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Sexual reproduction

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Sexual reproduction is a process that creates a new organism by combining the genetic material of two organisms. It occurs in eukaryotes.[1][2] Prokaryotes display processes similar to sex (mechanisms for lateral gene transfer such as conjugation, transformation and transduction), but they do not lead to reproduction. Prokaryotes reproduce through asexual reproduction. In multicellular eukaryote organisms, an individual is created anew; in prokaryotes, the initial cell has additional or transformed genetic material. In a process called genetic recombination, genetic material (DNA) originating from two different individuals join up so that homologous sequences are aligned with each other, and this is followed by exchange of genetic information. After the new recombinant chromosome is formed, it is passed on to progeny.

Sexual reproduction is the primary method of reproduction for the vast majority of macroscopic organisms, including almost all animals and plants. The evolution of sexual reproduction is a major puzzle. The first fossilized evidence of sexual reproduction in eukaryotes is from the Stenian period, about 1 to 1.2 billion years ago.[3] There are two main processes during sexual reproduction in eukaryotes: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the restoration of the original number of chromosomes. During meiosis, the chromosomes of each pair usually exchange genetic information to achieve homologous recombination. Evolutionary thought proposes several explanations for why sexual reproduction developed and why it is maintained. These reasons include fighting the accumulation of deleterious mutations, increasing rate of adaptation to changing environments[4] (see the red queen hypothesis), dealing with competition (see the tangled bank hypothesis) or as an adaptation for repairing DNA damage and masking deleterious mutations.[5][6][7] While these ideas about why sexual reproduction has been maintained are generally supported, the ultimate size of the population determines if sexual reproduction is entirely beneficial. Larger populations appear to respond more quickly to benefits obtained through sexual reproduction than smaller population sizes. [8]maintenance of sexual reproduction has been explained by theories that work at several levels of selection, though some of these models remain controversial. New models presented in recent years, however, suggest a basic advantage for sexual reproduction in slowly reproducing, complex organisms, exhibiting characteristics that depend on the specific environment that the given species inhabit, and the particular survival strategies that they employ.[9]

In the first stage of sexual reproduction, "meiosis", the number of chromosomes is reduced from a diploid number (2n) to a haploid number (n). During "fertilization", haploid gametes come together to form a diploid zygote and the original number of chromosomes is restored.

Bacteria and archaea

Three distinct processes in prokaryotes are regarded as similar to eukaryotic sex: 1) bacterial transformation involves the incorporation of foreign DNA into the bacterial chromosome, 2) bacterial conjugation is a transfer of plasmid DNA between bacteria, but the plasmids are rarely incorporated into the bacterial chromosome, 3) Gene transfer and genetic exchange in archaea.

Bacterial transformation involves the recombination of genetic material and its function is mainly associated with DNA repair. Bacterial transformation is a complex process encoded by numerous bacterial genes, and is a bacterial adaptation for DNA transfer.[5][6] This process occurs naturally in at least 40 bacterial species.[10] For a bacterium to bind, take up, and recombine exogenous DNA into its chromosome, it must enter a special physiological state referred to as competence (see Natural competence). Sexual reproduction in early single-celled eukaryotes may have evolved from bacterial transformation,[7] or from a similar process in archaea (see below).

On the other hand, bacterial conjugation is a type of direct transfer of DNA between two bacteria through an external appendage called the conjugation pilus.[11] Bacterial conjugation is controlled by plasmid genes that are adapted for spreading copies of the plasmid between bacteria. The infrequent integration of a plasmid into a host bacterial chromosome, and the subsequent transfer of a part of the host chromosome to another cell do not appear to be bacterial adaptations.[5][12]

Exposure of hyperthermophilic archaeal Sulfolobus species to DNA damaging conditions induces cellular aggregation accompanied by high frequency genetic marker exchange.[13][14] Ajon et al.[14] hypothesized that this cellular aggregation enhances species-specific DNA repair by homologous recombination. DNA transfer in Sulfolobus may be an early form of sexual interaction similar to the more well-studied bacterial transformation systems that also involve species-specific DNA transfer leading to homologous recombinational repair of DNA damage.

