Oomycota is a class of economically important heterokontophyte algae referred to as "water moulds.

" Because of their parasitic and saprophytic lifestyles, coupled with their obvious lack of chloroplasts, the water moulds were originally classified as a group of (mostly) aquatic fungi. Eventually, they were reclassified as protists because of the facts that their cell walls are composed of cellulose, and not chitin, and that their nuclei are diploid and not haploid. Detailed examinations of their cellular anatomy, as well as DNA comparisons, conclude that these organisms are related to the diatoms and brown algae.

An oomycote is a filamentous organism that grows on/within its substrate and/or host in a manner reminescent of mold. Because it is an alga, one refers to the entire organism as a "thallus". The oomycotes are economically important, as the group contains pathogenic species that attack several important crops. The most famous water mould is the bane of potato agriculture, the infamous Potato Blight, Phytophthora infestans. Plant diseases result in billions of dollars in damage to agricultural crops each year. One of the groups of organisms that cause many serious plant diseases has long been known as the Oomycota or oomycetes, traditionally classified in the phycomycetes or lower fungi. The phycomycetes are an informal group that, in addition to the Oomycota, has historically included such diverse organisms as the slime molds, chytrids, zygomycetes or bread molds, and arbuscular mycorrhizae. The higher fungi have traditionally included the ascomycetes and the basidiomycetes. Examples of ascomycetes, or sac fungi, include the discomycetes (e.g. morels), pyrenomycetes (including the cause of chestnut blight), deuteromycetes or imperfect fungi, and yeasts. The basidiomycetes, or club fungi, include rust fungi, smut fungi, mushrooms, gasteromycetes, polypores, and jelly fungi. However, over the past three decades knowledge about relationships among groups of fungi has increased greatly such that we now realize that many of the traditional morphology-based groupings no longer reflect phylogenetic relationships. The Oomycota or Peronosporomycota consist of more than 800 species that may be saprobic or parasitic on terrestrial or aquatic plants and animals. One member of the Oomycota has greatly influenced history, namely Phytophthora infestans, the cause of late blight of potatoes. As a result of the famine in Ireland caused by this disease, about 1 million people died and another 1.5 million emigrated (Alexopoulos, et al. 1997). A number of other plant diseases are caused by species of Phytophthora, including sudden oak death and ramorum blight caused by P. ramorum, cacao black pod (P. megakarya), and many root rot diseases, such as black shank of tobacco. Other members of the group include pathogens such as Pythium aphanidermatum, the cause of cottony blight of turf grasses; Peronospora tabacina (tobacco blue mold); Plasmopara viticola (downy mildew of grape); Plasmopara halstedii (sunflower downy mildew); and many others. Finally, a group traditionally placed in the oomycetes is the Saprolegniales, or water molds, which cause diseases of fish and other aquatic vertebrates.

Characteristics of the Oomycota

The Oomycota have long been considered fungi because they obtain their nutrients via absorption and many of them produce the filamentous threads known as mycelium

characteristic of many fungi. The Oomycota now are classified as a distinct group based on a number of unique characteristics (Table 1). All members of the Oomycota undergo oogamous reproduction, meaning that diploid oospores are produced as zygotes following fertilization of haploid oospheres by haploid gametes. These oospores may be large and solitary (Figure 1) or smaller and numerous inside the oogonium. None of the true Fungi produce oospores.

Table 1. Major distinctions between the Oomycota in the Chromista and the true Fungi (Chytridiomycota, Glomeromycota, Zygomycota, Ascomycota, Basidiomycota) Character Oomycota Heterogametangia. Fertilization of oospheres by nuclei from antheridia forming oospores. True Fungi Oospores not produced; sexual reproduction results in zygospores, ascospores or basidiospores

Sexual reproduction

Nuclear state of vegetative mycelium Cell wall composition Type of flagella on zoospores, if produced Mitochondria

Diploid

Haploid or dikaryotic

Beta glucans, cellulose Heterokont, of two types, one whiplash, directed posteriorly, the other fibrous, ciliated, directed anteriorly With tubular cristae

