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===寄生虫和有性生殖的宿主===

===寄生虫和有性生殖的宿主===

[[Host–parasite coevolution]] is the coevolution of a [[host (biology)|host]] and a [[parasite]]. A general characteristic of many viruses, as [[obligate parasite]]s, is that they coevolved alongside their respective hosts. Correlated mutations between the two species enter them into an evolution arms race. Whichever organism, host or parasite, that cannot keep up with the other will be eliminated from their habitat, as the species with the higher average population fitness survives. This race is known as the [[Red Queen hypothesis]]. The Red Queen hypothesis predicts that sexual reproduction allows a host to stay just ahead of its parasite, similar to the [[Red Queen's race]] in ''[[Through the Looking-Glass]]'': "it takes all the running ''you'' can do, to keep in the same place". The host reproduces sexually, producing some offspring with immunity over its parasite, which then evolves in response.

[[Host–parasite coevolution|宿主和寄生者的共同演化]]是[[host (biology)|宿主]]和[[parasite|寄生者]]的共同演化。<ref name="Woolhouse">{{cite journal |doi=10.1038/ng1202-569 |last1=Woolhouse |first1=M. E. J. |last2=Webster |first2=J. P. |last3=Domingo |first3=E. |last4=Charlesworth|first4=B. |last5=Levin |first5=B. R. |title=Biological and biomedical implications of the coevolution of pathogens and their hosts |journal=[[Nature Genetics]] |date=December 2002 |pmid=12457190 |volume=32 |issue=4 |pages=569–77 |url=http://www.era.lib.ed.ac.uk/bitstream/1842/689/2/Charlesworth_Woolhouse.pdf|hdl=1842/689 |s2cid=33145462 |hdl-access=free }}</ref>许多病毒作为[[obligate parasite|专性寄生者]]的一个普遍特征是它们与各自宿主的共同演化。这两个物种之间的相关突变使它们进入了进化的军备竞赛。无论是哪种生物、宿主或寄生物，如果不能跟上其他生物的步伐，它们就会从它们的栖息地消失，而平均适合度较高的物种会幸存下来。这种竞争的假说被称为[[Red Queen hypothesis|红皇后假说]]。<ref>{{cite journal |author=Van Valen, L. |date=1973 |title=A New Evolutionary Law |journal=Evolutionary Theory |volume=1 |pages=1–30}} cited in: [http://pespmc1.vub.ac.be/REDQUEEN.html The Red Queen Principle]</ref>红皇后假说预测有性生殖可以让寄主在寄生者之前保持领先，就像''[[Through the Looking-Glass|爱丽丝镜中奇遇]]''的[[Red Queen's race|红皇后比赛]]一样：“尽你所能跑而仍保留在同一个地方。”。<ref>{{cite book |last=Carroll |first=Lewis |author-link=Lewis Carroll |orig-year=1871 |title=Through the Looking-glass: And what Alice Found There |url=https://books.google.com/books?id=cJJZAAAAYAAJ |publisher=Macmillan |date=1875 |page=42 |quote=it takes all the running ''you'' can do, to keep in the same place.}}</ref>宿主进行有性繁殖，产生一些对寄生物具有免疫力的后代，然后进化为应对措施。<ref>{{cite journal |doi=10.1038/srep10004 |last=Rabajante |first=J. |display-authors=etal |title=Red Queen dynamics in multi-host and multi-parasite interaction system |journal=[[Scientific Reports]] |year=2015 |volume=5 |pages=10004 |pmid=25899168 |pmc=4405699|bibcode=2015NatSR...510004R}}</ref>

 

