Fish reproduction

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Typical egg of a bony fish, about 1 mm in diameter. Fish eggs are often released into the water column, where they drift as zooplankton

Fish reproductive organs include testes and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused.[1] There may also be a range of secondary organs that increase reproductive fitness. The genital papilla is a small, fleshy tube behind the anus in some fishes, from which the sperm or eggs are released; the sex of a fish often can be determined by the shape of its papilla.

Anatomy[edit]

Testicles[edit]

Most male fish have two testes of similar size. In the case of sharks, the testes on the right side is usually larger. The primitive jawless fish have only a single testis, located in the midline of the body, although even this forms from the fusion of paired structures in the embryo.[2]

Under a tough membranous shell, the tunica albuginea, the testis of some teleost fish, contains very fine coiled tubes called seminiferous tubules. The tubules are lined with a layer of cells (germ cells) that from puberty into old age, develop into sperm cells (also known as spermatozoa or male gametes). The developing sperm travel through the seminiferous tubules to the rete testis located in the mediastinum testis, to the efferent ducts, and then to the epididymis where newly created sperm cells mature (see spermatogenesis). The sperm move into the vas deferens, and are eventually expelled through the urethra and out of the urethral orifice through muscular contractions.

However, most fish do not possess seminiferous tubules. Instead, the sperm are produced in spherical structures called sperm ampullae. These are seasonal structures, releasing their contents during the breeding season, and then being reabsorbed by the body. Before the next breeding season, new sperm ampullae begin to form and ripen. The ampullae are otherwise essentially identical to the seminiferous tubules in higher vertebrates, including the same range of cell types.[2]

In terms of spermatogonia distribution, the structure of teleosts testes has two types: in the most common, spermatogonia occur all along the seminiferous tubules, while in Atherinomorph fish they are confined to the distal portion of these structures. Fish can present cystic or semi-cystic spermatogenesis in relation to the release phase of germ cells in cysts to the seminiferous tubules lumen.[1]

Ovaries[edit]

Many of the features found in ovaries are common to all vertebrates, including the presence of follicular cells and tunica albuginea There may be hundreds or even millions of fertile eggs present in the ovary of a fish at any given time. Fresh eggs may be developing from the germinal epithelium throughout life. Corpora lutea are found only in mammals, and in some elasmobranch fish; in other species, the remnants of the follicle are quickly resorbed by the ovary.[2] The ovary of teleosts is often contains a hollow, lymph-filled space which opens into the oviduct, and into which the eggs are shed.[2] Most normal female fish have two ovaries. In some elasmobranchs, only the right ovary develops fully. In the primitive jawless fish, and some teleosts, there is only one ovary, formed by the fusion of the paired organs in the embryo.[2]

Fish ovaries may be of three types: gymnovarian, secondary gymnovarian or cystovarian. In the first type, the oocytes are released directly into the coelomic cavity and then enter the ostium, then through the oviduct and are eliminated. Secondary gymnovarian ovaries shed ova into the coelom from which they go directly into the oviduct. In the third type, the oocytes are conveyed to the exterior through the oviduct.[3] Gymnovaries are the primitive condition found in lungfish, sturgeon, and bowfin. Cystovaries characterize most teleosts, where the ovary lumen has continuity with the oviduct.[1] Secondary gymnovaries are found in salmonids and a few other teleosts.

Physiology[edit]

See also: Reproductive organs in fish, Spawning, Ichthyoplankton

Oogonia development in teleosts fish varies according to the group, and the determination of oogenesis dynamics allows the understanding of maturation and fertilisation processes. Changes in the nucleus, ooplasm, and the surrounding layers characterize the oocyte maturation process.[1]

Postovulatory follicles are structures formed after oocyte release; they do not have endocrine function, present a wide irregular lumen, and are rapidly reabsorbed in a process involving the apoptosis of follicular cells. A degenerative process called follicular atresia reabsorbs vitellogenic oocytes not spawned. This process can also occur, but less frequently, in oocytes in other development stages.[1]

Some fish are hermaphrodites, having both testes and ovaries either at different phases in their life cycle or, as in hamlets, have them simultaneously.

Over 97% of all known fish are oviparous,[4] that is, the eggs develop outside the mother's body. Examples of oviparous fish include salmon, goldfish, cichlids, tuna, and eels. In the majority of these species, fertilisation takes place outside the mother's body, with the male and female fish shedding their gametes into the surrounding water. However, a few oviparous fish practice internal fertilisation, with the male using some sort of intromittent organ to deliver sperm into the genital opening of the female, most notably the oviparous sharks, such as the horn shark, and oviparous rays, such as skates. In these cases, the male is equipped with a pair of modified pelvic fins known as claspers.

Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of 1 millimetre (0.039 in). The eggs are generally surrounded by the extraembryonic membranes but do not develop a shell, hard or soft, around these membranes. Some fish have thick, leathery coats, especially if they must withstand physical force or desiccation. These type of eggs can also be very small and fragile.

The newly hatched young of oviparous fish are called larvae. They are usually poorly formed, carry a large yolk sac (for nourishment) and are very different in appearance from juvenile and adult specimens. The larval period in oviparous fish is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on zooplankton prey, a process which depends on typically inadequate zooplankton density, starving many larvae.

In ovoviviparous fish the eggs develop inside the mother's body after internal fertilisation but receive little or no nourishment directly from the mother, depending instead on the yolk. Each embryo develops in its own egg. Familiar examples of ovoviviparous fish include guppies, angel sharks, and coelacanths.

