The perivitellinefluid is an extracellular fluid found in the eggs of most gastropods and constitutes the main source of nutrition and defense for their embryos. It replaces the egg yolk of other animals, which in snail eggs is reduced to non-nutritive proteinaceous granules with putative enzymatic function. [1]
During embryonic development the perivitelline fluid is ingested macropinocytotically by the embryos and the resulting phagosomes fuse with β-granules containing hydrolytic enzymes, which digest the perivitelline fluid components. [1] [2] [3] [4] [5]
The perivitelline fluid is synthesized by the albumen gland of female snails (also known as albumen gland-capsule gland complex or uterine gland), an accessory gland from the reproductive tract. Fertilized oocytes enter the albumen gland and, on their way out, are coated with the perivitelline fluid. [1] [6] [7] The amount of perivitelline fluid per egg vary considerably among species. [1] However, the amount of perivitelline fluid per egg is constant within a given species. [8] In this regard, it has been shown in Pomacea apple snails that during the reproductive season, when the nutrient precursors decrease in the albumen gland due to successive ovipositions, females tend to reduce the number of eggs per clutch but not the amount allocated to each egg. [9]
The perivitelline fluid contains predominantly galactogen, proteins, and calcium. [8] [10] [11] [12]
Carbohydrate is invariably the most abundant component of the perivitelline fluid. Specifically, the eggs of most gastropod accumulate the polysaccharide galactogen, [8] [10] [11] [12] [13] which would provide the main energy source for the developing embryo. A small amount of soluble glucose was also detected in some species. [11] [12]
Proteins, called perivitellins, are the second most abundant component of the perivitelline fluid. Perivitellins are also a source of nutrients for snail embryos [11] [14] and play a role in protection against pathogens [12] [14] [15] [16] and predators, and include non-digestible perivitellins, toxins and protease inhibitors. [12] [17] [18] [19] [20] [21] [22] [23] [24] [25] These proteins were thoroughly studied in apple snails from the genus Pomacea , where they were originally grouped in two most abundant protein fractions perivitellin-1 or PV1, perivitellin-2 or PV2 (comprising approximately 70% of total protein), and a heterogeneous fraction dubbed perivitellin-3 or PV3 fraction. [26] [27] Recent proteomic analyses, however, showed that the perivitelline fluid from Pomacea snails has between 34-38 different proteins with a wide variety of functions. [28] [29] [30]
Lipids are a minor component, mostly represented by membrane lipids, indicating that snails do not use lipids as a major energy reserve during reproduction. [11] [12] Apart from structural lipids, some eggs also contain carotenoid pigments, notably astaxanthin. [12] [17] [27] These lipidic pigments have been associated with antioxidant and photoprotective functions, [27] [31] [32] and also provide Pomacea eggs with the typical bright color that would function as a warning signal (i.e. aposematism) to deter predators. [27] [33] [34] [35]
Among the inorganic components, calcium ion is the most abundant in the perivitelline fluid. As these snails have direct development, calcium needs to be stored to allow the snail to develop the shell during organogenesis. Besides, calcium is the main component of the eggshell of those snails with aerial oviposition. [36]
Ampullariidae, whose members are commonly known as apple snails, is a family of large freshwater snails that includes the mystery snail species. They are aquatic gastropod mollusks with a gill and an operculum. These snails simultaneously have a gill and a lung as functional respiratory structures, which are separated by a division of the mantle cavity. This adaptation allows these animals to be amphibious. Species in this family are considered gonochoristic, meaning that each individual organism is either male or female.
A trypsin inhibitor (TI) is a protein and a type of serine protease inhibitor (serpin) that reduces the biological activity of trypsin by controlling the activation and catalytic reactions of proteins. Trypsin is an enzyme involved in the breakdown of many different proteins, primarily as part of digestion in humans and other animals such as monogastrics and young ruminants. Serpins – including trypsin inhibitors – are irreversible and suicide substrate-like inhibitors.
Pila is a genus of large freshwater snails with an operculum, African and Asian apple snails, aquatic gastropod mollusks in the family Ampullariidae, the apple snails.
