SYSTEMATIC BIOLOGY

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PHYLUM OOMYCOTA

INTRODUCTION TO THE OOMYCOTA

Oomycota (o-o-mi-KO-ta) is derived from two Greek roots that mean egg (oario -ωάριο) and fungus (mykes -μύκης).  The reference is to the fungus-like habit of the organism and its typical oogamous sexual reproduction. 

The water molds are among the most economically important organisms in the heterokonts.  Although some like Saprolegnia (Figure A) are used for the commercial production of acetone and other small organics, most are important because of the damage that they do to agriculture.  The most notable outcome of economic importance was the great potato famine of Ireland (1845-1850) caused by Phytophthora (Figure B)The famine began in September of 1845 with the leaves of potato plants appearing blackened.  Soon, however, the feeding filaments (hyphae) had coursed through the plants, which turned to stinking, nauseous masses of slime.  The spores released by the plants infected in the initial assault spread over the island of Ireland, and the crop of potatoes, the principle food crop, failed.  The potato blight persisted through the next four years, a time called The great Hunger during which as many as 5 million may have died from starvation.  Other members of the oomycotes cause the familiar downy mildew of Lilac.

Hyphochytridium (Figure B) is in a group that is reduced in form and resembles a chytrid, but only superficially so.  Unlike other microbial eukaryotic filamentous organisms, these tend to be diploid with a haploid phase that is reduced or restricted to the gametes.  

Saprolegnia_cave-utoronto.jpg (45933 bytes)

A. Oogonia of Saprolegnia.

phytophthora-gc-ca.jpg (22103 bytes)

B. Labeled photomicrograph of Phytophthora.

hyphochytridium-bsu.jpg (52436 bytes)

C. Hyphochytridium grown from a spore in 24 hrs from an infected pollen grain.

Images taken from:
A: http://www.botany.utoronto.ca/ResearchLabs/MallochLab/Malloch/Moulds/Classification.html
B: http://www.inspection.gc.ca/english/plaveg/pestrava/phyram/images/fig17.jpg
C: http://www.bsu.edu/classes/ruch/msa/barr.html

SYNOPTIC DESCRIPTION OF THE OOMYCOTA

The following description comes from Margulis and Schwartz (1988 and 1998), Dick (1990), Fuller (1990), Barr and Desaulniers (1989), and Beakes (1989).

I. SYNONYMS: Oomycetes, water molds, phycomycota.

II. NUMBER: >800 species.

III. PHYLUM CHARACTERISTICS:

A. Structure and Physiology

Cell Form: Unicellular to siphonaceous filaments (with or without crosswalls).

Flagella: Zoospores with one anteriorly-directed tinsel flagellum or with two flagella, one anteriorly-directed tinsel flagellum and a recurrent whiplash flagellum.

Basal bodies: The two basal bodies lie at 180o in the oomycetes; single functional basal body attached to "centriole" in hyphochytridiomycetes.

Cell covering: Cell covering of cellulose and/or chitin.

Chloroplasts: Not present.

Food reserves: Not known.

Mitochondria: Tubular cristae.

Golgi: Present.

Nucleus: Diploid.

Centrioles: Present and outside nucleus.

Inclusions and ejectile organelles:

Not present.

B. Mitosis, Meiosis and Life History

Mitosis: Closed or open with polar fenestrae.

Meiosis: Gametic.

Sexual reproduction and life history:  Isogamous to oogamous.  Life histories of Saprolegnia and Phytophthora are representative of the phylum.