Plants

Main article: Plant reproduction

Animals typically produce gametes directly by meiosis. Male gametes are called sperm, and female gametes are called eggs or ova. In animals, fertilization follows immediately after meiosis. Plants on the other hand have mitosis occurring in spores, which are produced by meiosis. The spores germinate into the gametophyte phase. The gametophytes of different groups of plants vary in size; angiosperms have as few as three cells in pollen, and mosses and other so called primitive plants may have several million cells. Plants have an alternation of generations where the sporophyte phase is succeeded by the gametophyte phase. The sporophyte phase produces spores within the sporangium by meiosis.

Flowering plants

Flowers are the sexual organs of flowering plants.

Flowering plants are the dominant plant form on land and they reproduce by sexual and asexual means. Often their most distinguishing feature is their reproductive organs, commonly called flowers. The anther produces pollen grains which contain the male gametophytes (sperm). For pollination to occur, pollen grains must attach to the stigma of the female reproductive structure (carpel), where the female gametophytes (ovules) are located inside the ovary. After the pollen tube grows through the carpel's style, the sex cell nuclei from the pollen grain migrate into the ovule to fertilize the egg cell and endosperm nuclei within the female gametophyte in a process termed double fertilization. The resulting zygote develops into an embryo, while the triploid endosperm (one sperm cell plus two female cells) and female tissues of the ovule give rise to the surrounding tissues in the developing seed. The ovary, which produced the female gametophyte(s), then grows into a fruit, which surrounds the seed(s). Plants may either self-pollinate or cross-pollinate. Nonflowering plants like ferns, moss and liverworts use other means of sexual reproduction.

In 2013, flowers dating from the Cretaceous (100 million years before present) were found encased in amber, the oldest evidence of sexual reproduction in a flowering plant. Microscopic images showed tubes growing out of pollen and penetrating the flower's stigma. The pollen was sticky, suggesting it was carried by insects.[15]

Ferns

Ferns mostly produce large diploid sporophytes with rhizomes, roots and leaves; and on fertile leaves called sporangium, spores are produced. The spores are released and germinate to produce short, thin gametophytes that are typically heart shaped, small and green in color. The gametophytes or thallus, produce both motile sperm in the antheridia and egg cells in separate archegonia. After rains or when dew deposits a film of water, the motile sperm are splashed away from the antheridia, which are normally produced on the top side of the thallus, and swim in the film of water to the archegonia where they fertilize the egg. To promote out crossing or cross fertilization the sperm are released before the eggs are receptive of the sperm, making it more likely that the sperm will fertilize the eggs of different thallus. A zygote is formed after fertilization, which grows into a new sporophytic plant. The condition of having separate sporephyte and gametophyte plants is called alternation of generations. Other plants with similar reproductive means include the Psilotum, Lycopodium, Selaginella and Equisetum.

Bryophytes

The bryophytes, which include liverworts, hornworts and mosses, reproduce both sexually and vegetatively. They are small plants found growing in moist locations and like ferns, have motile sperm with flagella and need water to facilitate sexual reproduction. These plants start as a haploid spore that grows into the dominate form, which is a multicellular haploid body with leaf-like structures that photosynthesize. Haploid gametes are produced in antherida and archegonia by mitosis. The sperm released from the antherida respond to chemicals released by ripe archegonia and swim to them in a film of water and fertilize the egg cells thus producing a zygote. The zygote divides by mitotic division and grows into a sporophyte that is diploid. The multicellular diploid sporophyte produces structures called spore capsules, which are connected by seta to the archegonia. The spore capsules produce spores by meiosis, when ripe the capsules burst open and the spores are released. Bryophytes show considerable variation in their breeding structures and the above is a basic outline. Also in some species each plant is one sex while other species produce both sexes on the same plant.[16]

Fungi

Main article: Mating in fungi
Further information: Fungus § Reproduction

Fungi are classified by the methods of sexual reproduction they employ. The outcome of sexual reproduction most often is the production of resting spores that are used to survive inclement times and to spread. There are typically three phases in the sexual reproduction of fungi: plasmogamy, karyogamy and meiosis.