宿主和寄生者的共同演化是宿主和寄生者的共同演化。<ref name="Woolhouse">{{cite journal |doi=10.1038/ng1202-569 |last1=Woolhouse |first1=M. E. J. |last2=Webster |first2=J. P. |last3=Domingo |first3=E. |last4=Charlesworth|first4=B. |last5=Levin |first5=B. R. |title=Biological and biomedical implications of the coevolution of pathogens and their hosts |journal=[[Nature Genetics]] |date=December 2002 |pmid=12457190 |volume=32 |issue=4 |pages=569–77 |url=http://www.era.lib.ed.ac.uk/bitstream/1842/689/2/Charlesworth_Woolhouse.pdf|hdl=1842/689 |s2cid=33145462 |hdl-access=free }}</ref>许多病毒作为专性寄生者的一个普遍特征是它们与各自宿主的共同演化。这两个物种之间的相关突变使它们进入了进化的军备竞赛。无论是哪种生物、宿主或寄生物，如果不能跟上其他生物的步伐，它们就会从它们的栖息地消失，而平均适合度较高的物种会幸存下来。这种竞争的假说被称为红皇后假说。<ref>{{cite journal |author=Van Valen, L. |date=1973 |title=A New Evolutionary Law |journal=Evolutionary Theory |volume=1 |pages=1–30}} cited in: [http://pespmc1.vub.ac.be/REDQUEEN.html The Red Queen Principle]</ref>红皇后假说预测有性生殖可以让寄主在寄生者之前保持领先，就像''爱丽丝镜中奇遇''的红皇后比赛一样：“尽你所能跑而仍保留在同一个地方。”。<ref>{{cite book |last=Carroll |first=Lewis |author-link=Lewis Carroll |orig-year=1871 |title=Through the Looking-glass: And what Alice Found There |url=https://books.google.com/books?id=cJJZAAAAYAAJ |publisher=Macmillan |date=1875 |page=42 |quote=it takes all the running ''you'' can do, to keep in the same place.}}</ref>宿主进行有性繁殖，产生一些对寄生物具有免疫力的后代，然后进化为应对措施。<ref>{{cite journal |doi=10.1038/srep10004 |last=Rabajante |first=J. |display-authors=etal |title=Red Queen dynamics in multi-host and multi-parasite interaction system |journal=[[Scientific Reports]] |year=2015 |volume=5 |pages=10004 |pmid=25899168 |pmc=4405699|bibcode=2015NatSR...510004R}}</ref>

 

寄生者和宿主的关系可能导致了有性生殖的流行，而不是更有效率的无性生殖。看起来，当寄生者感染宿主时，有性生殖提供了一个更好的机会来发展抵抗性（通过下一代的变异） ，这种可导致适应性的有性生殖变异性在无性生殖中则看不到，这样就更容易会产生另一代易受同一寄生者的感染的机体。<ref>{{cite web |title=Sexual reproduction works thanks to ever-evolving host, parasite relationships |website=PhysOrg |url=https://phys.org/news/2011-07-sexual-reproduction-ever-evolving-host-parasite.html |date=7 July 2011}}</ref><ref>{{cite journal |author1=Morran, L.T. |author2=Schmidt, O.G. |author3=Gelarden, I.A. |author4=Parrish, R.C. II |author5= Lively, C.M. |title=Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex |journal=Science |volume=333 |issue=6039 |pages=216–8 |date=8 July 2011 |id=Science.1206360 |bibcode=2011Sci...333..216M |doi=10.1126/science.1206360 |pmid=21737739 |pmc=3402160}}</ref><ref>{{cite encyclopedia |author=Hogan, C. Michael |date=2010 |url=https://editors.eol.org/eoearth/wiki/Virus |title=Virus |encyclopedia=Encyclopedia of Earth |editor=Cutler Cleveland |editor2=Sidney Draggan}}</ref>宿主和寄生者之间的共同演化可能相应导致了在普遍的群体中的许多遗传多样性，包括血浆多态性、蛋白多态性和组织相容性系统。<ref>{{cite journal |author1=Anderson, R. |author2=May, R. |date=October 1982 |title=Coevolution of hosts and parasites |journal=Parasitology |volume=85 |issue=2 |pages=411–426 |doi=10.1017/S0031182000055360 |pmid=6755367}}</ref>

