate at the anterior end
b. 2° zoospores – both flagella emerge at the equator of the
zoospore
3. 2° zoospores then germinate to produce a new
individual which develops characteristic
siphonaceous, non-septate ‘hyphal’ filaments
cyst
2° spore
1° spore
Asexual cycle
Sexual cycle
Phylum Oomycota
Sexual Cycle involves
1. The apposition of the swollen tips of two hyphal
filaments, which each behave as an antheridium and
oogonium respectively.
2. Migration of nuclei takes place through a tube from
the ‘antheridium’ to the ‘oogonium’.
3. Following syngamy, the zygote develops into a thick-
walled cellulosic oospore , which behave as a resting
stage
4. The oospores then germinate under suitable
conditions to produce a diploid vegetative filament.
Xanthophyta
Morphology ranges from free-living or attached
unicells → colonies → unbranched or branched
filaments → siphonous forms
Predominantly in freshwater; also, soil, brackish waters
and marine habitats
Chlorophylls a, c1, and c2 and a range of xanthophylls,
but not fucoxanthin - no chl. b (cf. Chlorophyta)
Chloroplasts are yellowish-green, discoidal, parietal
choroplasts
Reproduction sexual and asexual
Yellow-green colour due to β-carotene and
xanthophylls such as violaxanthin, heteroxanthin, and
vaucherioxanthin
Xanthophyta
Primarily freshwater or soil algae
Mostly freshwater
e.g. Vaucheria sp. (type specimen)
Some coccoid but mostly siphonous filamentous form
Multiple discoid chloroplasts in each cell
Heterokont flagella anteriorly inserted in motile antherozoids &
spores
Yellow-green (literally “yellow plant”)
2° chloroplasts
Chls. a & c (low concentration)
No fucoxanthin
Storage CHO is chrysolaminarin
Cell wall impregnated with silica (coarse to the feel)
Xanthophyta
Vaucheria is a relatively common, siphonaceous
xanthophyte
Can be confused as a green alga
the correct placement of the Vaucheria recognized only on the
basis of careful pigment analysis and ultrastructural study
Reproduction is oogamous
Mitosis in Vaucheria involves a long spindle, which is
persistent far into telophase
Xanthophyta – siphonous habit
Xanthophyta
oogonium
antheridium
oogonium
Vaucheria sp.
Xanthophyta
oospores
within Empty
fertilised antheridium
oogonium
Vaucheria sp.
Supergroup SAR cont’d
Very large & diverse group consisting
of four kingdoms:
✓ Kingdom Alveolata
✓ Kingdom Heterokontae / Stramenopila
3. Kingdom Hacrobiae
4. Kingdom Rhizariae
Kingdom Hacrobiae
(Eukaryomonadae)
Kingdom Hacrobiae
Derived from the first two letters of the
dominant taxa:
1. HA (Haptomonada) and
2. CR (Cryptomonada)
3. Bios (life)
There are two main phyla in this Kingdom:
✓ Cryptophyta
✓ Haptophyta
− Centrohelomonada
Kingdom Hacrobiae
A large group of unicells with tremendous
diversity
Important 1° producers and heterotrophs
Particularly abundant in in oligotrophic lakes
and oceans
Recently proposed group
Based exclusively on molecular data since no single character
that unites these groups on the basis of morphology has been
identified
A significant apomorphy is the nucleomorph of these groups
Phylum Cryptophyta
Exclusively unicellular, asymmetrical cell
Leads to unbalanced rotating around the longitudinal axis while
swimming
Significant component of the nannoplankton (2-20 μm)
freshwater & inshore marine communities
Strong adaptability to light and temperature
Sandwich-layered cell envelope
Pellicle on either side of the cell boundary membrane
A subapical invagination associated with
two heterokont flagella, and,
ejectosomes in between the pellicular plates
Nucleus located posteriorly in the cell
Cryptophyta
Ejectosomes within the groove / gullet
discharged when mechanical / chemical stressors irritate the
cells
Grazing
Physical attack
A change in pH, temperature and/or osmotic pressure
composed of a tightly coiled spool of protein
undergo a very rapid, irreversible conformational shift.