Animals

Further information: Reproductive system § Animals, Fertilization in animals, and Animal sexual behavior

Insects

Further information: Insect § Reproduction and development
Australian Emperor laying egg, guarded by the male

Insect species make up more than two-thirds of all extant animal species, and most insect species use sex for reproduction, though some species are facultatively parthenogenetic. Many species have sexual dimorphism, while in others the sexes look nearly identical. Typically they have two sexes with males producing spermatozoa and females ova. The ova develop into eggs that have a covering called the chorion, which forms before internal fertilization. Insects have very diverse mating and reproductive strategies most often resulting in the male depositing spermatophore within the female, which stores the sperm until she is ready for egg fertilization. After fertilization, and the formation of a zygote, and varying degrees of development; the eggs are deposited outside the female in many species, or in some, they develop further within the female and live born offspring are produced.

Birds

Further information: Avian reproduction

Mammals

Further information: Reproductive system § Mammals, Sex organ § Mammals, Rut (mammalian reproduction), and Carnivora § Reproductive system

There are three extant kinds of mammals: Monotremes, Placentals and Marsupials, all with internal fertilization. In placental mammals, offspring are born as juveniles: complete animals with the sex organs present although not reproductively functional. After several months or years, the sex organs develop further to maturity and the animal becomes sexually mature. Most female mammals are only fertile during certain periods during their estrous cycle, at which point they are ready to mate. Individual male and female mammals meet and carry out copulation.[citation needed] For most mammals, males and females exchange sexual partners throughout their adult lives.[17][18][19]

Placental mammals

Male placental mammals
Further information: Human male reproductive system, Canine reproduction § Male, and Stallion § Reproductive anatomy
Wikimedia Commons has media related to Mammal male reproductive system.

The mammalian male reproductive system contains two main divisions, the penis and the testicles, the latter of which is where sperm are produced. In humans, both of these organs are outside the abdominal cavity, but they can be primarily housed within the abdomen in other animals. For instance, a dog's penis is covered by a penile sheath except when mating. Having the testicles outside the abdomen best facilitates temperature regulation of the sperm, which require specific temperatures to survive. The external location may also cause a reduction in the heat-induced contribution to the spontaneous mutation rate in male germinal tissue.[20] Sperm are the smaller of the two gametes and are generally very short-lived, requiring males to produce them continuously from the time of sexual maturity until death. The produced sperm are stored in the epididymis until ejaculation. The sperm cells are motile and they swim using tail-like flagella to propel themselves towards the ovum. The sperm follows temperature gradients (thermotaxis)[21] and chemical gradients (chemotaxis) to locate the ovum.

Female placental mammals
Further information: Human female reproductive system, Canine reproduction § Female, and Horse anatomy § Mare
Wikimedia Commons has media related to Mammal female reproductive system.

The mammalian female reproductive system likewise contains two main divisions: the vagina and uterus, which act as the receptacle for the sperm, and the ovaries, which produce the female's ova. All of these parts are always internal. The vagina is attached to the uterus through the cervix, while the uterus is attached to the ovaries via the Fallopian tubes. At certain intervals, the ovaries release an ovum, which passes through the fallopian tube into the uterus.

If, in this transit, it meets with sperm, the egg selects sperm with which to merge; this is termed fertilization. The fertilization usually occurs in the oviducts, but can happen in the uterus itself. The zygote then implants itself in the wall of the uterus, where it begins the processes of embryogenesis and morphogenesis. When developed enough to survive outside the womb, the cervix dilates and contractions of the uterus propel the fetus through the birth canal, which is the vagina.

