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1
1.1
Ovaries
Anatomy
Testes
1.3
1 ANATOMY
Eggs
Salmon eggs
in dierent stages of development. In some only a few
cells grow on top of the yolk, in the lower right the blood
vessels surround the yolk and in the upper left the black
eyes are visible.
Diagram of a Salmon Fry hatching. The larva has grown around the remains
sh egg: A. vitelline membrane B. chorion C. yolk D. oil of the yolk and the remains of the soft, transparent egg are discarded.
globule E. perivitelline space F. embryo
The eggs of sh and amphibians are jellylike. Cartilagenous sh (sharks, skates, rays, chimaeras) eggs are
fertilized internally and exhibit a wide variety of both
internal and external embryonic development. Most sh
species spawn eggs that are fertilized externally, typically
with the male inseminating the eggs after the female
lays them. These eggs do not have a shell and would dry
out in the air. Even air-breathing amphibians lay their
eggs in water, or in protective foam as with the Coast
foam-nest treefrog, Chiromantis xerampelina.
This young
male spinner shark has claspers, a modication to the
pelvic ns which also function as intromittent organs
1.4
Intromittent organs
3.1
Ovuliparity
3
In sh, fertilisation of eggs can be either external or internal. In many species of sh, ns have been modied
to allow Internal fertilisation. Similarly, development of
the embryo can be external or internal, although some
species show a change between the two at various stages
of embryo development. Thierry Lod described reproductive strategies in terms of the development of the zygote and the interrelationship with the parents; there are
ve classications - ovuliparity, oviparity, ovo-viviparity,
histotrophic viviparity and hemotrophic viviparity.[10]
3.1 Ovuliparity
eggs (ova),
Similar organs with similar characteristics are found Ovuliparity means the female lays unfertilised
[10]
which
must
then
be
externally
fertilised.
Examples
of
in other shes, for example the andropodium in the
ovuliparous
sh
include
salmon,
goldsh,
cichlids,
tuna
[7]
Hemirhamphodon or in the Goodeidae.
and eels. In the majority of these species, fertilisation
Claspers are found on the males of cartilaginous shes. takes place outside the mothers body, with the male and
They are the posterior part of the pelvic ns that have female sh shedding their gametes into the surrounding
also been modied to function as intromittent organs, and water.
are used to channel semen into the females cloaca during
copulation. The act of mating in sharks usually includes
raising one of the claspers to allow water into a siphon 3.2 Oviparity
through a specic orice. The clasper is then inserted
into the cloaca, where it opens like an umbrella to anchor Main article: Oviparity
its position. The siphon then begins to contract expelling
water and sperm.[8][9]
Oviparity is where fertilisation occurs internally and so
the female sheds zygotes (or newly developing embryos)
into the water,[10] often with important outer tissues
2 Physiology
added. Over 97% of all known sh are oviparous,[11] In
oviparous sh, internal fertilisation requires the male to
Oogonia development in teleosts sh varies according to use some sort of intromittent organ to deliver sperm into
the group, and the determination of oogenesis dynam- the genital opening of the female. Examples include the
ics allows the understanding of maturation and fertilisa- oviparous sharks, such as the horn shark, and oviparous
tion processes. Changes in the nucleus, ooplasm, and rays, such as skates. In these cases, the male is equipped
the surrounding layers characterize the oocyte maturation with a pair of modied pelvic ns known as claspers.
process.[1]
Marine sh can produce high numbers of eggs which are
Postovulatory follicles are structures formed after oocyte often released into the open water column. The eggs have
release; they do not have endocrine function, present a an average diameter of 1 millimetre (0.039 in). The eggs
wide irregular lumen, and are rapidly reabsorbed in a are generally surrounded by the extraembryonic memprocess involving the apoptosis of follicular cells. A de- branes but do not develop a shell, hard or soft, around
generative process called follicular atresia reabsorbs vitel- these membranes. Some sh have thick, leathery coats,
logenic oocytes not spawned. This process can also oc- especially if they must withstand physical force or descur, but less frequently, in oocytes in other development iccation. These type of eggs can also be very small and
fragile.
stages.[1]
Some sh are hermaphrodites, having both testes and
ovaries either at dierent phases in their life cycle or, as
in hamlets, have them simultaneously.
