A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comprise more than half of all microscopic plankton. [1] There are two types of eukaryotic mixotrophs: those with their own chloroplasts, and those with endosymbionts—and those that acquire them through kleptoplasty or through symbiotic associations with prey or enslavement of their organelles. [2]
Possible combinations are photo- and chemotrophy, litho- and organotrophy (osmotrophy, phagotrophy and myzocytosis), auto- and heterotrophy or other combinations of these. Mixotrophs can be either eukaryotic or prokaryotic. [3] They can take advantage of different environmental conditions. [4]
If a trophic mode is obligate, then it is always necessary for sustaining growth and maintenance; if facultative, it can be used as a supplemental source. [3] Some organisms have incomplete Calvin cycles, so they are incapable of fixing carbon dioxide and must use organic carbon sources.
Organisms may employ mixotrophy obligately or facultatively.
Amongst plants, mixotrophy classically applies to carnivorous, hemi-parasitic and myco-heterotrophic species. However, this characterisation as mixotrophic could be extended to a higher number of clades as research demonstrates that organic forms of nitrogen and phosphorus — such as DNA, proteins, amino-acids or carbohydrates — are also part of the nutrient supplies of a number of plant species. [6]
Mixotrophy is less common among animals than among plants and microbes, but there are many examples of mixotrophic invertebrates and at least one example of a mixotrophic vertebrate.
To characterize the sub-domains within mixotrophy, several very similar categorization schemes have been suggested. Consider the example of a marine protist with heterotrophic and photosynthetic capabilities: In the breakdown put forward by Jones, [19] there are four mixotrophic groups based on relative roles of phagotrophy and phototrophy.
An alternative scheme by Stoeker [18] also takes into account the role of nutrients and growth factors, and includes mixotrophs that have a photosynthetic symbiont or who retain chloroplasts from their prey. This scheme characterizes mixotrophs by their efficiency.
Another scheme, proposed by Mitra et al., specifically classifies marine planktonic mixotrophs so that mixotrophy can be included in ecosystem modeling. [20] This scheme classified organisms as:
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The Acantharea (Acantharia) are a group of radiolarian protozoa, distinguished mainly by their strontium sulfate skeletons. Acantharians are heterotrophic marine microplankton that range in size from about 200 microns in diameter up to several millimeters. Some acantharians have photosynthetic endosymbionts and hence are considered mixotrophs.
Plankton are the diverse collection of organisms found in water that are unable to propel themselves against a current. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish, and baleen whales.
Zooplankton are the animal component of the planktonic community, having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequently, they drift or are carried along by currents in the ocean, or by currents in seas, lakes or rivers.
Primary nutritional groups are groups of organisms, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the sources of carbon can be of organic or inorganic origin.
A heterotroph is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but not producers. Living organisms that are heterotrophic include all animals and fungi, some bacteria and protists, and many parasitic plants. The term heterotroph arose in microbiology in 1946 as part of a classification of microorganisms based on their type of nutrition. The term is now used in many fields, such as ecology in describing the food chain.
The green sulfur bacteria are a phylum, Chlorobiota, of obligately anaerobic photoautotrophic bacteria that metabolize sulfur.
Photoheterotrophs are heterotrophic phototrophs—that is, they are organisms that use light for energy, but cannot use carbon dioxide as their sole carbon source. Consequently, they use organic compounds from the environment to satisfy their carbon requirements; these compounds include carbohydrates, fatty acids, and alcohols. Examples of photoheterotrophic organisms include purple non-sulfur bacteria, green non-sulfur bacteria, and heliobacteria. These microorganisms are ubiquitous in aquatic habitats, occupy unique niche-spaces, and contribute to global biogeochemical cycling. Recent research has also indicated that the oriental hornet and some aphids may be able to use light to supplement their energy supply.
Osmotrophy is a feeding mechanism involving the movement of dissolved organic compounds by osmosis for nutrition. Organisms that use osmotrophy are called osmotrophs. Some mixotrophic microorganisms use osmotrophy to derive some of their energy. Osmotrophy is used by many diverse organisms. Organisms that use osmotrophy include bacteria, many species of protists and most fungi. Some macroscopic animals like molluscs, sponges, corals, brachiopods and echinoderms may use osmotrophic feeding as a supplemental food source.
Hatena arenicola is a species of single-celled eukaryotes discovered in 2000, and first reported in 2005. It was discovered by Japanese biologists Noriko Okamoto and Isao Inouye at the University of Tsukuba, and they gave the scientific description and formal name in 2006. The species is a flagellate, and can resemble a plant at one stage of its life, in which it carries a photosynthesizing alga inside itself, or an animal, acting as predator in another stage of its life. Researchers believe that this organism is in the process of secondary endosymbiosis, in which one organism is incorporated into another, resulting in a completely new life form.