projectile pops suddenly out of the cell, causing the cell to recoil
as a result
defense mechanism or
function in predation
Phylum Cryptophyta
Autotrophic : Heterotrophic taxa ~ 1:1
Even some autotrophic ones ingest prey (mixotrophy)
Chloroplasts
4 - membraned (3°) chloroplast with CER
Two different light harvesting complexes:
1. Chlorophylls a and c2 on the outer membranes of the thylakoids
2. Phycobiliproteins in the lumen of the thylakoid
No phycobilisomes (as in CNB, Glaucophyta & Rhodophyta)
Abundant β–carotene, fucoxanthin and other xanthophylls
Pyrenoid within chloroplast
Nucleomorph located between the CER & chloroplast envelope
periplastidial space containing
starch grains, 80S ribosomes and a nucleomorph (~ 3 chromosomes)
Phylum Cryptophyta
four genomes in cryptophyte cells
✓ Host nucleus genome
✓ Mitochondrial genome
✓ Plastid genome
✓ Nucleomorph genome
Storage
Starch in the periplastidial space
Oil droplets in the cytoplasm
Some, such as Cyanomonas, have endosymbiotic cyanobacteria
(cyanelles) which function as chloroplasts
Mitochondria with flat cristae
Can form major blooms with devastating effect on wildlife
e. g. Cyanomonas is also bloom-forming → toxins
Cryptophyta
Nucleomorphs of certain cryptomonads can be
isolated
The nucleomorph gene sequences are highly
divergent from those of the host nucleus
Support the hypotheses that the nucleomorph is the
remnant of a foreign nucleus
Most of the plastid proteins are coded for by the host
nucleus with a minority coded for by the
nucleomorph
Cryptophyte DNA Compartments
Chloroplast*
(remnant nucleus of red algal
origin)
Cryptophyta
Reproduction is generally asexual
but sexual reproduction has recently been documented
Colonial:
Some form sessile ("palmelloid") stages (Dinobryon sp.)
Colonial form – (Synura sp.)
Large nucleus at central to posterior end
Mitosis is semi-open
Partial breakdown of nuclear envelope with the division
spindle at the cell poles, spindle persistent into telophase
Cytokinesis: proceeds longitudinally from the rear along cell
axis, starting from posterior end (cf. Euglenophyta)
Cryptomonas sp.
Cryptomonas sp.
Chroomonas sp.
Generalized structure of a cryptophyte
Contractile
vacuole
ejectosomes
invagination
ribosomes
chloroplast envelope Rhizostyle (microtubules)
thylakoid membrane mitochondrion
nucleomorph
oil droplet
Starch grain
Periplastidial space Golgi apparatus
Nucleus with nucleolus
CER
Generalized structure of a cryptophyte
Cryptophyta – Cell boundary
A periplast
sandwich-layered cell boundary
consists of proteinaceous layers on either side of the
plasma membrane
Morphology of a cryptophyte
cell:
The periplastidial space
Nucleomorphs
Cryptophyte plastid in Guillardia theta Chlorarachniophyte plastid in Bigelowiella natans
Crytophyta taxonomy
1. Chrysomonads
Cellulosic, freshwater forms
Loricate forms
2. Silicomonads
Marine silicated forms with silicaceous scales, spines
or bristles external to the plasma membrane
Encystment to form statospores → fossil
record
Cryptophyta taxonomy
Three orders:
1. Goniomonadales: obligate heterotroph, phagocytic
– colourless, no plastids: Goniomonas
2. Cryptomonadales: strictly photoautotrophic
– reddish protoplasm w/ Cr-phycoerythrin: Cryptomonas
3. Chroomonales: osmotrophic photoautotrophs (i.e.,
mixotrophic)
– Cells blue-green in colour w/ Cr-phycocyanin: Chroomonas