Reproductive strategies
Egg of lamprey
Egg of catshark (mermaids purse)
Egg of bullhead shark
Egg of chimaera
3 REPRODUCTIVE STRATEGIES
3.3
Hermaphroditism
Ovoviviparity
3.4
Viviparity
Female
groupers change their sex to male if no male is available
3.6
Sexual parasitism
4 INBREEDING
the species. They live and remain reproductively functional as long as the female lives, and can take part in
multiple spawnings.[27] This extreme sexual dimorphism
ensures, when the female is ready to spawn, she has a mate
immediately available.[29] Multiple males can be incorporated into a single individual female with up to eight males
in some species, though some taxa appear to have a one
male per female rule.[27]
One explanation for the evolution of sexual parasitism
is that the relative low density of females in deep-sea
environments leaves little opportunity for mate choice
among anglersh. Females remain large to accommodate
fecundity, as is evidenced by their large ovaries and eggs.
Males would be expected to shrink to reduce metabolic
costs in resource-poor environments and would develop
highly specialized female-nding abilities. If a male manages to nd a female parasitic attachment, then it is ultimately more likely to improve lifetime tness relative to
free living, particularly when the prospect of nding future mates is poor. An additional advantage to parasitism
is that the males sperm can be used in multiple fertilizations, as he stays always available to the female for mating. Higher densities of male-female encounters might
correlate with species that demonstrate facultative parasitism or simply use a more traditional temporary contact
mating.[30]
3.7
Parthenogenesis
to the ospring. Because gynogenetic species are all female, activation of their eggs requires mating with males
of a closely related species for the needed stimulus. The
Amazon molly, (pictured), reproduces by gynogenesis.
3.8 Others
The elkhorn sculpin (Alcichthys elongatus) is a marine
teleost with a unique reproductive mode called internal gametic association. Sperm are introduced into the
ovary by copulation and then enter the micropylar canal
of ovulated eggs in the ovarian cavity. However, actual
sperm-egg fusion does not occur until the eggs have been
released into sea water.[36]
4 Inbreeding
4.1 Inbreeding depression
The eect of inbreeding on reproductive behavior was
studied in the poeciliid sh Heterandria formosa.[37] One
generation of full-sib mating was found to decrease reproductive performance and likely reproductive success
of male progeny. Other traits that displayed inbreeding
depression were ospring viability and maturation time
of both males and females.
7
fertilisation.[40] In competitions between sperm from an
unrelated male and from a full sibling male, a significant bias in paternity towards the unrelated male was
observed.[40]
Inbreeding depression is considered to be due largely
to the expression of homozygous deleterious recessive
mutations.[41] Outcrossing between unrelated individuals
results in the benecial masking of deleterious recessive
mutations in progeny.[42]
Sexual strategies
Within two or three days, the vulnerable goldsh eggs hatch into
larvae, and rapidly develop into fry
Goldsh
Spawning strategies
Spawning grounds
Examples
Female goldsh spawn (discharge) eggs into the water, encouraged by male goldsh who simultaneously discharge sperm
which externally fertilizes the eggs
10
but more often she will simply devour all the eggs that [7]
she manages to catch). Once the female has released all
of her eggs, she is chased away from the males territory,
as it is likely that she'll eat the eggs due to hunger.[44] The
eggs then remain in the males care. He keeps them in [8]
the bubble nest, making sure none fall to the bottom and
repairing the nest as needed. Incubation lasts for 2436 [9]
hours, and the newly hatched larvae remain in the nest for
the next 23 days, until their yolk sacs are fully absorbed.
Afterwards the fry leave the nest and the free-swimming
stage begins.[45]
[10]
Siamese ghting sh build bubble nests of varying
sizes.
A pair of Siamese ghting sh spawning under their
bubble nest.