Chitin. Cellulose rarely present If flagellum produced, usually of only one type: posterior, whiplash With flattened cristae

Another distinction is the cell wall composition. In the Oomycota, the cell walls are composed of beta glucans and cellulose rather than chitin as in the true Fungi. In addition the Oomycota produce motile zoospores with two kinds of flagella, one of which is a whiplash flagellum oriented posteriorly while the other is called a tinsel flagellum, which has a fibrous, ciliated structure and is oriented anteriorly (Figure 2). Behavior of zoospores of Phytophthora is illustrated in a video at this link. The occurrence of two kinds of flagella places these organisms in a group known as heterokonts. Although some true Fungi, namely the Chytridiomycota, produce stages with motile zoospores, their flagella are only of one kind, the posterior whiplash type.

Figure 1

Figure 2 A fourth major difference between the Oomycota and the true Fungi is that the vegetative cells of the Oomycota generally consist of coenocytic hyphae (hyphae without septa, i.e., without cross-walls), which contain diploid nuclei -- these organisms exist primarily in a diploid state. This is unlike the true Fungi in which most of the mycelium is divided into cells by cross-walls, with each cell containing one, two, or more haploid nuclei. An exception are the Zygomycota, true Fungi with haploid nuclei but which produce coenocytic hyphae. Some members of the Oomycota produce unicellular thalli, a characteristic they share with some kinds of true Fungi. Studies with the transmission electron microscope (TEM) in the late 1970s revealed differences in ultrastructural characteristics between the Oomycota and true Fungi. One distinguishing characteristic is that members of the Oomycota have mitochondria with tubular cristae and protoplasmic and nuclear-associated microtubules, while the true Fungi have flattened mitochondrial cristae. As more groups of organisms were examined using the TEM a relationship was hypothesized between the Oomycota and the heterokont algae (Table 1). This was the first modern evidence that the Oomycota should be regarded as colorless algae rather than true Fungi. Interestingly, as noted in Alexopoulos et al., 1997, about 150 years ago Pringsheim (1858) suggested, in reference to the Saprolegniales [water molds], that the oomycetes were allied with certain algae.

Relationships of the Oomycota based on Molecular Results

As new tools for determining phylogenetic relationships were developed, especially those using molecular sequence data, they were applied to questions such as whether the Oomycota are more closely related to the heterokont algae or the true Fungi. Determining relationships using DNA sequence data is based on comparing sequence similarities in gene regions such as small or large subunits of the nuclear ribosomal DNA. Each base pair is regarded as a

character and the sequence of base pairs is aligned such that analogous characters are compared. Mathematical algorithms running on powerful computers are used to analyze changes among the thousands of base pairs of gene regions. By determining how many changes have occurred between different groups of organisms, it is possible to estimate relative distance among groups and to develop an evolutionary tree that reflects those relationships. Results from a number of studies using molecular sequence data, combined with the ultrastructural similarities, confirm unequivocally that the Oomycota share a common ancestor with the other members of the heterokont algae or Chromista (Figure 3). In addition to the Oomycota, two smaller groups, the Hyphochytridiomycota and Labyrinthulomycota appear to be most closely related to the Chromista. The Chromista include several kinds of algae, namely the Phaeophyta or brown algae, Xanthophyta or yellow-green algae, Chrysophyta or golden algae, and Bacillariophyta or diatoms, as well as several smaller groups. The heterokont algae are distinctive among the algae in having the same two kinds of flagella as occur in the Oomycota, tubular mitochondrial cristae, and other ultrastructural similarities. The photosynthetic members of the Chromista possess chlorophyll c and other pigments that are not found in any other group. Some controversy still remains about exactly what to call this group of organisms. Most authors refer to the kingdom Chromista, phylum Heterokonta, while others place them in the kingdom Straminipila (sometimes written as Stramenopila, Patterson & Sogin, 1992).

