Heterokonts are a group of protists
(formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryote
Most are algae
, ranging from the giant multicellular kelp
to the unicellular diatom
s, which are a primary component of plankton
. Other notable members of the Stramenopiles
include the (generally) parasitic oomycete
s, including ''Phytophthora
'' of Great Famine
of Ireland infamy and ''Pythium
'' which causes seed rot and damping off.
The name "heterokont" refers to the type of motile life cycle stage, in which the flagellated cells
possess two differently arranged flagella
In 1899, Alexander Luther created "Heterokontae" for some algae with unequal flagella, today called Xanthophyceae
. Later, some authors (e.g., Copeland, 1956) would include other groups in Heterokonta, expanding its sense. The term continues to be applied in different ways, leading to Heterokontophyta being applied also to the phylum Ochrophyta
. The term 'Stramenopile' was introduced in 1989 by Patterson to overcome ambiguities that had (and continue to be) developed with the use of the term 'heterokont'. Consequently, heterokonts may be referred to as stramenopiles.
The term 'heterokont' first emerged in the context of 19th century phycology. Over time, the scope of application has changed; especially when in the 1970's as ultrastructural studies revealed greater diversity among the algae with chromoplasts (= chlorophylls a and c) than had previously been recognized. At the same time, a protistological perspective was replacing the 19th century one based on the division of unicellular eukaryotes along inappropriate botanical/zoological lines. One consequence was that an array of heterotrophic organisms, many of which had not been previously considered as 'heterokonts' were seen as being related to the 'core heterokonts' (i.e. those having anterior flagella with stiff hairs). Newly recognized relatives included the parasitic opalines, proteromonads, and actinophryid heliozoa. They joined other heterotrophic protists, such as bicosoecids, labyrinthulids, and oomycete fungi, that were included by some as heterokonts and excluded by others. Rather than continue to use a name whose meaning had changed over time and was hence ambiguous, the name 'stramenopile' was introduced to refer to the clade of protists that had tripartite stiff (usually flagellar) hairs and all their descendents. Molecular studies confirm that the genes that code for the proteins of these hairs are exclusive to stramenopiles. As the concept of 'Stramenopile' is based on a presumed apomorphy, it is stable and robust even when its composition changes. There is a widespread presumption, as here, that the terms 'stramenopile' and 'heterokont' are synonyms. They are not because they are defined differently and despite compositional overlap, most applications of the names imply differing compositions.
Many heterokonts are unicellular flagellate
s, and most others produce flagellated cells at some point in their lifecycles, for instance as gamete
s or zoospore
s. The name heterokont now refers to the characteristic form of these cells, which typically have two unequal flagella. The anterior flagellum is covered with one or two rows of lateral hairs or mastigonemes
, which are tripartite (i.e. with a flexible basal insertion, a stiff hollow component, and tipped with fine delicate hairs ), while the posterior flagellum is smooth, and usually shorter, or sometimes reduced to a basal body. The flagella are inserted subapically or laterally, and are usually supported by four microtubule
roots in a distinctive pattern. Opalines have many rows of flagella which do not have flagellar hairs. They are closely related to endosymbiotic proteromonad flagellates some of which have tripartite hairs extending from the body surface.
Mastigonemes are manufactured from glycoprotein
s in the endoplasmic reticulum
before being transported to the anterior flagellar surface. When the hairy flagellum beats, the stiff mastigonemes are forced backwards and this creates a retrograde current, pulling the cell through the water or bringing in food.
The term mastigonemes refers to various types of flagellar hairs, but those of stramenopiles have a distinctive structure. It was treated as the evolutionary innovation that defined the stramenopiles, and although not initially so, it is increasingly treated as the defining characteristic of the heterokonts. Mastigonemes have been lost in a few heterkont lines, most notably the diatoms, opalines, and actinphryid heliozoa.
Many heterokonts are algae with chloroplast
s surrounded by four membranes, which are counted from the outermost to the innermost membrane. The first membrane is continuous with the host's chloroplast endoplasmic reticulum
, or cER. The second membrane presents a barrier between the lumen
of the cER and the primary endosymbiont or chloroplast, which represents the next two membranes, within which the thylakoid
membranes are found. This arrangement of membranes has led to the hypothesis that heterokont chloroplasts were obtained from the reduction of a symbiotic red algal
eukaryote, which had arisen by evolutionary divergence from the monophyletic primary endosymbiotic ancestor that is thought to have given rise to all eukaryotic photoautotrophs
. The chloroplasts characteristically contain chlorophyll a
and chlorophyll c
, and usually the accessory pigment fucoxanthin
, giving them a golden-brown or brownish-green colour. Because of this colour, they are referred to as 'chromoplasts' distinguishing them from chlorophyll B containing plastids of green algae, their descendants the higher plants, and euglenids.