The ova, which are the female sex cells, are much larger than the sperm and are normally formed within the ovaries of the fetus before its birth. They are mostly fixed in location within the ovary until their transit to the uterus, and contain nutrients for the later zygote and embryo. Over a regular interval, in response to hormonal signals, a process of oogenesis matures one ovum which is released and sent down the Fallopian tube. If not fertilized, this egg is released through menstruation in humans and other great apes, and reabsorbed in other mammals in the estrus cycle.

Gestation
The initial stages of human embryogenesis.
Main articles: Gestation and Pregnancy (mammals)

Gestation, called pregnancy in humans, is the period of time during which the fetus develops, dividing via mitosis inside the female. During this time, the fetus receives all of its nutrition and oxygenated blood from the female, filtered through the placenta, which is attached to the fetus' abdomen via an umbilical cord. This drain of nutrients can be quite taxing on the female, who is required to ingest slightly higher levels of calories. In addition, certain vitamins and other nutrients are required in greater quantities than normal, often creating abnormal eating habits. The length of gestation, called the gestation period, varies greatly from species to species; it is 40 weeks in humans, 56–60 in giraffes and 16 days in hamsters.

Birth
Main article: Birth

Once the fetus is sufficiently developed, chemical signals start the process of birth, which begins with contractions of the uterus and the dilation of the cervix. The fetus then descends to the cervix, where it is pushed out into the vagina, and eventually out of the female. The newborn, which is called an infant in humans, should typically begin respiration on its own shortly after birth. Not long after, the placenta is passed as well. Most mammals eat this, as it is a good source of protein and other vital nutrients needed for caring for the young. The end of the umbilical cord attached to the young's abdomen eventually falls off on its own.

Monotremes

Further information: Monotreme § Reproductive system

Monotremes, only five species of which exist, all from Australia and New Guinea, are mammals that lay eggs. They have one opening for excretion and reproduction called the cloaca. They hold the eggs internally for several weeks, providing nutrients, and then lay them and cover them like birds.

Marsupials

Further information: Marsupial reproductive system

Marsupials' reproductive systems differ markedly from those of placental mammals.[22] The female develops a kind of yolk sac in her womb which delivers nutrients to the embryo. Embryos of some marsupials additionally form placenta-like organs that connect them to the uterine wall, although it is not certain that they transfer nutrients from the mother to the embryo.[23] Pregnancy is very short, typically 4 to 5 weeks, and the embryo is born at a very young stage of development.

Fish

Further information: Fish reproductive anatomy

The vast majority of fish species lay eggs that are then fertilized by the male,[24] some species lay their eggs on a substrate like a rock or on plants, while others scatter their eggs and the eggs are fertilized as they drift or sink in the water column. Some fish species use internal fertilization and then disperse the developing eggs or give birth to live offspring. Fish that have live-bearing offspring include the Guppy and Mollies or Poecilia. Fishes that give birth to live young can be ovoviviparous, where the eggs are fertilized within the female and the eggs simply hatch within the female body, or in seahorses, the male carries the developing young within a pouch, and gives birth to live young.[25] Fishes can also be viviparous, where the female supplies nourishment to the internally growing offspring. Some fish are hermaphrodites, where a single fish is both male and female and can produce eggs and sperm. In hermaphroditic fish, some are male and female at the same time while in other fish they are serially hermaphroditic; starting as one sex and changing to the other. In at least one hermaphroditic species, self-fertilization occurs when the eggs and sperm are released together. Internal self-fertilization may occur in some other species.[26] One fish species does not reproduce by sexual reproduction but uses sex to produce offspring; Poecilia formosa is a unisex species that uses a form of parthenogenesis called gynogenesis, where unfertilized eggs develop into embryos that produce female offspring. Poecilia formosa mate with males of other fish species that use internal fertilization, the sperm does not fertilize the eggs but stimulates the growth of the eggs which develops into embryos.[27]

