ultrastructuralThe other definition describes protists primarily by functional or biological criteria: protists are essentially those eukaryotes that are never multicellular,[29] that either exist as independent cells, or if they occur in colonies, do not show differentiation into tissues (but vegetative cell differentiation may occur restricted to sexual reproduction, alternate vegetative morphology, and quiescent or resistant stages, such as cysts);[33] this definition excludes many brown, multicellular red and green algae, which may have tissues.
The taxonomy of protists is still changing. Newer classifications attempt to present monophyletic groups based on morphological (especially ultrastructural),[34][35][36] biochemical (chemotaxonomy)[37][38] and DNA sequence (molecular research) information.[39][40] However, there are sometimes discordances between molecular and morphological investigations; these can be categorized as two types: (i) one morphology, multiple lineages (e.g. morphological convergence, cryptic species) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity, multiple life-cycle stages).[41]
Because the protists as a whole are paraphyletic, new systems often split up or abandon the kingdom, instead treating the protist groups as separate lines of eukaryotes. The recent scheme by Adl et al. (2005)[33] does not recognize formal ranks (phylum, class, etc.) and instead treats groups as clades of phylogenetically related organisms. This is intended to make the classification more stable in the long term and easier to update. Some of the main groups of protists, which may be treated as phyla, are listed in the taxobox, upper right.[42] Many are thought to be monophyletic, though there is still uncertainty. For instance, the Excavata are probably not monophyletic and the chromalveolates are probably only monophyletic if the haptophytes and cryptomonads are excluded.[43]
Nutrition can vary according to the type of protist. Most eukaryotic algae are autotrophic, but the pigments were lost in some groups.[vague] Other protists are heterotrophic, and may present phagotrophy, osmotrophy, saprotrophy or parasitism. Some are mixotrophic. Some protists that do not have / lost chloroplasts/mitochondria have entered into endosymbiontic relationship with other bacteria/algae to replace the missing functionality. For example, Paramecium bursaria and Paulinella have captured a green alga (Zoochlorella) and a cyanobacterium respectively that act as replacements for chloroplast. Meanwhile, a protist, Mixotricha paradoxa that has lost its mitochondria uses endosymbiontic bacteria as mitochondria and ectosymbiontic hair-like bacteria (Treponema spirochetes) for locomotion.
Many protists are flagellate, for example, and filter feeding can take place where flagellates find prey. Other protists can engulf bacteria and other food particles, by ext
Many protists are flagellate, for example, and filter feeding can take place where flagellates find prey. Other protists can engulf bacteria and other food particles, by extending their cell membrane around them to form a food vacuole and digesting them internally in a process termed phagocytosis.
For most important cellular structures and functions of animal and plants, it can be found a heritage among protists.[44]
Reproduction
Some protists reproduce sexually using gametes, while others reproduce asexually by binary fission.
Some species, for example Plasmodium falciparum, have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually.[45] However, it is unclear how frequently sexual reproduction causes genetic exchange between different strains of Plasmodium in nature and most
Some protists reproduce sexually using gametes, while others reproduce asexually by binary fission.
Some species, for example Plasmodium falciparum, have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually.Some species, for example Plasmodium falciparum, have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually.[45] However, it is unclear how frequently sexual reproduction causes genetic exchange between different strains of Plasmodium in nature and most populations of parasitic protists may be clonal lines that rarely exchange genes with other members of their species.[46]
Eukaryotes emerged in evolution more than 1.5 billion years ago.[47] The earliest eukaryotes were likely protists. Although sexual reproduction is widespread among extant eukaryotes, it seemed unlikely until recently, that sex could be a primordial and fundamental characteristic of eukaryotes. A principal reason for this view was that sex appeared to be lacking in certain pathogenic protists whose ancestors branched off early from the eukaryotic family tree. However, several of these protists are now known to be capable of, or to recently have had the capability for, meiosis and hence sexual reproduction. For example, the common intestinal parasite Giardia lamblia was once considered to be a descendant of a protist lineage that predated the emergence of meiosis and sex. However, G. lamblia was recently found to have a core set of genes that function in meiosis and that are widely present among sexual eukaryotes.[48] These results suggested that G. lamblia is capable of meiosis and thus sexual reproduction. Furthermore, direct evidence for meiotic recombination, indicative of sex, was also found in G. lamblia.[49]
The pathogenic parasitic protists of the genus Leishmania have been shown to be capable of a sexual cycle in the invertebrate vector, likened to the meiosis undertaken in the trypanosomes.[50]
Trichomonas vaginalis, a parasitic protist, is not known to undergo meiosis, but when Malik et al.[51] tested for 29 genes that function in meiosis, they found 27 to be present, including 8 of 9 genes specific to meiosis in model eukaryotes. These findings suggest that T. vaginalis may be capable of meiosis. Since 21 of the 29 meiotic genes were also present in G. lamblia, it appears that most of these meiotic genes were likely present in a common ancestor of T. vaginalis and G. lamblia. These two species are descendants of protist lineages that are highly divergent among eukaryotes, leading Malik et al.[51] to suggest that these meiotic genes were likely present in a common ancestor of all eukaryotes.