One-day-old Siamese ghting sh larvae in a bubble
nest - their yolk sacs have not yet been absorbed
A 15-day-old free-swimming fry of a Siamese ghting sh
See also
Evolution of sexual reproduction
10
References
REFERENCES
[32] Vrijenhoek, R.C., R.M. Dawley, C.J. Cole, and J.P. Bogart. 1989. A list of the known unisexual vertebrates,
pp. 19-23 in: Evolution and Ecology of Unisexual Vertebrates. R.M. Dawley and J.P. Bogart (eds.) Bulletin 466,
New York State Museum, Albany, New York
[33] Chapman, Demian D.; Shivji, Mahmood S.; Louis, Ed;
Sommer, Julie; Fletcher, Hugh; Prodhl, Paulo A. (2007).
Virgin birth in a hammerhead shark. Biology Letters.
3 (4): 425427. doi:10.1098/rsbl.2007.0189. PMC
2390672 . PMID 17519185.
[34] Robinson, D. P.; Baverstock, W.; Al-Jaru, A.; Hyland, K.; Khazanehdari, K. A. (2011). Annually recurring parthenogenesis in a zebra shark Stegostoma fasciatum". Journal of Fish Biology. 79 (5): 1376
1382. doi:10.1111/j.1095-8649.2011.03110.x. PMID
22026614.
[35] Chapman, D. D.; Firchau, B.; Shivji, M. S. (2008).
Parthenogenesis in a large-bodied requiem shark, the
blacktip. Journal of Fish Biology. 73 (6): 14731477.
doi:10.1111/j.1095-8649.2008.02018.x.
[28] Gould, Stephen Jay (1983). Hens Teeth and Horses Toes.
New York: W. W. Norton & Company. ISBN 0-39301716-8.
[29] Theodore W. Pietsch. Precocious sexual parasitism in
the deep sea ceratioid anglersh, Cryptopsaras couesi
Gill. Archived from the original on 28 August 2008. Retrieved 31 July 2008.
[30] Miya, Masaki; Pietsch, Theodore W; Orr, James W;
Arnold, Rachel J; Satoh, Takashi P; Shedlock, Andrew M;
Ho, Hsuan-Ching; Shimazaki, Mitsuomi; Yabe, Mamoru;
Nishida, Mutsumi (1 January 2010). Evolutionary history of anglershes (Teleostei: Lophiiformes): a mitogenomic perspective. BMC Evolutionary Biology. 10 (1):
58. doi:10.1186/1471-2148-10-58. PMC 2836326 .
PMID 20178642.
[31] Hubbs, C. L.; Hubbs, L. C. (1932).
Apparent parthenogenesis in nature, in a form
of sh of hybrid origin.
Science.
76
Bibcode:1932Sci....76..628H.
(1983):
628630.
doi:10.1126/science.76.1983.628. PMID 17730035.
[39] Gallardo JA, Neira R (2005). Environmental dependence of inbreeding depression in cultured Coho salmon
(Oncorhynchus kisutch): aggressiveness, dominance and
intraspecic competition. Heredity (Edinb). 95 (6):
44956. doi:10.1038/sj.hdy.6800741. PMID 16189545.
[40] Fitzpatrick JL, Evans JP (2014). Postcopulatory inbreeding avoidance in guppies. J. Evol. Biol. 27 (12):
258594. doi:10.1111/jeb.12545. PMID 25387854.
[41] Charlesworth D, Willis JH (2009). The genetics of inbreeding depression. Nat. Rev. Genet. 10 (11): 78396.
doi:10.1038/nrg2664. PMID 19834483.
[42] Bernstein H, Hopf FA, Michod RE (1987). The molecular basis of the evolution of sex. Adv. Genet. 24: 32370.
doi:10.1016/s0065-2660(08)60012-7. PMID 3324702.
[43] http://www.carp.me.uk Carp Spawning Information
[44] Leong, Paul (2004).
http://www.cbsbettas.org/doc/
articles/Tips_on_Spawning_Bettas.htm. Retrieved on
March 13, 2009.
10
12
[45] Rainwate FL and Miller EJ (1967) Courtship and reproductive behavior of the Siamese ghting sh, Betta splendens Regan" Proceedings of the Oklahoma Academy of
Science, Oklahoma State University.
11
Further references
12
External links
EXTERNAL LINKS
11
13
13.1
13.2
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13.3
Content license