寄生者和宿主的关系可能导致了有性生殖的流行，而不是更有效率的无性生殖。看起来，当寄生者感染宿主时，有性生殖提供了一个更好的机会来发展抵抗性（通过下一代的变异） ，这种可导致适应性的有性生殖变异性在无性生殖中则看不到，这样就更容易会产生另一代易受同一寄生者的感染的机体。<ref>{{cite web |title=Sexual reproduction works thanks to ever-evolving host, parasite relationships |website=PhysOrg |url=https://phys.org/news/2011-07-sexual-reproduction-ever-evolving-host-parasite.html |date=7 July 2011}}</ref><ref>{{cite journal |author1=Morran, L.T. |author2=Schmidt, O.G. |author3=Gelarden, I.A. |author4=Parrish, R.C. II |author5= Lively, C.M. |title=Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex |journal=Science |volume=333 |issue=6039 |pages=216–8 |date=8 July 2011 |id=Science.1206360 |bibcode=2011Sci...333..216M |doi=10.1126/science.1206360 |pmid=21737739 |pmc=3402160}}</ref><ref>{{cite encyclopedia |author=Hogan, C. Michael |date=2010 |url=https://editors.eol.org/eoearth/wiki/Virus |title=Virus |encyclopedia=Encyclopedia of Earth |editor=Cutler Cleveland |editor2=Sidney Draggan}}</ref>宿主和寄生者之间的共同演化可能相应导致了在普遍的群体中的许多遗传多样性，包括血浆多态性、蛋白多态性和组织相容性系统。<ref>{{cite journal |author1=Anderson, R. |author2=May, R. |date=October 1982 |title=Coevolution of hosts and parasites |journal=Parasitology |volume=85 |issue=2 |pages=411–426 |doi=10.1017/S0031182000055360 |pmid=6755367}}</ref>

{{Main|Brood parasitism}}

{{Main|Brood parasitism}}

[[Brood parasite|Brood parasitism]] demonstrates close coevolution of host and parasite, for example in some [[cuckoo]]s. These birds do not make their own nests, but lay their eggs in nests of other species, ejecting or killing the eggs and young of the host and thus having a strong negative impact on the host's reproductive fitness. Their eggs are camouflaged as eggs of their hosts, implying that hosts can distinguish their own eggs from those of intruders and are in an evolutionary arms race with the cuckoo between camouflage and recognition. Cuckoos are counter-adapted to host defences with features such as thickened eggshells, shorter incubation (so their young hatch first), and flat backs adapted to lift eggs out of the nest.

[[Brood parasite|巢穴寄生]]表明了宿主和寄生者密切的共同演化，例如一些[[cuckoo|杜鹃]]。这些鸟不自己筑巢，而是在其他物种的巢中产卵，排除或杀死寄主的卵和幼鸟，从而对寄主的生殖良性产生严重的负面影响。它们的卵伪装成它们寄主的卵，这意味寄主是能够区分自己的卵和入侵者的卵的，并且与杜鹃处于一种伪装和识别之间进化的军备竞赛中。杜鹃与寄主相反，具有加厚的蛋壳、较短的孵化期（所以它们的幼鸟会先孵化）和适于将蛋提出巢外的平脊等特征。<ref name="Weiblen">{{cite web |last1=Weiblen |first1=George D. |title=Interspecific Coevolution |url=http://geo.cbs.umn.edu/Weiblen2003.pdf |publisher=Macmillan |date=May 2003}}</ref><ref>{{cite journal |last1=Rothstein |first1=S.I |year=1990 |title=A model system for coevolution: avian brood parasitism |journal=Annual Review of Ecology and Systematics |volume=21 |pages=481–508 |doi=10.1146/annurev.ecolsys.21.1.481}}</ref><ref>{{Cite book|title=Cuckoo : cheating by nature|last=Davies, N. B. (Nicholas B.), 1952-|others=McCallum, James (Wildlife artist)|date=7 April 2015|isbn=978-1-62040-952-7|edition=First U.S.|location=New York, NY|oclc=881092849}}</ref>

