HAP2 (hapless 2), also known as GCS1 (generative cell-specific protein 1), is a family of

membrane fusion protein Membrane fusion proteins (not to be confused with chimeric or fusion proteins) are proteins that cause fusion of biological membranes. Membrane fusion is critical for many biological processes, especially in eukaryotic development and viral entry ...

s found in the

sperm cell Sperm (: sperm or sperms) is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail k ...

of diverse

eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

s including ''

Toxoplasma ''Toxoplasma gondii'' () is a species of parasitic alveolate that causes toxoplasmosis. Found worldwide, ''T. gondii'' is capable of infecting virtually all warm-blooded animals, but members of the cat family (felidae) are the only known d ...

'',

thale cress ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally ...

, and fruit flies. This protein is essential for

gamete A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...

fusion, and therefore

fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...

, in these organisms. By eukaryotic taxonomy, it is found in: * Plant in the broad sense (

Archaeplastida The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced ) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It als ...

) ** Land plants (pollen – "male" gamete) -

, numerous other species including

rice Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...

** Green algae ("minus" mating type) - ''

Chlamydomonas ''Chlamydomonas'' ( ) is a genus of green algae consisting of about 150 species of unicellular organism, unicellular flagellates, found in stagnant water and on damp soil, in freshwater, seawater, and even in snow as "snow algae". ''Chlamydom ...

'' ** Red algae - ''

Cyanidioschyzon ''Cyanidioschyzon merolae'' is a small (2μm), club-shaped, unicellular Ploidy, haploid Rhodophyta, red alga adapted to high sulfur acidic hot spring environments (pH 1.5, 45 °C). The cellular architecture of ''C. merolae'' is extremely sim ...

'' * Alveolates **

Apicomplexan The Apicomplexa (also called Apicomplexia; single: apicomplexan) are organisms of a large phylum of mainly parasitic alveolates. Most possess a unique form of organelle structure that comprises a type of non-photosynthetic plastid called an apico ...

s (male gametocytes) *** ''

Plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...

'' *** ''

'' **

Ciliophora The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different ...

ns - ''

Tetrahymena ''Tetrahymena'' is a genus of free-living ciliates, examples of unicellular eukaryotes. The genus Tetrahymena is the most widely studied member of its phylum. It can produce, store and react with different types of hormones. ''Tetrahymena'' cel ...

'' *

Euglenozoa Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by four major groups, ''i.e.,'' Kinetoplastea, ...

ns - ''

Trypanosoma ''Trypanosoma'' is a genus of kinetoplastids (class Trypanosomatidae), a monophyletic group of unicellular parasitic flagellate protozoa. Trypanosoma is part of the phylum Euglenozoa. The name is derived from the Ancient Greek ''trypano-'' (b ...

'' * Amorphea ** Opisthokonta *** Animals **** Cnidarians - ''Hydra'' **** Arthropods - Insects - fruit flies and bees **** Acorn worms - '' Saccoglossus'' ** Amoebozoa - ''

Dictyostelium discoideum ''Dictyostelium discoideum'' is a species of soil-dwelling Amoeboid, amoeba belonging to the phylum Amoebozoa, infraphylum Mycetozoa. Commonly referred to as slime mold, ''D. discoideum'' is a eukaryote that transitions from a collection of unic ...

'' slime mold (in a gene-duplicated setup with 3 mating types) For a full list of organisms in which it occurs, consul
InterPro #IPR018928 - taxonomy

Origin

HAP2/GCS1 belongs to the broader group of fusexins, which includes such proteins as ''

C. elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' ( ...

'' developmental fusogens EFF-1 and AFF-1, haloarchaeal Fsx1, and viral "class II" fusion proteins. In an unrooted phylogenetic tree from 2021, HAP2/GCS1 and EFF-1/AFF-1 occupy two ends of the tree, the middle being occupied by viral sequences; this suggests that they may have been acquired separately. The latest structure-based unrooted phylogenetic tree of Brukman ''et al.'' (2022), which takes into account the newly-discovered archaeal sequences, shows that Fsx1 groups with HAP2/GCS1, and that they are separated from EFF-1 by a number of viral sequences. Based on where the root is placed, a number of different hypotheses regarding the history of these families – their horizontal transfer and vertical inheritance – can be generated. Older comparisons excluding archaeal sequences would strongly favor an interpretation where HAP2/GCS1 is acquired from a virus, but the grouping of Fsx1 with HAP2/GCS1 has allowed the possibility of a much more ancient source.

References

{{Reflist Protein families