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''Galdieria sulphuraria'' is an
extremophilic An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...
unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
. It is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' Galdieria''. It is known for its broad metabolic capacities, including
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
and heterotrophic growth on over 50 different extracellular carbon sources. The members of the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
Cyanidiophyceae Cyanidiophyceae is a class of unicellular red algae within subdivision Cyanidiophytina, and contain a single plastid, one to three mitochondria, a nucleus, a vacuole and floridean starch. Most are extremophiles inhabiting acid hot springs. The ma ...
are among the most
acidophilic Acidophiles or acidophilic organisms are those that thrive under highly acidic conditions (usually at pH 5.0 or below). These organisms can be found in different branches of the tree of life, including Archaea, Bacteria,Becker, A.Types of Bacteria ...
known photosynthetic organisms, and the growth conditions of ''G. sulphuraria'' – pH between 0 and 4, and
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied o ...
s up to 56 °C – are among the most extreme known for
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. Analysis of its
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
suggests that its
thermoacidophilic A thermoacidophile is an extremophilic microorganism that is both thermophilic and acidophilic; i.e., it can grow under conditions of high temperature and low pH. The large majority of thermoacidophiles are archaea (particularly the Thermoproteot ...
adaptations derive from horizontal gene transfer from
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
and
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, another rarity among eukaryotes.


History and taxonomy

Published descriptions of thermoacidophilic unicellular algae date to the mid-19th century. The earliest description of an organism corresponding to the modern ''G. sulphuraria'' was published in 1899 by an Italian scientist, A. Galdieri, who gave it the name ''Pleurococcus sulphurarius''. The
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of thermoacidophilic algae was revised in 1981, which introduced the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' Galdieria'' and gave the organism its modern designation. ''G. sulphuraria'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
for this genus. The group to which ''G. sulphuraria'' belongs, the
Cyanidiophyceae Cyanidiophyceae is a class of unicellular red algae within subdivision Cyanidiophytina, and contain a single plastid, one to three mitochondria, a nucleus, a vacuole and floridean starch. Most are extremophiles inhabiting acid hot springs. The ma ...
, is the most deeply branching subgroup of the
rhodophyta Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
(red algae), meaning they were the earliest to diverge in the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary history of this group.


Metabolism

''G. sulphuraria'' is noted for its extreme
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
flexibility: it is capable of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
and can also grow heterotrophically on a wide variety of carbon sources, including diverse
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s. Over 50 different carbon sources that support growth have been reported. Careful measurements of its growth patterns under laboratory conditions suggest that it is not a true mixotroph capable of using both energy sources at the same time; rather, it prefers heterotrophic growth conditions and downregulates photosynthesis after extended exposure to extracellular carbon sources. Analysis of the ''G. sulphuraria'' photosystem I complex, a key photosynthetic component, suggests a structure intermediate between the homologous complexes in
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
and
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
s. Although most red algae use
floridean starch Floridean starch is a type of a storage glucan found in glaucophytes and in red algae (or rhodophytes), in which it is usually the primary sink for fixed carbon from photosynthesis. It is found in grains or granules in the cell's cytoplasm and is c ...
as a storage
glucan A glucan is a polysaccharide derived from D-glucose, linked by glycosidic bonds. Glucans are noted in two forms: alpha glucans and beta glucans. Many beta-glucans are medically important. They represent a drug target for antifungal medications of ...
, ''G. sulphuraria'' uses a highly unusual form of
glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one o ...
which is among the most highly branched glycogens known, has very short branch lengths, and forms particles of unusually low
molecular weight A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
. These properties are believed to be metabolic adaptations to extreme environmental conditions, although the precise mechanism is unclear.


Habitat and ecology

''G. sulphuraria'' is unusual for a
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
in being
thermoacidophilic A thermoacidophile is an extremophilic microorganism that is both thermophilic and acidophilic; i.e., it can grow under conditions of high temperature and low pH. The large majority of thermoacidophiles are archaea (particularly the Thermoproteot ...
– that is, capable of growing at both high temperature and low pH. It grows well in a pH range of 0–4 and at temperatures up to 56 °C, close to the approximately 60 °C sometimes cited as the likely maximum for eukaryotic life. It is also highly tolerant of high
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
concentrations and of toxic metals. It is found in naturally acidic
hot spring A hot spring, hydrothermal spring, or geothermal spring is a spring produced by the emergence of geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow bodies of magma (molten rock) or by circ ...
s, in solfataric environments, and in polluted environments; It is also found in
endolithic ecosystem An endolith or endolithic is an organism (archaeon, bacterium, fungus, lichen, algae or amoeba) that is able to acquire the necessary resources for growth in the inner part of a rock, mineral, coral, animal shells, or in the pores between min ...
s, where light is scarce and its heterotrophic metabolic capacities are particularly important. Laboratory tests indicate that it is capable of actively acidifying its environment.


Genome

The ''G. sulphuraria''
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
contains evidence of extensive horizontal gene transfer (HGT) from thermoacidophilic
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
and
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, explaining the origin of its
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
to this environment. At least 5% of its proteome is likely to be derived from HGT. This is highly unusual for a eukaryote; relatively few well-substantiated examples exist of HGT from prokaryotes to eukaryotes. The genome of its
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
is also exceptionally small and has a very high
GC skew GC skew is when the nucleotides guanine and cytosine are over- or under-abundant in a particular region of DNA or RNA. GC skew is also a statistical method for measuring strand-specific guanine overrepresentation. In equilibrium conditions (wi ...
, while the genome of its
plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosy ...
s is of normal size but contains an unusual number of
stem-loop Stem-loop intramolecular base pairing is a pattern that can occur in single-stranded RNA. The structure is also known as a hairpin or hairpin loop. It occurs when two regions of the same strand, usually complementary in nucleotide sequence when ...
structures. Both of these properties are proposed to be adaptations for the organism's polyextremophilic environment. By comparison to '' Cyanidioschyzon merolae'' – a unicellular thermoacidophilic red alga that is obligately photoautotrophic – the ''G. sulphuraria'' genome contains a large number of genes associated with carbohydrate metabolism and cross-membrane transport.


Biotechnology

Because of its ability to tolerate extreme environments and grow under a wide variety of conditions, ''G. sulphuraria'' has been considered for use in bioremediation projects. For example, it has been tested for the ability to recover
precious metal Precious metals are rare, naturally occurring metallic chemical elements of high economic value. Chemically, the precious metals tend to be less reactive than most elements (see noble metal). They are usually ductile and have a high lustre. ...
s, recover
rare-earth metal The rare-earth elements (REE), also called the rare-earth metals or (in context) rare-earth oxides or sometimes the lanthanides (yttrium and scandium are usually included as rare earths), are a set of 17 nearly-indistinguishable lustrous silve ...
s, and remove
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
and
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
from various waste streams.


References


External links

* * https://microbewiki.kenyon.edu/index.php/Galdieria_sulphuraria {{Authority control Thermophiles Acidophiles Species described in 1982 Cyanidiophyceae