C. Ecology: Parasitic or saprobic in water or soil.

SYSTEMATICS OF THE OOMYCOTA

Note that the Hyphochytrids (Pr-25 and Pr-21) and Oomycetes (Pr-27 and Pr-20) are treated as separate phyla by Margulis and Schwartz (1988 and 1998, respectively), Dick (1990) and Fuller (1990). Indeed, Fuller (1990) explores the reasons for lumping or separating them and concludes that the similarities in lysine synthesis and mitochondria cannot counterbalance differences in cell wall chemistry, mitosis, centrioles and rRNA. Perhaps, such differences do not really occur.  Patterson (1999) places them together in the Stramenopiles (Heterokonts).  Indeed, the tree of Sogin and Patterson (Tree of Life Project) has the two groups closely linked on the same clade within the heterokonts.  Barr and Desaulniers (1989) further suggest that the similarities in their flagellar structures warrant their association.  I have followed their lead and lumped both groups into a single phylum, Oomycota.

Beakes (1989) reports that the oomycetes have ultrastructural and biochemical affinities with the brown algae and eustigs as well as obvious (but perhaps misleading) morphological similarities with the xanthophyte, Vaucheria. Dick (1990) agrees that the siphonaceous form in the Oomycota is not only convergent with that of the Xanthophyta but highly advanced within the Oomycota.  Still, they have obvious ties to other members of the heterokonts.

HIERARCHICAL CLASSIFICATION OF THE OOMYCOTA

TAXONOMY OF THE PHYLUM OOMYCOTA. This system is a modification of Dick (1990), Barr and Desaulniers (1989),  and Beakes (1989).

CLASS OOMYCOTEA

Holocarpic or eucarpic; zoospores with anterior tinsel flagellum and recurrent whiplash flagellum; two types of zoospores may be produced in succession, a primary zoospore with both flagella originating at the anterior end and a secondary zoospore with the two flagella originating laterally; zoospores germinate to produce a body with cellulose and glucan walls, with siphonaceous filaments (called hyphae) coarse and non-septate; nuclear division closed, with persistent nucleolus and intranuclear spindle with poles near pairs of centrioles oriented at 180o to each other; diploid with sexual reproduction by apposition of two hyphae cut off by septa at their tips, followed by a migration of nuclei through channels from one (differentiated as an antheridium) to the other (differentiated as an oogonium), and the formation of thick-walled (chitinous) oospores; oospores develop by the release of zoospores; parasitic and free-living. This class has 4 orders.

ORDER SAPROLEGNIALES

Holocarpic or eucarpic with broad hyphae forming an extensive "mycelium"; zoosporangia long and cylindrical; zoospores often dimorphic, sometimes only secondary zoospores present; oogonia usually contain several oospheres (eggs). The water molds.

Saprolegnia, Achlya, Dictyuchus, Leptolegnia, Aphanomyces.

ORDER LEPTOMITIALES

Small group of water molds with constrictions of the wall which can become blocked giving the appearance of septa, otherwise it is very similar to SAPROLEGANES.

Leptomitus, Apodachlya.

ORDER LAGENIDALES

Small holocarpic body of 1 or a few cells; zoosporangia may release a vesicle in which secondary zoospores are formed; 1 oosphere per oogonium.

Lagenidium.

ORDER PERONOSPORALES

Eucarpic; zoosporangia usually globular, produced on special hyphae, detachable as a wind-blown distributive phase; zoosporangia may also function as conidia, germinating directly to produce a germ tube or release secondary zoospores; 1 oosphere per oogonium; mostly terrestrial species in soil or on vascular plants. The downy mildews.

Peronospora, Albugo, Phytophthora, Pythium, Plasmopara.

CLASS HYPHOCHYTRIDEA

Holocarpic or simple eucarpic, with walls of chitin and sometimes also cellulose; zoospores with only a single anterior tinsel flagellum and a barren basal body, formed within an inoperculate sporangium or in a plasmodial mass released from a zoosporangium; nucleus at mitosis perforated at the poles and with centrioles in orthogonal pairs; gametes similar, flagellate; sexual or asexual resting spores that release zoospores on germination; parasitic or saprobic; freshwater or marine. A single order, HYPHOCHYTRIALES, with characters of the class.

Hyphochytrium, Rhizidiomyces, Anisolpidium, Canteriomyces, Latrostium.


This page is maintained by Jack R. Holt.  Last revised: 02/20/2008.