Some species of fish are viviparous. In such species the mother retains the eggs and nourishes the embryos. Typically, viviparous fish have a structure analogous to the placenta seen in mammals connecting the mother's blood supply with that of the embryo. Examples of viviparous fish include the surf-perches, splitfins, and lemon shark. Some viviparous fish exhibit oophagy, in which the developing embryos eat other eggs produced by the mother. This has been observed primarily among sharks, such as the shortfin mako and porbeagle, but is known for a few bony fish as well, such as the halfbeak Nomorhamphus ebrardtii.[5] Intrauterine cannibalism is an even more unusual mode of vivipary, in which the largest embryos eat weaker and smaller siblings. This behavior is also most commonly found among sharks, such as the grey nurse shark, but has also been reported for Nomorhamphus ebrardtii.[5]

In many species of fish, fins have been modified to allow Internal fertilisation.

Aquarists commonly refer to ovoviviparous and viviparous fish as livebearers.

  • Many fish species are hermaphrodites. Synchronous hermaphrodites possess both ovaries and testes at the same time. Sequential hermaphrodites have both types of tissue in their gonads, with one type being predominant while the fish belongs to the corresponding gender.

Strategies[edit]

Sexual strategies[edit]

Spawning strategies[edit]

Spawning grounds[edit]

See also: Spawning ground

Examples[edit]

Female goldfish spawn (discharge) eggs into the water, encouraged by male goldfish who simultaneously discharge sperm which externally fertilizes the eggs
Within two or three days, the vulnerable goldfish eggs hatch into larvae, and rapidly develop into fry
Goldfish

Goldfish, like all cyprinids, are egg-layers. They usually start breeding after a significant temperature change, often in spring. Males chase females, prompting them to release their eggs by bumping and nudging them. As the female goldfish spawns her eggs, the male goldfish stays close behind fertilizing them. Their eggs are adhesive and attach to aquatic vegetation. The eggs hatch within 48 to 72 hours. Within a week or so, the fry begins to assume its final shape, although a year may pass before they develop a mature goldfish colour; until then they are a metallic brown like their wild ancestors. In their first weeks of life, the fry grow quickly—an adaptation born of the high risk of getting devoured by the adult goldfish.

Carp

A member of the Cyprinidae family, carp spawn in times between April and August, largely dependent upon the climate and conditions they live in. Oxygen levels of the water, availability of food, size of each fish, age, number of times the fish has spawned before and water temperature are all factors known to effect when and how many eggs each carp will spawn at any one time.[6]

Siamese fighting fish

Prior to spawning, male Siamese fighting fish build bubble nests of varying sizes at the surface of the water. When a male becomes interested in a female, he will flare his gills, twist his body, and spread his fins. The female darkens in colour and curves her body back and forth. The act of spawning takes place in a "nuptial embrace" where the male wraps his body around the female, each embrace resulting in the release of 10-40 eggs until the female is exhausted of eggs. The male, from his side, releases milt into the water and fertilization takes place externally. During and after spawning, the male uses his mouth to retrieve sinking eggs and deposit them in the bubble nest (during mating the female sometimes assists her partner, but more often she will simply devour all the eggs that she manages to catch). Once the female has released all of her eggs, she is chased away from the male's territory, as it is likely that she'll eat the eggs due to hunger.[7] The eggs then remain in the male's care. He keeps them in the bubble nest, making sure none fall to the bottom and repairing the nest as needed. Incubation lasts for 24–36 hours, and the newly hatched larvae remain in the nest for the next 2–3 days, until their yolk sacs are fully absorbed. Afterwards the fry leave the nest and the free-swimming stage begins.[8]

References[edit]

  1. ^ a b c d e Guimaraes-Cruz, Rodrigo J., Rodrigo J.; Santos, José E. dos; Santos, Gilmar B. (July–Sept 2005). "Gonadal structure and gametogenesis of Loricaria lentiginosa Isbrücker (Pisces, Teleostei, Siluriformes)". Rev. Bras. Zool. 22 (3): 556–564. doi:10.1590/S0101-81752005000300005. ISSN 0101-8175.  Check date values in: |date= (help)
  2. ^ a b c d e Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 385–386. ISBN 0-03-910284-X. 
  3. ^ Brito, M.F.G.; Bazzoli, N. (2003). "Reproduction of the surubim catfish (Pisces, Pimelodidae) in the São Francisco River, Pirapora Region, Minas Gerais, Brazil". Arquivo Brasileiro de Medicina Veterinária e Zootecnia 55 (5): 624–633. doi:10.1590/S0102-09352003000500018. ISSN 0102-0935. 
  4. ^ Peter Scott: Livebearing Fishes, p. 13. Tetra Press 1997. ISBN 1-56465-193-2
  5. ^ a b Meisner, A & Burns, J: Viviparity in the Halfbeak Genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae). Journal of Morphology 234, pp. 295–317, 1997
  6. ^ http://www.carp.me.uk Carp Spawning Information
  7. ^ Leong, Paul (2004). http://www.cbsbettas.org/doc/articles/Tips_on_Spawning_Bettas.htm. Retrieved on March 13, 2009.
  8. ^ Rainwate FL and Miller EJ (1967) "Courtship and reproductive behavior of the Siamese fighting fish, Betta splendens Regan" Proceedings of the Oklahoma Academy of Science, Oklahoma State University.

Further references[edit]

External links[edit]