Myco-heterotrophy is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food from parasitism upon fungi rather than from photosynthesis. A myco-heterotroph is the parasitic plant partner in this relationship. Myco-heterotrophy is considered a kind of cheating relationship and myco-heterotrophs are sometimes informally referred to as "mycorrhizal cheaters". This relationship is sometimes referred to as mycotrophy, though this term is also used for plants that engage in mutualistic mycorrhizal relationships.
Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.
Predatory dinoflagellates are predatory heterotrophic or mixotrophic alveolates that derive some or most of their nutrients from digesting other organisms. About one half of dinoflagellates lack photosynthetic pigments and specialize in consuming other eukaryotic cells, and even photosynthetic forms are often predatory.
A protist or protoctist is any eukaryotic organism that is not an animal, plant, or fungus. Protists do not form a natural group, or clade, but an artificial grouping of several independent clades that evolved from the last eukaryotic common ancestor.
Marine microorganisms are defined by their habitat as microorganisms living in a marine environment, that is, in the saltwater of a sea or ocean or the brackish water of a coastal estuary. A microorganism is any microscopic living organism or virus, that is too small to see with the unaided human eye without magnification. Microorganisms are very diverse. They can be single-celled or multicellular and include bacteria, archaea, viruses and most protozoa, as well as some fungi, algae, and animals, such as rotifers and copepods. Many macroscopic animals and plants have microscopic juvenile stages. Some microbiologists also classify viruses as microorganisms, but others consider these as non-living.
Vitrella brassicaformis (CCMP3155) is a unicellular alga belonging to the eukaryotic supergroup Alveolata. V. brassicaformis and its closest known relative, Chromera velia, are the only two currently described members of the phylum Chromerida, which in turn constitutes part of the taxonomically unranked group Colpodellida. Chromerida is phylogenetically closely related to the phylum Apicomplexa, which includes Plasmodium, the agent of malaria. Notably, both V. brassicaformis and C. velia are photosynthetic, each containing a complex secondary plastid. This characteristic defined the discovery of these so-called 'chromerids,' as their photosynthetic capacity positioned them to shed light upon the evolution of Apicomplexa's non-photosynthetic parasitism. Both genera lack chlorophyll b or c; these absences link the two taxonomically, as algae bearing only chlorophyll a are rare amid the biodiversity of life. Despite their similarities, V. brassicaformis differs significantly from C. velia in morphology, lifecycle, and accessory photosynthetic pigmentation. V. brassicaformis has a green color, with a complex lifecycle involving multiple pathways and a range of sizes and morphologies, while Chromera has a brown color and cycles through a simpler process from generation to generation. The color differences are due to differences in accessory pigments.
Picozoa, Picobiliphyta, Picobiliphytes, or Biliphytes are protists of a phylum of marine unicellular heterotrophic eukaryotes with a size of less than about 3 micrometers. They were formerly treated as eukaryotic algae and the smallest member of photosynthetic picoplankton before it was discovered they do not perform photosynthesis. The first species identified therein is Picomonas judraskeda. They probably belong in the Archaeplastida as sister of the Rhodophyta.
Dinophysis acuminata is a marine plankton species of dinoflagellates that is found in coastal waters of the north Atlantic and Pacific oceans. The genus Dinophysis includes both phototrophic and heterotrophic species. D. acuminata is one of several phototrophic species of Dinophysis classed as toxic, as they produce okadaic acid which can cause diarrhetic shellfish poisoning (DSP). Okadiac acid is taken up by shellfish and has been found in the soft tissue of mussels and the liver of flounder species. When contaminated animals are consumed, they cause severe diarrhoea. D. acuminata blooms are constant threat to and indication of diarrhoeatic shellfish poisoning outbreaks.
Dinoflagellates are eukaryotic plankton, existing in marine and freshwater environments. Previously, dinoflagellates had been grouped into two categories, phagotrophs and phototrophs. Mixotrophs, however include a combination of phagotrophy and phototrophy. Mixotrophic dinoflagellates are a sub-type of planktonic dinoflagellates and are part of the phylum Dinoflagellata. They are flagellated eukaryotes that combine photoautotrophy when light is available, and heterotrophy via phagocytosis. Dinoflagellates are one of the most diverse and numerous species of phytoplankton, second to diatoms.
Rhodelphis is a single-celled archaeplastid that lives in aquatic environments and is the sister group to red algae and possibly Picozoa. While red algae have no flagellated stages and are generally photoautotrophic, Rhodelphis is a flagellated predator containing a non-photosynthetic plastid. This group is important to the understanding of plastid evolution because they provide insight into the morphology and biochemistry of early archaeplastids. Rhodelphis contains a remnant plastid that is not capable of photosynthesis, but may play a role in biochemical pathways in the cell like heme synthesis and iron-sulfur clustering. The plastid does not have a genome, but genes are targeted to it from the nucleus. Rhodelphis is ovoid with a tapered anterior end bearing two perpendicularly-oriented flagella.
Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.