Most basal heterokonts are colorless. This suggests that they diverged before the acquisition of chloroplasts within the group. However, fucoxanthin-containing chloroplasts are also found among the haptophyte
s. This led to hypotheses that all organisms with chlorophylla/c containing chloroplasts have a common phylogenetic
history with cryptomonad
s, and should be taxonomically grouped as the Chromista
. Molecular studies do not confirm that the stramenopiles, haptophytes, and cryptomonads are sister taxa. The current consensus is that the ancestral stramenopiles / heterokonts were heterotrophic and acquired chloroplats after their defining feature (the tripartite hairs) appeared.
As noted above, classification varies considerably. Originally, the heterokont algae was used only for Xanthophytes. The concept morphed to include more lineages and considered by some as part of the kingdom Plantae and later, by others, as within the Protista. An example is:
::Class Chrysophyceae (golden alga
::Class Bacillariophyceae (diatom
:Division Phaeophyta (brown alga
In this scheme, the Chrysophyceae is conceived of a very extensive group (Chrysophyte ''sensu lato'') that was paraphyletic
- as the diatoms and brown algae evolved from within the chrysophytes. Over time, various lineages have been given their own classes and often divisions. Recent systems often treat these as classes within a single division, called the Heterokontophyta, Chromophyta, or Ochrophyta. This is not universal, however; Round ''et al.'' treat the diatoms as a division.
The discovery that oomycete
s and hyphochytrid
s are related to these algae, rather than fungi, as previously thought, has led many authors to include these two groups among the heterokonts. Should it turn out that they evolved from colored ancestors, the heterokont group would be paraphyletic in their absence. Once again, however, usage varies. David J. Patterson
named this extended group the stramenopiles, characterized by the presence of tripartite mastigonemes, mitochondria
with tubular crista
e, and open mitosis
. He used the stramenopiles as a prototype for a classification without Linnaean rank. Their composition has been essentially stable, but their use within ranked systems varies.
treats the heterokonts as identical in composition with the stramenopiles; this is the definition followed here. He has proposed placing them in a separate kingdom, the Chromalveolata
, together with the haptophytes, cryptomonads, and alveolates. This is one of the most common revisions to the five-kingdom system
, but has not been adopted, because Chromalveolata is not a monophyletic group. A few treat the Chromalveolata as identical in composition with the heterokonts, or list them as a kingdom Stramenopila.
Some sources divide the heterokonts into the autotrophic Ochrophyta
and heterotrophic Bigyra
However, some modifications to these classifications have been suggested.
The name Heterokonta can be confused with the (much older) name Heterokontae, which is generally equivalent to the Xanthophyceae
, a limited subset of the Heterokonta.
The simplified classification of the group according to Adl ''et al.'' (2012), in which heterokonts (using the term Stramenopiles) are part of a larger clade that embraces Alveolates
** Stramenopiles , 1989, emend. ''et al.'', 2005
, 1926, emend. , 1997 (Slopalinida
'' , 1911
, 1926, emend. , 1998
''et al.'', 2002
, 2001 (Öomycetes
, 1897, emend. , 1976)
1874, emend. 1926
'' , 1999 (Bolidophyceae
in et al., 1999)
, in ''et al.'', 1998
''et al.'', 2003
, 1950, emend. , 1980
, 1930, emend. , 1935 (Heterokontae
, 1899, Heteromonadea
, 1983, Xanthophyta
, 1886 (not , 1891, not , 1894)
'' ''et al.'', in ''et al.'', 2003 (Schizocladales
''et al.'', 2003) (M)
, 1821 (= Bacillariophyta
Based on the following works of Ruggiero et al. 2015 & Silar 2016.
File:Four_common_forms_of_Blastocystis_hominis_Valzn.jpg|''Blastocystis hominis'' (Blastocystea)
File:Parasite140015-fig2 Protoopalina pingi (Opalinidae) Microscopy.tif|''Protoopalina pingi'' (Opalinea)
File:Peronospora sparsa.JPG| ''Peronospora sparsa'' (Oomycetes)
File:Aardappel Phytophthora Fresco.jpg|Potatoes with ''Phytophthora'' (Oomycetes)
File:Diatomeas w.jpg|Diatoms (Bacillariophyceae)
File:Dictyocha speculum.jpg|Silicoflagellate (Dictyochophyceae)
File:Actinophrys sol.jpg|''Actinophrys sol'' (Actinophryida)
File:15 3klein2.jpg|''Nannochloropsis'' sp. (Eustigmatophyceae)
File:Dinobryon sp.jpg|''Dinobryon'' sp. (Chrysophyceae)
File:Synura.jpg|''Synura'' sp. (Synurophyceae)
File:Gonyostomum-cells.JPG|''Gonyostomum semen'' (Raphidophyceae)
File:Pacific rockweed, Olympic National Park, USA.jpg|''Fucus distichus'' (Phaeophyceae)
File:FMIB_48487_Pelagophycus_Porra,_or_Elk_Kelp.jpeg|''Pelagophycus porra'' (Phaeophyceae)
Tree of Life Web Project: Stramenopiles