Figure 3

True Fungi or Eumycota

The true Fungi, or Eumycota, are now restricted to five major groups, each of which is regarded as a phylum in the Kingdom Fungi. One of these groups is the Chytridiomycota or chytrids. In some ways the chytrids are similar to the Saprolegniales in the Oomycota in that they are regarded as water molds. They feed on small-celled organisms and debris in aquatic environments. Another group of true Fungi is the Zygomycota or zygomycetes that include the bread molds (Mucorales). Their mostly non-septate mycelium gives them a superficial resemblance to the Oomycota. The arbuscular mycorrhizae were once regarded as part of the Zygomycota because they produce what appear to be zygospores. However, molecular data have shown that these root-associated fungi are quite distinct and should be regarded as their own phylum, the Glomeromycota. These fungi are ancient in origin, known from the fossil record to be associated with primitive land plants from the Devonian (Remy et al., 1994).

The largest and most well known of the true Fungi are the Ascomycota or ascomycetes and the Basidiomycota or basidiomycetes. The Ascomycota include most of the lichenized fungi, yeast fungi, morels, cup fungi, and many kinds of microfungi such as pyrenomycetes and their asexual states, also referred to as mitotic or anamorphic states. In the past these have been variously referred to as asexual fungi, deuteromycetes or fungi imperfecti. These asexually reproducing ascomycetes include the hyphomycetes and coelomycetes many of which can now be placed among their relatives in the Ascomycota. The Basidiomycota or basidiomycetes include the rust and smut fungi most of which are obligate plant pathogens. Mushrooms, gasteromycetes, jelly fungi and polypores are also members of the Basidiomycota. A recent noteworthy discovery is that the true Fungi are more closely related to animals than to plants (Baldauf & Palmer, 1993; Wainright et al., 1993; Figure 3), thus explaining why it is so difficult to develop antibiotics that are effective against human fungal pathogens without deleterious effects on humans.

The Remaining non-Fungi

A few other groups of organisms previously regarded as fungi are now known to belong outside the true Fungi (Figure 3). The Plasmodiophoromycota, including Plasmodiophora brassicae, cause of cabbage club root, have proven difficult to place in the tree of life but they are not related to the true Fungi. Some evidence suggests that they too share a common ancestor with the Chromista but this is not conclusive. For some years it has been known that the Myxomycota or true slime molds, the Dictyosteliomycota or dictyostelid cellular slime molds, and the Acrasiomycota or acrasid cellular slime molds are not related to the true Fungi. Recent studies suggest that the Myxomycota represent an independent evolutionary lineage that diverged prior to the emergence of the crown groups of organisms that include the Fungi, Animalia, Plantae, Chromista, and Alveolates (Rhodophyta).

Conclusion
Most mycologists have not abandoned the study of the Oomycota and still define the organisms they study as eukaryotic, heterotrophic osmotrophs in which assimilation takes place through a cell wall (Dick, 1997). Adaptations of these organisms to obtaining their nutrients by absorption have resulted in considerable morphological convergence among them as exemplified by the similarity of the oomycetous white rusts to the uredinia of the true rust Fungi. Understanding the evolutionary relationships among these groups of organisms contributes greatly to our ability to develop strategies to control the diseases these organisms cause.

Literature Cited
Alexopoulos, C.J., C.W. Mims, and M. Blackwell. 1997. Introductory Mycology. John Wiley & Sons, Inc., New York, NY, USA. Baldauf, S.L., and J.D. Palmer. 1993. Animals and fungi are each other's closest relatives: congruent evidence from multiple proteins. Proceedings of the National Academy of Sciences U.S.A. 90:11558-62.