 

巢穴寄生表明了宿主和寄生者密切的共同演化，例如一些杜鹃。这些鸟不自己筑巢，而是在其他物种的巢中产卵，排除或杀死寄主的卵和幼鸟，从而对寄主的生殖良性产生严重的负面影响。它们的卵伪装成它们寄主的卵，这意味寄主是能够区分自己的卵和入侵者的卵的，并且与杜鹃处于一种伪装和识别之间进化的军备竞赛中。杜鹃与寄主相反，具有加厚的蛋壳、较短的孵化期（所以它们的幼鸟会先孵化）和适于将蛋提出巢外的平脊等特征。<ref name="Weiblen">{{cite web |last1=Weiblen |first1=George D. |title=Interspecific Coevolution |url=http://geo.cbs.umn.edu/Weiblen2003.pdf |publisher=Macmillan |date=May 2003}}</ref><ref>{{cite journal |last1=Rothstein |first1=S.I |year=1990 |title=A model system for coevolution: avian brood parasitism |journal=Annual Review of Ecology and Systematics |volume=21 |pages=481–508 |doi=10.1146/annurev.ecolsys.21.1.481}}</ref><ref>{{Cite book|title=Cuckoo : cheating by nature|last=Davies, N. B. (Nicholas B.), 1952-|others=McCallum, James (Wildlife artist)|date=7 April 2015|isbn=978-1-62040-952-7|edition=First U.S.|location=New York, NY|oclc=881092849}}</ref>

=== 拮抗性的共同演化 ===

=== 拮抗性的共同演化 ===

Antagonistic coevolution is seen in the [[harvester ant]] species ''[[Pogonomyrmex barbatus]]'' and ''[[Pogonomyrmex rugosus]]'', in a relationship both parasitic and mutualistic. The queens are unable to produce worker ants by mating with their own species. Only by crossbreeding can they produce workers. The winged females act as parasites for the males of the other species as their sperm will only produce sterile hybrids. But because the colonies are fully dependent on these hybrids to survive, it is also mutualistic. While there is no genetic exchange between the species, they are unable to evolve in a direction where they become too genetically different as this would make crossbreeding impossible.

拮抗性的共同演化关系在[[harvester ant|收获蚁]]品种中的红收获蚁（''[[Pogonomyrmex barbatus]]''）和罗纹须蚁（''[[Pogonomyrmex rugosus]]''）之间可以看到，它们之间既有寄生关系也有互惠关系。蚁后无法通过与同类交配来繁殖工蚁。只有通过杂交，它们才能繁殖工蚁。有翅膀的雌性对其他物种的雄性像寄生一样，因为它们的精子只会繁殖不育的杂种。但由于殖民完全依赖于这些杂交种的生存，这也是互惠互利的。虽然两个物种之间没有基因交换，但它们不能朝向基因差异太大的方向演化，因为这将使杂交繁殖变得不可能。<ref name="Herrmann Cahan pp. 20141771–20141771">{{cite journal |last1=Herrmann |first1=M. |last2=Cahan |first2=S. H. |title=Inter-genomic sexual conflict drives antagonistic coevolution in harvester ants |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1797 |date=29 October 2014 |doi=10.1098/rspb.2014.1771 |pmid=25355474 |pages=20141771 |pmc=4240986}}</ref>

 

拮抗性的共同演化关系在红收获蚁（''Pogonomyrmex barbatus''）和罗纹须蚁（''Pogonomyrmex rugosus''）之间可以看到，它们之间既有寄生关系也有互惠关系。蚁后无法通过与同类交配来繁殖工蚁。只有通过杂交，它们才能繁殖工蚁。有翅膀的雌性对其他物种的雄性像寄生一样，因为它们的精子只会繁殖不育的杂种。但由于殖民完全依赖于这些杂交种的生存，这也是互惠互利的。虽然两个物种之间没有基因交换，但它们不能朝向基因差异太大的方向演化，因为这将使杂交繁殖变得不可能。<ref name="Herrmann Cahan pp. 20141771–20141771">{{cite journal |last1=Herrmann |first1=M. |last2=Cahan |first2=S. H. |title=Inter-genomic sexual conflict drives antagonistic coevolution in harvester ants |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1797 |date=29 October 2014 |doi=10.1098/rspb.2014.1771 |pmid=25355474 |pages=20141771 |pmc=4240986}}</ref>