Reptiles

Further information: Reptiles § Reproduction

Mollusks

Further information: Mollusk § Reproduction

See also

Template:Wikipedia books

Notes

  1. ^ Lodé, Thierry (2011). "Sex is not a solution for reproduction: the libertine bubble theory". BioEssays. 33 (6): 419–422. doi:10.1002/bies.201000125. PMID 21472739.
  2. ^ Lodé, Thierry (2012). "Sex and the origin of genetic exchanges". Trends in Evolutionary Biology. 4. doi:10.4081/eb.2012.e1.
  3. ^ N.J. Buttefield (2000). "Bangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes". Paleobiology. 26 (3): 386–404. doi:10.1666/0094-8373(2000)026<0386:BPNGNS>2.0.CO;2.
  4. ^ Gray, J. C.; Goddard, M. R. (2012). Bonsall, Michael (ed.). "Gene-flow between niches facilitates local adaptation in sexual populations". Ecology Letters. 15 (9): n/a. doi:10.1111/j.1461-0248.2012.01814.x.
  5. ^ a b c Michod RE, Bernstein H, Nedelcu AM; Bernstein; Nedelcu (May 2008). "Adaptive value of sex in microbial pathogens" (PDF). Infect. Genet. Evol. 8 (3): 267–85. doi:10.1016/j.meegid.2008.01.002. PMID 18295550.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. ^ a b Bernstein, Harris; Bernstein, Carol (2010). "Evolutionary Origin of Recombination during Meiosis". BioScience. 60 (7): 498. doi:10.1525/bio.2010.60.7.5.
  7. ^ a b Bernstein H, Bernstein C, Michod RE. (2012) "DNA Repair as the Primary Adaptive Function of Sex in Bacteria and Eukaryotes". Chapter 1, pp. 1–50, in DNA Repair: New Research, Editors S. Kimura and Shimizu S. Nova Sci. Publ., Hauppauge, New York. Open access for reading only. ISBN 978-1-62100-756-2
  8. ^ Colegrave, N (2002). "Sex releases the speed limit on evolution". Nature. 420 (6916): 664–6. Bibcode:2002Natur.420..664C. doi:10.1038/nature01191. PMID 12478292.
  9. ^ Kleiman, Maya; Tannenbaum, Emmanuel (2009). "Diploidy and the selective advantage for sexual reproduction in unicellular organisms". Theory in Biosciences. 128 (4): 249–85. doi:10.1007/s12064-009-0077-9. PMID 19902285.
  10. ^ Lorenz, MG; Wackernagel, W (1994). "Bacterial gene transfer by natural genetic transformation in the environment". Microbiological reviews. 58 (3): 563–602. PMC 372978. PMID 7968924.
  11. ^ Lodé, T (2012). "Have Sex or Not? Lessons from Bacteria". Sexual development : genetics, molecular biology, evolution, endocrinology, embryology, and pathology of sex determination and differentiation. 6 (6): 325–8. doi:10.1159/000342879. PMID 22986519.
  12. ^ Krebs, JE; Goldstein, ES; Kilpatrick, ST (2011). Lewin's GENES X. Boston: Jones and Bartlett Publishers. pp. 289–292. ISBN 9780763766320.
  13. ^ Fröls S, Ajon M, Wagner M, Teichmann D, Zolghadr B, Folea M, Boekema EJ, Driessen AJ, Schleper C, Albers SV. (2008). UV-inducible cellular aggregation of the hyperthermophilic archaeon Sulfolobus solfataricus is mediated by pili formation. Mol Microbiol 70(4):938-952. doi: 10.1111/j.1365-2958.2008.06459.x. PMID 18990182
  14. ^ a b Ajon M, Fröls S, van Wolferen M, Stoecker K, Teichmann D, Driessen AJ, Grogan DW, Albers SV, Schleper C; Fröls; Van Wolferen; Stoecker; Teichmann; Driessen; Grogan; Albers; Schleper (November 2011). "UV-inducible DNA exchange in hyperthermophilic archaea mediated by type IV pili". Mol. Microbiol. 82 (4): 807–17. doi:10.1111/j.1365-2958.2011.07861.x. PMID 21999488. {{cite journal}}: Invalid |display-authors=9 (help)CS1 maint: multiple names: authors list (link)