Dick, M.W. 1997. Fungi, flagella and phylogeny. Mycological Research 101: 385-94. Patterson, D.J., and M.L. Sogin. 1992. Eukaryote Origins and Protistan Diversity. Pp. 13-46. In: The Origin and Evolution of Prokaryotic and Eukaryotic Cells. Eds. H. Hartman and K. Matsuno. World Scientific, Singapore. Pringsheim, N. 1858. Beitrge zur Morphologie und Systematik der Algen II. Die Saprolegnieen. Jahrbuch fr Wissenschaftlichen Botanik 1:284-304. Remy, W., T.N. Taylor, H. Hass, and H. Kerp. 1994. Four hundred-million-year-old vesicular arbuscular mycorrhizae. Proceedings of the National Academy of Sciences U.S.A. 91: 118413. Wainright, P.O., G. Hinkle, M.L. Sogin, and S.K. Stickel. 1993. Monophyletic origins of the Metazoa: an evolutionary link with fungi. Science 260:340-2. Oomycota (o-o-mi-KO-ta) is derived from two Greek roots that mean egg (oario ) and fungus (mykes -). The reference is to the fungus-like habit of the organism and its typical oogamous sexual reproduction. Winter defined the class, Oomycetes, we gave it the proper ending for a phylum of a fungus-like organism. In addition, the description of this group is broader than the original Oomycetes. This description includes hyphochytrids and several heterokont taxa with close associations with the Oomycetes.

INTRODUCTION TO THE OOMYCOTA The water molds are among the most economically important organisms in the heterokonts. Although some like Saprolegnia (Figure 1) 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 2). 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. The life cycles of the Oomycota generally are eucarpic, but some may be holocarpic. Details of the life histories can best be seen in Figure 3, Life Cycle of Saprolegnia. Note that there is an asexual cycle (Figure 3-A-H) and a sexual cycle (Figure 3-I-O). In the asexual cycle, the organism produces specialized sporangia (Figure 3-B) from which emerge zoospores with typical heterokont flagella (an anterior tinsel flagellum and a recurrent whiplash flagellum; see Figure 3-C). Notably, they have two types of zoospores which may be produced in succession: a primary zoospore (Figure 3-C) with

both flagella originating at the anterior end and a secondary zoospore (Figure 3-F) with the two flagella originating laterally. Zoospores then germinate to produce a body with cellulose and glucan walls (Figure 3-H and -A). The filaments are siphonaceous filaments (called hyphae), coarse, and non-septate. Nuclear division is closed, with a persistent nucleolus and an intranuclear spindle with poles near pairs of centrioles oriented at 180o to each other. In the vegetative state, they are diploid organisms with gametic meiosis. Sexual reproduction begins with the apposition of two hyphae that are cut off by septa at their tips (Figure 3-J-K). Figure 3-L illustrates the migration of nuclei through channels from one (differentiated as an antheridium) to the other (differentiated as an oogonium). Following fertilization, the zygote develops into a thick-walled (chitinous) oospore (Figure 3-M-N). The oospores are resting stages that germinate by the production of a diploid vegetative filament (Figure 3-O).

FIGURE 1

FIGURE 2

FIGURE 3. Life Cycle of Saprolegnia. A-H is the asexual cycle and I-O is the sexual cycle. See the text for a detailed explanation of the stages. Image taken from Bold et al. (1987)

FIGURE 4

FIGURE 6

THE HYPHOCHYTRIDS

Hyphochytridium (Figure 4) is in a group that is reduced in form and resembles a chytrid, but only superficially so. These organisms range from parasitic to saprobic and can be found in freshwater or marine environments. Like Saprolegnia, these tend to be diploid with a haploid phase that is reduced or restricted to the gametes. They tend to be holocarpic and have walls of chitin and sometimes also cellulose. In the asexual cycle, zoospores have 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. During mitosis the nucleus is perforated at the poles with centrioles in orthogonal pairs. During the sexual cycle gametes are motile and similar (isogametes or anisogametes). As in Figure 3, the hyphochytrids make both sexual and asexual resting spores. However, upon germination, both spores release zoospores.

FIGURE 7

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. Most of the recent molecular phylogenetic analyses (e.g. Andersen 2004; Brown and Sorhannus 2010; Figure 5) show the Oomycota as the sister to the photosynthetic heterokonts. Basal taxa in the clade that includes the Oomycota are Pirsonia (Figure 6) and Developayella (Figure 7). A examination of these taxa, because they occur near the point at which a heterotrophic motile heterokont entered into secondary endosymbiosis to become photosynthetic, is great interest.