Pila ampullacea, is a species of freshwater snail with an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails.
Pomacea is a genus of freshwater snails with gills and an operculum, aquatic gastropod mollusks in the family Ampullariidae, the apple snails. The genus is native to the Americas; most species in this genus are restricted to South America.
Pomacea canaliculata, commonly known as the golden apple snail or the channeled apple snail, is a species of large freshwater snail with gills and an operculum, an aquatic gastropod mollusc in the family Ampullariidae, the apple snails. South American in origin, this species is considered to be in the top 100 of the "World's Worst Invasive Alien Species". It is also ranked as the 40th worst alien species in Europe and the worst alien species of gastropod in Europe.
Pomacea haustrum, common name the titan applesnail, is a species of large freshwater snail with an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the applesnails.
Pomacea diffusa, common name the spike-topped apple snail, is a species of freshwater snail, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails.
Pomacea paludosa, common name the Florida applesnail, is a species of freshwater snail with an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails.
The AB toxins are two-component protein complexes secreted by a number of pathogenic bacteria, though there is a pore-forming AB toxin found in the eggs of a snail. They can be classified as Type III toxins because they interfere with internal cell function. They are named AB toxins due to their components: the "A" component is usually the "active" portion, and the "B" component is usually the "binding" portion. The "A" subunit possesses enzyme activity, and is transferred to the host cell following a conformational change in the membrane-bound transport "B" subunit. These proteins consist of two independent polypeptides, which correspond to the A/B subunit moieties. The enzyme component (A) enters the cell through endosomes produced by the oligomeric binding/translocation protein (B), and prevents actin polymerisation through ADP-ribosylation of monomeric G-actin.
Pomacea maculata is a species of large freshwater snail with an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails.
Galactogen is a polysaccharide of galactose that functions as energy storage in pulmonate snails and some Caenogastropoda. This polysaccharide is exclusive of the reproduction and is only found in the albumen gland from the female snail reproductive system and in the perivitelline fluid of eggs.
Perivitellins are egg proteins found in the perivitelline fluid of many gastropods. They are multifunctional complexes providing the developing embryo with nutrition, protection from the environment, and defense against predators.
Ovorubin is the most abundant perivitellin of the perivitelline fluid from Pomacea canaliculata snail eggs. This glyco-lipo-caroteno protein complex is a approx. 300 kDa multimer of a combination of multiple copies of six different ~30 kDa subunits.
Pomacea maculata perivitellin-1 (PmPV1) is the most abundant perivitellin found in the perivitelline fluid from Pomacea maculata snail eggs. This glyco-lipo-caroteno protein is an approx. 294 kDa multimer of a combination of multiple copies of six different ~30 kDa subunits. PmPV1 account >60% of the total proteins found in the Pomacea maculata eggs.
Scalarin (PsSC) is the most abundant perivitellin of the perivitelline fluid from Pomacea scalaris eggs. This glyco-lipo-caroteno protein is an approx. 380 kDa multimer combining multiple copies of six different 24-35 kDa subunits.
Perivitellin-2 (PV2) is a pore-forming toxin present in the egg perivitelline fluid of the apple snails Pomacea maculata (PmPV2) and Pomacea canaliculata (PcPV2). This protein, called perivitellin, is massively accumulated in the eggs. As a toxin PV2 protects eggs from predators, but it also nourishes the developing snail embryos.
Pomacea columellaris is a South American species of freshwater snail in the apple snail family, Ampullariidae.
Pomacea fasciata is a freshwater snail in the Ampullariidae family. It is located in Jamaica, Guadeloupe, and Hispaniola. It lays white eggs.
Temptin is a protein that acts as a water-borne pheromone in the marine gastropod mollusk Aplysia californica. It is an abundant protein that is synthesized in the albumen gland, and is released in the egg cords during oviposition, along with other proteins called attractin, seductin and enticin. Together, they make up a complex of water-soluble proteins that act together to attract mates for reproduction and induce spawning.