Based on a phylogenetic analysis, Dacks and Roger proposed that facultative sex was present in the common ancestor of all eukaryotes.[52]
This view was further supported by a study of amoebae by Lahr et al.[53] Amoeba have generally been regarded as asexual protists. However, these authors describe evidence that most amoeboid lineages are anciently sexual, and that the majority of asexual groups likely arose recently and independently. Early researchers (e.g., Calkins) have interpreted phenomena related to chromidia (chromatin granules free in the cytoplasm) in amoeboid organisms as sexual reproduction.[54]
Protists generally reproduce asexually under favorable environmental conditions, but tend to reproduce sexually under stressful conditions, such as starvation or heat shock.[55] Oxidative stress, which is associated with the production of reactive oxygen species leading to DNA damage, also appears to be an important factor in the induction of sex in protists.[55]
Some commonly found Protist pathogens such as Toxoplasma gondii are capable of infecting and undergoing asexual reproduction in a wide variety of animals – which act as secondary or intermediate host – but can undergo sexual reproduction only in the primary or definitive host (for example: felids such as domestic cats in this case).[56][57][58]
Free-living Protists occupy almost any environment that contains liquid water. Many protists, such as algae, are photosynthetic and are vital primary producers in ecosystems, particularly in the ocean as part of the plankton. Protists make up a large portion of the biomass in both marine and terrestrial environments.[59]
Other protists include pathogenic species, such as the kinetoplastid Trypanosoma brucei, which causes sleeping sickness, and species of the apicomplexan Plasmodium, which cause malaria.
Parasitism: role as pathogens
See also:
kinetoplastid Trypanosoma brucei, which causes
sleeping sickness, and species of the
apicomplexan Plasmodium, which cause
malaria.
Some protists are significant parasites of animals (e.g.; five species of the parasitic genus Plasmodium cause malaria in humans and many others cause similar diseases in other vertebrates), plants[60][61] (the oomycete Phytophthora infestans causes late blight in potatoes)[62] or even of other protists.[63][64] Protist pathogens share many metabolic pathways with their eukaryotic hosts. This makes therapeutic target development extremely difficult – a drug that harms a protist parasite is also likely to harm its animal/plant host. A more thorough understanding of protist biology may allow these diseases to be treated more efficiently. For example, the apicoplast (a nonphotosynthetic chloroplast but essential to carry out important functions other than photosynthesis) present in apicomplexans provides an attractive target for treating diseases caused by dangerous pathogens such as plasmodium.
Recent papers have proposed the use of viruses to treat infections caused by protozoa.[65][66]
Researchers from the Agricultural Research Service are taking advantage of protists as pathogens to control red imported fire ant (Solenopsis invicta) populations in protozoa.[65][66]
Researchers from the Agricultural Research Service are taking advantage of protists as pathogens to control red imported fire ant (Solenopsis invicta) populations in Argentina. Spore-producing protists such as Kneallhazia solenopsae (recognized as a sister clade or the closest relative to the fungus kingdom now)[67] can reduce red fire ant populations by 53–100%.[68] Researchers have also been able to infect phorid fly parasitoids of the ant with the protist without harming the flies. This turns the flies into a vector that can spread the pathogenic protist between red fire ant colonies.[69]
Many protists have neither hard parts nor resistant spores, and their fossils are extremely rare or unknown. Examples of such groups include the apicomplexans,[70] most ciliates,[71] some green algae (the Klebsormidiales),[72] choanoflagellates,[73] oomycetes,[74] brown algae,[75] yellow-green algae,[76] Excavata (e.g., euglenids).[77] Some of these have been found preserved in amber (fossilized tree resin) or under unusual conditions (e.g., Paleoleishmania, a kinetoplastid).
Others are relatively common in the fossil record,[78] as the diatoms,[79] Others are relatively common in the fossil record,[78] as the diatoms,[79] golden algae,[80] haptophytes (coccoliths),[81] silicoflagellates, tintinnids (ciliates), dinoflagellates,[82] green algae,[83] red algae,[84] heliozoans, radiolarians,[85] foraminiferans,[86] ebriids and testate amoebae (euglyphids, arcellaceans).[87] Some are even used as paleoecological indicators to reconstruct ancient environments.
More probable eukaryote fossils begin to appear at about 1.8 billion years ago, the acritarchs, spherical fossils of likely algal protists.[88] Another possible representative of early fossil eukaryotes are the Gabonionta.