Pirsonia is similar to taxa in the Oomycota in that it is fungus-like in its life history and ecology. Figure 6 shows the margin of a heavily infected centric diatom with the liberation of propagules from the parasite. Curiously, the motile cells that they produce have two flagella, each of which has flagellar hairs. The anterior flagellum has the typical heterokont tripartite tubular hairs, but the posteriorly-directed flagellum has simple flagellar hairs. Although Kuhn et al. (2004) show Pirsonia as sister to the Oomycota, Yubuki et al. (2010) have Pirsonia emerging as sister to Oomycota + Ochrophyta (Photosynthetic Heterokonts). Developayella (Figure 7) is a free-living gliding flagellate with a ventral feeding groove. It does seem to be able to attach to the substrate by long strands of cytoplasm, which is reminiscent of the amoeboid silicoflagellates. Though the form of the cell would suggest that they should occupy a more basal position in the Oomycota clade, actual analyses have produced variable results. Andersen (2004) and Riisberg et al. (2009) have Developayella emerge as basal or sister to the Oomycota. However, Developayella appears within the Oomycota (Grant et al. 2009; Yubuki et al. 2010), as sister to the Ochrophytes (Kostka et al. 2004), and within the heterotrophic heterokonts, but not associated with the Oomycota (Kostka et al. 2007).

FIGURE 5. A cladogram of the Oomycota sensu latu (taxa in shaded box) within the heterotrophic taxa of the heterokonts (taxa in bold). This is a consensus of many analyses including Andersen (2004), Riisberg et al. (2009), and Brown and Sorhannus (2010).

FURTHER READING: DISCOVERY OF THE DOMAINS OF LIFE

INTRODUCTION TO THE DOMAIN EUKARYA

LITERATURE CITED Adl, S. M., A. G. B. Simpson, M. A. Farmer, R. A. Andersen, O. R. Anderson, J. R. Barta, S. S. Bowser, G. Brugerolle, R. A. Fensome, S. Fredericq, T. Y. James, S. Karpov, P. Kugrens, J. Krug, C. E. Lane, L. A. Lewis, J. Lodge, D. H. Lynn, D. G. Mann, R. M. McCourt, L. Mendoza, O. Moestrup, S. E. Mozley-Standridge, T. A. Nerad, C. A. Shearer, A. V. Smirnov, F. W. Spiegel, and M. F. J. R. Taylor. 2005. The new higher level classification of eukaryotes with emphasis on the taxonomy of protists. Journal of Eukaryotic Microbiology. 52(5): 399-451. Andersen R. A. 2004a. Biology and systematics of heterokont and haptophyte algae. American Journal of Botany. 91(10): 1508-1508. Barr, D. J. S. and N. L. Desaulniers. 1989. The flagellar apparatus of the Oomycetes and Hyphochytridiomycetes. In: Green, J. C., B. S. C. Leadbeater, and W. L. Diver, eds. The chromophyte algae; problems and perspectives. Systematics Association Special Volume No. 38. Clarendon Press. Oxford. pp. 343-355. Beakes, G. W. 1989. Oomycete Fungi: their phylogeny and relationship to chromophyte algae. In: Green, J. C., B. S. C. Leadbeater, and W. L. Diver, eds. The chromophyte algae: problems and perspectives. Systematics Association Special Volume No. 38. Clarendon Press. Oxford . pp. 325-342. Bold, H. C., C. J. Alexopoulos, and T. Delevoryas. 1987. Morphology of Plants and Fungi. 5th edition. HarperCollins Publishers. New York. Brown, J. W. and U. Sorhannus. 2010. A molecular genetic timescale for the diversification of autotrophic stramenopiles (Ochrophyta): substantive underestimation of putative fossil ages. PLoS ONE 5(9): e12759. doi:10.1371/journal.pone.0012759. Dick, M. W. 1990. Oomycota. In: Margulis, L., J.O. Corliss, M. Melkonian, and D.J. Chapman, eds. 1990. Handbook of the Protoctista; the Structure, Cultivation, Habits and Life Histories of the Eukaryotic Microorganisms and Their Descendants Exclusive of Animals, Plants and Fungi. Jones and Bartlett Publishers. Boston . pp. 661-685. Fuller, M.S. 1990. Hyphochytridiomycota. In: Margulis, L., J. O. Corliss, M. Melkonian, and D. J. Chapman, eds. 1990. Handbook of the Protoctista; the structure, cultivation, habits and life histories of the eukaryotic microorganisms and their descendants exclusive of animals, plants and fungi. Jones and Bartlett Publishers. Boston. pp. 380-387. Grant, J., Y. I. Tekle, O. R. Anderson, D. J. Patterson, and L. A. Katz. 2009. Multigene evidence for the placement of a heterotrophic amoeboid lineage Leukarachnion sp. among photosynthetic stramenopiles. Protist. 160. 376-385. [C]

