Pregnancy has been traditionally defined as the period of time eggs are incubated in the body after the egg-sperm union. Although the term often refers to placental mammals, it has also been used in the titles of many international, peer-reviewed, scientific articles on fish. Consistent with this definition, there are several modes of reproduction in fish, providing different amounts of parental care. In ovoviviparity, there is internal fertilization and the young are born live but there is no placental connection or significant trophic (feeding) interaction; the mother's body maintains gas exchange but the unborn young are nourished by egg yolk. There are two types of viviparity in fish. In histotrophic viviparity, the zygotes develop in the female's oviducts, but she provides no direct nutrition; the embryos survive by eating her eggs or their unborn siblings. In hemotrophic viviparity, the zygotes are retained within the female and are provided with nutrients by her, often through some form of placenta. In seahorses and pipefish, it is the male that becomes pregnant.
Types of reproduction and pregnancy
Pregnancy has been traditionally defined as the period during which developing embryos are incubated in the body after egg-sperm union. Despite strong similarities between viviparity in mammals, researchers have historically been reluctant to use the term "pregnancy" for non-mammals because of the highly developed form of viviparity in eutherians. Recent research into physiological, morphological and genetic changes associated with fish reproduction provide evidence that incubation in some species is a highly specialized form of reproduction similar to other forms of viviparity. Although the term "pregnancy" often refers to eutherian animals, it has also been used in the titles of many international, peer-reviewed, scientific articles on fish, e.g. Five modes of reproduction can be differentiated in fish based on relations between the zygote(s) and parents:
Ovuliparity: Fertilization of eggs is external; zygotes develop externally. Oviparity: Fertilization of eggs is internal; zygotes develop externally as eggs with large vitellus. Ovoviviparity: Fertilization is internal; zygotes are retained in the female (or male) but without major trophic (feeding) interactions between zygote and parents (there may be minor interactions, such as maintenance of water and oxygen levels). The embryos depend upon their yolk for survival. There are two types of viviparity among fish.
Histotrophic ("tissue eating") viviparity: The zygotes develop in the female's oviducts, but she provides no direct nutrition. The embryos survive by eating her eggs or their unborn siblings. Hemotrophic ("blood eating") viviparity: The zygotes are retained within the female and are provided with nutrients by her, often through some form of placenta. Diagram
Ovoviviparous fish Examples of ovoviviparous fish are many of the squaliform sharks, which include sand sharks, mackerel sharks, nurse sharks, requiem sharks, dog sharks and hammerheads, among others, and the lobe finned coelacanth. Some species of rockfish (Sebastes) and sculpins (Comephoridae) produce rather weak larvae with no egg membrane and are also, by definition, ovoviviparous. Ovoviviparity occurs in most live-bearing bony fishes (Poeciliidae).
Viviparous fish Viviparous fish include the families Goodeidae, Anablepidae, Jenynsiidae, Poeciliidae, Embiotocidae, and some sharks (some species of the requiem sharks, Carcharinidae and the hammerheads, Sphyrnidae, among others). The halfbeaks, Hemiramphidae, are found in both marine and freshwaters and those species that are marine produce eggs with extended filaments that attach to floating or stationary debris, while those that are found in freshwater are viviparous with internal fertilization. The Bythitidae are also viviparous although one species, Dinematichthys ilucoeteoides, is reported to be ovoviviparous. Aquarists commonly refer to ovoviviparous and viviparous fish as "livebearers". Examples include guppies, mollies, moonfish, platys, four-eyed fish, and swordtails. All of these varieties exhibit signs of their pregnancy before the live fry are born. As examples, the female swordtail and guppy will both give birth to anywhere from 20 to 100 live young after a gestation period of four to six weeks, and mollies will produce a brood of 20 to 60 live young after a gestation of six to 10 weeks.
Nutrition during pregnancy Other terms relating to pregnancy in fish relate to the differences in the mode and extent of support the female gives the developing offspring. "Lecithotrophy" (yolk feeding) occurs when the mother provisions the oocyte with all the resources that it needs prior to fertilization, so the egg is independent of the mother. Many members of the fish family Poeciliidae are considered to be lecithotrophic, however, research is increasingly showing that others are matrotrophic. "Aplacental viviparity" occurs when the female retains the embryos during the entire time of development but without any transfer of nutrients to the young. The yolk sac is the only source of nutrients for the developing embryo. There are at least two exceptions to this; some sharks gain nourishment by eating unfertilized eggs produced by the mother (oophagy or egg eating) or by eating their unborn siblings (intra-uterine cannibalism). "Matrotrophy" (mother feeding) occurs when the embryo exhausts its yolk supply early in gestation and the mother provides additional nutrition. Post-fertilization transfer of nutrients has been reported in several species within the genera Gambusia and Poecilia, specifically, G. affinis, G. clarkhubbsi, G. holbrooki, G. gaigei, G. geiseri, G. nobilis, P. formosa, P. latipinna, and P. mexicana. Viviparous fish have developed several ways of providing their offspring with nutrition. "Embryotrophic" or "histrotrophic" nutrition occurs by the production of nutritive fluid, uterine milk, by the uterine lining, which is absorbed directly by the developing embryo. "Hemotrophic" nutrition occurs through the passing of nutritive substances between blood vessels of the mother and embryo that are in close proximity, i.e. a placenta-like organ similar to that found in mammals.
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