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distinction between unconditioned versus conditioned reward; indeed, most primary rewards are learned and thus conditioned (foods are primary rewards that are typically learnt). ... Pleasure is not only one of the three main reward functions but also provides a definition of reward. As homeostasis explains the functions of only a limited number of rewards, the prevailing reason why particular stimuli, objects, events, situations, and activities are rewarding may be pleasure. This applies first of all to sex (who would engage in the ridiculous gymnastics of reproductive activity if it were not for the pleasure) and to the primary homeostatic rewards of food and liquid, and extends to money, taste, beauty, social encounters and nonmaterial, internally set, and intrinsic rewards. ... Desire makes behavior purposeful and directs it towards identifiable goals. Thus desire is the emotion that helps to actively direct behavior towards known rewards, whereas pleasure is the passive experience that derives from a received or anticipated reward. Desire has multiple relations to pleasure; it may be pleasant in itself (I feel a pleasant desire), and it may lead to pleasure (I desire to obtain a pleasant object). Thus pleasure and desire have distinctive characteristics but are closely intertwined. They constitute the most important positive emotions induced by rewards. They prioritize our conscious processing and thus constitute important components of behavioral control. These emotions are also called liking (for pleasure) and wanting (for desire) in addiction research (471) and strongly support the learning and approach generating functions of reward. ... Some of the stimuli and events that are pleasurable in humans may not even evoke pleasure in animals but act instead through innate mechanisms. We simply do not know. Nevertheless, the invention of pleasure and desire by evolution had the huge advantage of allowing a large number of stimuli, objects, events, situations, and activities to be attractive. This mechanism importantly supports the primary reward functions in obtaining essential substances and mating partners.
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prostaglandins. Some mammalian OR genes are known to be expressed in other tissues, including the testis, tongue, brain and placenta17. However, the functional significance of such 'ectopic expression' of OR genes is not definitively known. TAARs are also expressed in the MOE. These receptors were first identified as brain receptors for the trace amines, a collection of amines that are present at low concentrations in the central nervous system18. TAARs were originally suspected to be involved in psychiatric disorders19 but are now known to function as a second class of olfactory receptors10. Some mouse TAARs recognize volatile amines that are present in urine, and it seems that the TAARs function to detect ligands associated with social cues10. ... Most mammals possess an additional olfactory organ called the vomeronasal organ (VnO). ... The VnO was previously thought to be a specialized organ for pheromone detection, but it is now known that the VnO and MOE share some overlapping functions22. ... However, at least one of the five V1R genes is expressed in the human olfactory mucosa72. Furthermore, a recent study suggests that these five genes can activate an OR-like signal transduction pathway in a heterologous expression system73. It is therefore possible that the products of these genes function as pheromone or olfactory receptors. Adult humans do not have a VnO but seem to be sensitive to pheromones74. Another interesting observation is that chickens have no functional or non-functional V1R and V2R genes or a VnO75, although birds use pheromones for mate choice and other behaviours76. It is possible that some OR genes in the MOE are able to detect pheromones, as in humans74,77.
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sexual behaviour develops over time as people learn to associate stimuli such as bodily features, personality, and contextual cues with genitally-induced sexual pleasure.7 Adolescence is arguably the most critical phase in sexual development ... Popular belief holds that humans also respond to some distal sexual incentive stimuli (breasts, pheromones) in an unconditioned manner, but this has been difficult to evaluate empirically (Box 1) ... Sexual wanting in both rats and humans involves interaction between gonadal hormones and external stimuli that become sexual incentives through association with genitally-induced sexual reward; pleasurable genital stimulation is thus a major factor in sexual learning ...
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humans. Both mouse Taar3 and Taar5 detect compounds (isoamylamine and trimethylamine, respectively) that are enriched in male versus female mouse urine. Isoamylamine in male urine is reported to act as a pheromone, accelerating puberty onset in female mice . The authors suggest the Taar family has a chemosensory function that is distinct from odorant receptors with a role associated with the detection of social cues. ... The evolutionary pattern of the TAAR gene family is characterized by lineage-specific phylogenetic clustering . These characteristics are very similar to those observed in the olfactory GPCRs and vomeronasal (V1R, V2R) GPCR gene families.
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human babies must rotate inside the birth canal in order to squeeze through the birthing canal and fit through the pelvic planes. This process is known as a rotational birth, and while it is not a process unique to humans, humans are unique in that nearly all human babies undergo this process out of necessity. A primary hypothesis for why this process and others occur, causing human births to be drastically more difficult than other mammals is known as the
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to recognize new ligands, with almost an entire teleost clade losing the canonical amine-recognition motif. Furthermore, while some TAARs detect aversive odors, TAAR-mediated behaviors can vary across species. ... The ability of particular TAARs to mediate aversion and attraction behavior provides an exciting opportunity for mechanistic unraveling of odor valence encoding.