Kostka, M., V. Hampl, I. Cepicka, and J. Flegr. 2004. Phylogenetic position of Protoopalina intestinalis based on SSU rRNA gene sequence. Molecular Phylogenetics and Evolution. 33: 220-224. Kostka, M., I. Cepocka, V. Hampl, and J. Flegr. 2007. Phylogenetic position of Karotomorpha and paraphyly of Proteromonadidae. Khn, S., L. Medlin, and G. Eller. 2004. Phylogenetic position of the parasitoid nanoflagellate Pirsonia inferred from Nuclear-encoded small subunit ribosomal DNA and a description of Pseudopirsonia n. gen. and Pseudopirsonia mucosa (Drebes) comb. nov. Protist. 155: 143-156. Margulis, L. and K. Schwartz. 1988. Five kingdoms, an illustrated guide to the phyla of life on earth. 2nd Edition. W.H. Freeman and Co. New York. Margulis, L. and K. Schwartz. 1998. Five kingdoms, an illustrated guide to the phyla of life on earth. 3rd Edition. W. H. Freeman and Company. New York. Patterson, D. J. 1999. The diversity of eukaryotes. American Naturalist. 154 (Suppl.): S96S124. Riisberg, I., R. J. S. Orr, R. Kluge, K. Shalchian-Tabrizi, H. A. Bowers, V. Patil, B. Edvardsen, and K. S. Jakobsen. 2009. Seven gene phylogeny of heterokonts. Protist. 160: 191-204. Sogin, M. L. and D. J. Patterson. 1995. Stramenopiles. Version 01 January 1995 (under construction). http://tolweb.org/Stramenopiles/2380/1995.01.01 In: The Tree of Life Web Project, http://tolweb.org/ Tong, S. M. 1995. Developayella elegans nov. gen., nov. spec., a new type of hetertrophic flagellate from marine plankton. European Journal of Protistology. 31: 24-31. Winter, G. 1879. ber ein natrliches System der Thallophyten. Hedwig a 1879. pt.1: 1 12. Yang, E. C., G. H. Boo, H. J. Kim, S. M. Cho, S. M. Boo, R. A. Andersen, and H. S. Yoon. 2012. Supermatrix data highlight the phylogenetic relationships of photosynthetic stramenopiles. Protist. 163: 217-231. Yubuki, N., B. S. Leander, and J. D. Silberman. 2010. Untrastructural and molecular phylogenetic position of a novel phagotrophic stramenopile from low oxygen environments: Rictus lutensis gen. et sp. nov. (Bicosoecida, incertae sedis). Protist. 161: 264-278.