  15. ^ Poinar Jr., George O; Chambers, Kenton L; Wunderlich, Joerg (10 December 2013). "Micropetasos, a new genus of angiosperms from mid-Cretaceous Burmese amber" (PDF). J. Bot. Res. Inst. Texas. 7 (2): 745–750. {{cite journal}}: Unknown parameter |laydate= ignored (help); Unknown parameter |laysummary= ignored (help)
  16. ^ Jon Lovett Doust; Lesley Lovett Doust (1988). Plant Reproductive Ecology: Patterns and Strategies. Oxford University Press. p. 290. ISBN 9780195063943.
  17. ^ Reichard, U.H. (2002). "Monogamy—A variable relationship" (PDF). Max Planck Research. 3: 62–7. Retrieved 24 April 2013.
  18. ^ Lipton, Judith Eve; Barash, David P. (2001). The Myth of Monogamy: Fidelity and Infidelity in Animals and People. San Francisco: W.H. Freeman and Company. ISBN 0-7167-4004-4.{{cite book}}: CS1 maint: multiple names: authors list (link)
  19. ^ Research conducted by Patricia Adair Gowaty. Reported by Morell, V. (1998). "Evolution of sex: A new look at monogamy". Science. 281 (5385): 1982–1983. doi:10.1126/science.281.5385.1982. PMID 9767050.
  20. ^ Baltz, RH; Bingham, PM; Drake, JW (1976). "Heat mutagenesis in bacteriophage T4: The transition pathway". Proceedings of the National Academy of Sciences of the United States of America. 73 (4): 1269–73. Bibcode:1976PNAS...73.1269B. doi:10.1073/pnas.73.4.1269. PMC 430244. PMID 4797.
  21. ^ Bahat, Anat; Tur-Kaspa, Ilan; Gakamsky, Anna; Giojalas, Laura C.; Breitbart, Haim; Eisenbach, Michael (2003). "Thermotaxis of mammalian sperm cells: A potential navigation mechanism in the female genital tract". Nature Medicine. 9 (2): 149–50. doi:10.1038/nm0203-149. PMID 12563318. {{cite journal}}: Unknown parameter |laydate= ignored (help); Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help)
  22. ^ Iowa State University Biology Dept. Discoveries about Marsupial Reproduction Anna King 2001. webpage (note shows code, html extension omitted)
  23. ^ "Family Peramelidae (bandicoots and echymiperas)".
  24. ^ BONY FISHES - Reproduction
  25. ^ M. Cavendish (2001). Endangered Wildlife and Plants of the World. Marshall Cavendish. p. 1252. ISBN 978-0-7614-7194-3. Retrieved 2013-11-03.
  26. ^ Orlando EF, Katsu Y, Miyagawa S, Iguchi T; Katsu; Miyagawa; Iguchi (2006). "Cloning and differential expression of estrogen receptor and aromatase genes in the self-fertilizing hermaphrodite and male mangrove rivulus, Kryptolebias marmoratus". Journal of Molecular Endocrinology. 37 (2): 353–365. doi:10.1677/jme.1.02101. PMID 17032750.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  27. ^ I. Schlupp, J. Parzefall, J. T. Epplen, M. Schartl; Parzefall; Epplen; Schartl (2006). "Limia vittata as host species for the Amazon molly: no evidence for sexual reproduction". Journal of Fish Biology. 48 (4): 792–795. doi:10.1111/j.1095-8649.1996.tb01472.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)

References

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