Ectocarpus Siliculosus
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''Ectocarpus siliculosus'' is a filamentous
brown alga Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate an ...
. Its genome was the first brown macroalgal genome to be sequenced, with the expectation that ''E. siliculosus'' will serve as a genetic and genomic model for brown macroalgae.


Ecology

The brown algae are members of the
stramenopiles Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
(along with organisms such as
diatoms A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
and
oomycetes Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the result ...
). The stramenopiles diverged from other major eukaryotic groups such as the
opisthokonts The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
(animals and fungi) and the
archaeplastida The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor gr ...
(which includes land plants) over a billion years ago. The brown algae are also important because they are one of only a small number of eukaryotic groups that have evolved complex multicellularity. The alga is unbranched and filamentous; it forms soft beards on larger plants or other firm substrata and grows up to 2 feet long. Its
thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms ...
is filamentous, initially organized as a main primary filament composed of elongated cells and round cells, from which branches differentiate. E. siliculosus is a tufted plant, often only one to a few cm tall, but in exceptional cases up to 20 cm. It has axes that are freely branched, and the main axis is not distinguishable. Filaments on E. siliculosus can grow up to 30μm in diameter, tapering toward the apices and sometimes forming terminal pseudo hairs.


Reproduction

E. siliculosus reproduction and growth involves two different patterns of early development, which begin with either a symmetric or an asymmetric division of the initial cell. Symmetric division leads to the development of a prostrate, basal structure before the erect thallus is formed. Asymmetric division leads to the immediate development of an erect thallus without the formation of a prostrate, basal structure (immediate differentiation). E. siliculosus alternates between two generational life cycles that differ in either being
sporophytes A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygot ...
( produce few laterals and develop from a branched prostrate base) or
gametophytes A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
( richly branched and devoid of a prostrate base). E. siliculosus gametophytes have an asymmetric initial cell division and immediate differentiation of an erect thallus. The alternation of the two different generations in E. siliculosus therefore alternates between symmetric and asymmetric cell divisions as well. E. siliculosus develops uniseriate filaments. It has a sporophyte body which is made up of the prostrate body and the upright body. The prostrate body is in turn composed of crawling filaments ( crawling filaments are made of Elongated (E) cells and Round (R) cells) which is a filament with E cells on the edges and R cells in the center. Then, there is a period of secondary growth in which axes develop  in the center of the primary filament and on the R cells. The upright filaments will grow from the prostrate body and differentiate into
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
.


E. siliculosus in Research

Brown algae have many unique characteristics in terms of their metabolism and cell biology. Ergo, brown algae and in particular, E. siliculosus, are often used for explorative research. Its genome was the first brown macroalgal genome to be sequenced, with the expectation that E. siliculosus will serve as a genetic and genomic model for brown macroalgae. In 2004, many laboratories, including the Station Biologique in Roscoff and Genoscope, began to sequence the genome of E. siliculosus. Ectocarpus has been used by researchers to study the evolution of complex multicellularity in brown algae. With the study of Ectocarpus came the discovery of multiple genetic and genomic resources that apply to all species of brown algae. Before, the lack of both the proper tools to study genome data and genome data itself halted the progress of understanding brown algal developmental processes at the molecular level. However, due to Ectocarpus being less complex, it is easier to study.    


Iron Storage and Bonding

E. siliculosus is able to accumulate high concentrations of iodide from seawater. The carbon storage system of brown algae is unusual, involving the accumulation of reserves of
mannitol Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to lo ...
and the β-1,3-glucan laminarin rather than α-1,4-glucans such as starch or glycogen. The mannitol pathway was probably most likely a speciation event in the brown algal lineage via a horizontal transfer event from
actinobacteria The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to so ...
, along with another key metabolic pathway in brown algae, alginate biosynthesis. This species of Ectocarpus has been shown to bind iron on its cells with non-specificity. This iron ion shell allows the algae to store and have a constant source of iron regardless of the conditions of the surrounding environment. This adaptation is important because this method of iron uptake is similar to that of terrestrial organisms and differs from the methods typically used in the marine environment such as
siderophores Siderophores (Greek: "iron carrier") are small, high-affinity iron-chelating compounds that are secreted by microorganisms such as bacteria and fungi. They help the organism accumulate iron. Although a widening range of siderophore functions is no ...
.


Sexual Characteristics

Ectocarpus is a species known for its evolution of sex-based gene expression. It has also been found to have a low level of phenotypic
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
. Having a low level of sexual dimorphism means that two sexes of a species do not have different characteristics. This has been backed up by the findings that Ectocarpus has female genes that evolve as rapidly as their male genes. This is also backed up by the findings that the consistency in the patterns that scientists found with its sexual systems relating to UV haploid systems.  


References

{{Taxonbar, from=Q2703732 Ectocarpales