Division: Oomycota The division Oomycota, unlike the Chytridiomycota, is a group of aquatic fungi that have cell walls

composed of cellulose and a diploid dominant lifecycle (=gametic lifecycle). The mycelium is coenocytic and produce septa only to separate the reproductive structures from the assimilative portion of the thallus. Asexual reproduction is by zoospores that are produced in zoosporangia. The zoospores produced are biflagellated with one flagellum of the whiplash type and the other of the tinsel type. Sexual reproduction is heterogamous and occurs by direct injection of the male nuclei (=sperms) from the antheridium into the eggs contained in the oogonium . A swimming sperm is absent in the Oomycota. This type of sexual reproduction is referred to as gametangial copulation. The eggs and sperms are products of meiosis and the only parts of the life cycle that are haploid. Only one class, the Oomycetes, is recognized in this division. The combination of cellulose cell wall, biflatellated zoospores, one flagellum of the tinsel type and the other of the whiplash type, and gametangial copulation are characteristics that are shared with some members of the algal divisions Phaeophyta and Chrysophyta. This has recently led to recognition of yet another kingdom, Stramenopila, which includes the divisions Bacillariophyta, Chrysophyta, Phaeophyta, Hyphochytridiomycota and Oomycota. These divisions are now thought to be derived from a common ancestor . This phylum of organisms differs in fundamental ways from the rest of the organisms in the Kingdom Fungi cell wall chemistry, lysine biosynthesis, rDNA sequences Basic form of thallus in most is the mycelium with hyphae, some form holocarpic thalli Significant economic group contains species that are important plant pathogens Evolutionary trends in Oomycota Vegetative thallus in some species, it is holocarpic, endobiotic most for well developed aseptate hyphae (eucarpic, polycentric) Nutrition some species are aquatic (both saprotrophic and parasitic). Others are highly specialized biotrophic parasites of higher plants Asexual reproduction In many, zoospores are produced in a zoosporangium, in some the number of zoospores/sporangium is reduced. In a few species, the sporangium functions as a conidium (germinates with a germ tube) Oomycota Contains one class Oomycetes Two subclasses: Saprolegniomycetidae one order Saprolegniales Peronosporomycetidae five orders, we will discuss representatives of two Pythiales Peronosporales Class: Oomycetes The characteristics of the class Oomycetes is the same as those of the division. We will examine two representatives in the Oomycota: Saprolegnia sp. (Saprolegniales) and Phytophthora palmivora (Peronosporales). Saprolegniales Vegetative thallus varies from relatively simple holocarpic thallus (parasitic on algae and fungi) to well developed mycelium Hyphae are aseptate Commonly called water molds Most are saprotrophs, some parasitize fish and fish eggs can cause large economic losses to fish

hatcheries Asexual reproduction Zoospores produced by long cylindrical zoosporangia typically formed at the hyphal tips (formed when immersed in water) Zoospores Two types of zoospores produced sequentially = dimorphic If they produce only one type of zoospore = monomorphic Primary zoospores are pear shaped, poor swimmers Secondary zoospores are oval to kidney shaped and better swimmers Primary & secondary zoospores Species vary in sequence of events in formation of primary and secondary zoospores (see handout) Zoospores Saprolegnia primary zoospore swims away before encysting, then forms secondary zoospore Achlya 1? zoospore encysts right outside zoosporangium, then forms 2? zoospore Dichtyuchus No 1? flagellated zoospores, they encyst in the zoosporangium, cysts germinate to form 2? zoospores or they may germinate with a germ tube Asexual reproduction May also produce gemmae irregularly shaped hyphal segments that separate from the thallus and can germinate (germ tube or zoospores) to form a new thallus Sexual life cycle For a long time the Oomycetes were thought to have a haploid vegetative phase like most other fungi Since nuclei are small, it is difficult to determine where meiosis takes place Microspectrophotometry allowed concentration of DNA in nuclei to be measured in situ indicated that vegetative mycelium is diploid, meiosis takes place in gametangia Exhibit a diploid life cycle (unusual for fungi Life cycle Vegetative hyphae are diploid Undergo asexual reproduction by formation of zoospores in zoosporangia Gametangia formed & meiosis occurs Homothallic & heterothallic species Male gametangium antheridium Female gametangium oogonium that contains multiple gametes oospheres in the Saprolegniales Antheridium grows to oogonium Forms fertilization tubes to oospheres Nuclei from antheridia migrate through fertilization tubes Plasmogamy and karyogamy take place Oosphere is now diploid Develops thick wall and becomes an oospore Oogonium Oospore can remain dormant Germinates to produce zoosporangium or 2n mycelium Saprolegnia: Asexual Reproduction: Zoosporangia are not differentiated, in this genus, from the mycelium and