==捕食者和猎物==

==捕食者和猎物==

{{Main|Predation}}

{{Main|Predation}}

[[Predator]]s and prey interact and coevolve: the predator to catch the prey more effectively, the prey to escape. The coevolution of the two mutually imposes [[selective pressure]]s. These often lead to an [[evolutionary arms race]] between prey and predator, resulting in [[anti-predator adaptation]]s.

s and prey interact and coevolve: the predator to catch the prey more effectively, the prey to escape. The coevolution of the two mutually imposes s. These often lead to an between prey and predator, resulting in s.

捕食者和猎物互动并共同演化：捕食者去更有效地捕捉猎物，猎物去逃离追捕。两者的共同演化相互施加这一选择压。这往往导致猎物和捕食者之间的进化军备竞赛，并导致反捕食者适应。<ref>{{cite web|title=Predator-Prey Relationships|url=https://necsi.edu/predator-prey-relationships|publisher=New England Complex Systems Institute|access-date=17 January 2017}}</ref>

[[Predator|捕食者]]和猎物互动并共同演化：捕食者去更有效地捕捉猎物，猎物去逃离追捕。两者的共同演化相互施加这一[[selective pressure|选择压]]。这往往导致猎物和捕食者之间的[[evolutionary arms race|进化军备竞赛]]，并导致[[anti-predator adaptation|反捕食者适应]]。<ref>{{cite web|title=Predator-Prey Relationships|url=https://necsi.edu/predator-prey-relationships|publisher=New England Complex Systems Institute|access-date=17 January 2017}}</ref>

The same applies to [[herbivore]]s, animals that eat plants, and the plants that they eat.  [[Paul R. Ehrlich]] and [[Peter H. Raven]] in 1964 proposed the theory of [[escape and radiate coevolution]] to describe the evolutionary diversification of plants and butterflies. In the [[Rocky Mountains]], [[red squirrel]]s and [[crossbill]]s (seed-eating birds) compete for seeds of the [[lodgepole pine]]. The squirrels get at pine seeds by gnawing through the cone scales, whereas the crossbills get at the seeds by extracting them with their unusual crossed mandibles. In areas where there are squirrels, the lodgepole's cones are heavier, and have fewer seeds and thinner scales, making it more difficult for squirrels to get at the seeds. Conversely, where there are crossbills but no squirrels, the cones are lighter in construction, but have thicker scales, making it more difficult for crossbills to get at the seeds. The lodgepole's cones are in an evolutionary arms race with the two kinds of herbivore.

The same applies to [[herbivore]]s, animals that eat plants, and the plants that they eat.  [[Paul R. Ehrlich]] and [[Peter H. Raven]] in 1964 proposed the theory of [[escape and radiate coevolution]] to describe the evolutionary diversification of plants and butterflies. In the [[Rocky Mountains]], [[red squirrel]]s and [[crossbill]]s (seed-eating birds) compete for seeds of the [[lodgepole pine]]. The squirrels get at pine seeds by gnawing through the cone scales, whereas the crossbills get at the seeds by extracting them with their unusual crossed mandibles. In areas where there are squirrels, the lodgepole's cones are heavier, and have fewer seeds and thinner scales, making it more difficult for squirrels to get at the seeds. Conversely, where there are crossbills but no squirrels, the cones are lighter in construction, but have thicker scales, making it more difficult for crossbills to get at the seeds. The lodgepole's cones are in an evolutionary arms race with the two kinds of herbivore.