resemble hyphal cells. However, they can be easily located since they are delimited by septa and have much denser protoplasm than the rest of the mycelium. Saprolegnia zoosporangia can be identified by the internal proliferation of a new zoosporangium inside of the previous one (Fig. 1). Compare this genus with Achyla, which as lateral proliferation of zoosporangia, next to and below the previous zoosporangium (Fig. 2).Two types of zoospores are produced: The primary zoospore, which is posteriorly biflagellated, is released from the zoosporangium, encyst and germinates to form the secondary zoospore which is reniform (=bean-shaped) and laterally flagellated. Figure 1: Zoosporangium of Saprolengia: Note proliferation of zoosporangium through previous zoosporangium. Youngest zoosporangium has dense protoplasm with basal septum.

Figure 2: Zoosporangium of Achyla: Note lateral proliferation of zoosporangium below previous zoosporangium. Primary zoospores do not have a swarming period. Instead, they encyst immediately upon being released from the zoosporangium. Sexual Reproduction: The oogonium (Fig. 3) contains from 2 to 20 eggs and typically have several antheridia attached to the oogonial wall at maturity. Sperm nuclei from the antheridium are injected directly to the eggs and form the zygote. The egg and sperm are the products of meiosis and are the only haploid stage in the life cycle.

Figure 3: Oogonia and Eggs of Achyla, with attached Antheridia. Note that there are more than a single egg in each oogonium, a characteristic of the Saprolegniales.

Peronosporomycetidae Sporangia may produce zoospores but only secondary zoospores or, Trend in this subclass is for the zoosporangia to produce smaller number of zoospores In some species, sporangia may germinate with a germ tube (called conidia) Pythiales Pythium a large genus that includes saprotrophs (soil and water) and facultative parasites of algae, fungi and plants One species causes damping off of seedlings grows intercellularly in stems and roots of seedlings and rots tissue

 Asexual reproduction by zoosporangia and zoospores (secondary zoospores only) Contents of zoosporangium empty into a vesicle, cleave and form zoospores outside zoosporangium Pythiales Phytophthora facultative plant parasites In plant host, form intercellular between cells - and intracellular penetrate cells - hyphae Intracellular hyphae produce haustoria exchange of nutrients from host to parasite Phytophthora Forms lemon shaped sporangia that detach from sporangiophore in asexual reproduction sporangia can germinate by forming zoospores or forming a germ tube directly Phytophthora palmivora: Asexual Reproduction: The Zoosporangium (Fig. 4-6) is typically broad-ellipsoid and well differentiated from the vegetative myucelium in this species. Because the zoosporangia are readily deciduous, they will usually not be attached to the mycelium when mounted on a microscope slide for observation. Only the secondary zoospore is produced in this species. The micrographs below were taken under phase interference optics, which is responsible for the dark appearance of the images. Figure 4: Intact zoosporangium of Phytophthora palmivora before release of zoospore. Note that it is morphologically differentiated from the hyphae, unlike those produced by Achyla and Saprolegnia.

Figure 5: Zoosporangium of Phytophthora palmivora with zoospores clustered together and oozing out of operculum.

Figure 6: Zoospores of Phytopthora palmivora coming apart. Two zoospores swiming away can be seen at the top of the screen. Sexual Reproduction: The oogonium in this species, unlike Saprolegnia, contains only one egg. Peronosporales Albugo obligate plant parasite, causes white rust Sporangia detach, dispersed by wind If moisture available, sporangia produce 4-12 zoospores Can germinate directly with a germ tube Peronosporales Peronospora and related genera obligate parasites of plants cause downy mildews In some species, sporangia always germinate with a germ tube, never form zoospores called conidia