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Once the fetus has sufficiently developed, chemical signals start the process of birth. This begins with contractions of the uterus and 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
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Importantly, three ligands identified activating mouse Taars are natural components of mouse urine, a major source of social cues in rodents. Mouse Taar4 recognizes β-phenylethylamine, a compound whose elevation in urine is correlated with increases in stress and stress responses in both rodents and
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OR genes are predominantly expressed in sensory neurons of the main olfactory epithelium (MOE) in the nasal cavity. Mammals detect many types of chemicals in the air and some in the water as odorants, whereas fishes recognize water-soluble molecules, such as amino acids, bile acids, sex steroids and
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presumably mediates aversion to trimethylamine, which is known to act as an hTAAR5 agonist and to possess a foul, fishy odor that is aversive to humans; however, hTAAR5 is not the only olfactory receptor that is responsible for trimethylamine olfaction in humans. As of
December 2015, hTAAR5-mediated
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While mice produce gender-specific amounts of urinary TMA levels and were attracted by TMA, this odor is repellent to rats and aversive to humans , indicating that there must be species-specific functions. ... Furthermore, a homozygous knockout of murine TAAR5 abolished the attraction behavior
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Roles for another receptor are supported by TAAR5-independent trimethylamine anosmias in humans . ... Several TAARs detect volatile and aversive amines, but the olfactory system is capable of discarding ligand-based or function-based constraints on TAAR evolution. Particular TAARs have mutated
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Sexual behavior follows hormonal imbalances, at least in men, but is also strongly based on pleasure. To acquire and follow these primary alimentary and mating rewards is the main reason why the brain's reward system has evolved in the first place. Note that "primary" reward does not refer to the
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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. Sperm are the smaller of the two gametes and are generally very short-lived, requiring males to
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Human babies are unique in the animal kingdom due to their large head size relative to their bodies. This has an effect on the birthing process for humans as the bipedal gait of a human causes the birthing canal to be relatively narrow and twisted in the middle. As a result, the vast majority of
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The sexual pleasure cycle adheres to the basic structure of pleasure cycles related to other rewards (such as food), and can therefore also be expressed in terms of motivation–consummation–satiety or wanting–liking–inhibition (Figure 1; Box 2).6,11,1 ... Similar to other forms of learning,
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In most female mammals, the act of copulation is controlled by several innate neurobiological processes, including the motor sexual reflex of lordosis. In males, the act of copulation is more complex, because some learning is necessary, but the innate processes (retrocontrol of
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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
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to TMA . Thus, it is concluded that TAAR5 itself is sufficient to mediate a behavioral response at least in mice. ... Whether the TAAR5 activation by TMA elicits specific behavioral output like avoidance behavior in humans still needs to be examined.
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Pfaff Donald W., Schwartz-Giblin Susan, Maccarthy
Margareth M., Kow Lee-Ming : Cellular and molecular mechanisms of female reproductive behaviors, in Knobil Ernest, Neill Jimmy D. : The physiology of reproduction, Raven Press, 2nd edition,
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Wallis MC, Waters PD, Delbridge ML, Kirby PJ, Pask AJ, Grützner F, Rens W, Ferguson-Smith MA, Graves JA, et al. (2007). "Sex determination in platypus and echidna: autosomal location of SOX3 confirms the absence of SRY from monotremes".
952:) occurs when an animal starts to associate bodily features, personality, contextual cues, and other stimuli with genitally-induced sexual pleasure. Once formed, these associations in turn impinge upon both sexual wanting and sexual liking.
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The duration of copulation varies significantly between mammal species, and may be correlated with body mass, lasting longer in large mammals than in small mammals. The duration of copulation may also be correlated with the length of the
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Giallombardo, Andres, 2009 New
Cretaceous mammals from Mongolia and the early diversification of Eutheria Ph.D. dissertation, Columbia University, 2009402 pages; AAT 3373736 (abstract) The origin of Placental Mammals, Cimolestidae,
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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".
964:, rhythmic movement of the pelvis, detection of female pheromones) are specific to copulation. These innate processes direct heterosexual copulation. Female lordosis behaviour became secondary in
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1193:"Self-Love Is Important, but We Mammals Are Stuck With Sex - Some female birds, reptiles and other animals can make a baby on their own. But for mammals like us, eggs and sperm need each other"
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are capable of detecting. While sex pheromones clearly play a role in modifying sexual behavior in some mammals, the capacity for general pheromone detection and the involvement of
735:. During gametogenesis in mammals many genes encoding proteins that take part in DNA repair mechanisms show enhanced or specialized expression These mechanisms include meiotic
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Meisel Robert L., Sachs
Benjamin D. : The physiology of male sexual behavior. In Knobil Ernest, Neill Jimmy D. The physiology of reproduction, Raven Press, 2nd edition, 1994
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of mammals are specialized to produce milk, a liquid used by newborns as their primary source of nutrition. The monotremes branched early from other mammals and do not have the
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Pacifici, Michela; Santini, Luca; Marco, Moreno Di; Baisero, Daniele; Francucci, Lucilla; Marasini, Gabriele
Grottolo; Visconti, Piero; Rondinini, Carlo (2013-11-13).
334:. 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.
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and is non-functional in humans. Mammals usually copulate in a dorso-ventral posture, although some primate species copulate in a ventro-vental posture.
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901:("inhibition"); these behavioral states are regulated in mammals by reward-based sexual learning, fluctuations in various neurochemicals (i.e.,
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and other nutrients are required in greater quantities than normal, often creating abnormal eating habits. The length of gestation, called the
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Nei M, Niimura Y, Nozawa M (December 2008). "The evolution of animal chemosensory receptor gene repertoires: roles of chance and necessity".
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979:. Despite the fact that humans do not possess this organ, adult humans appear to be sensitive to certain mammalian pheromones that putative
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1161:. This review also noted that the behavioral response evoked by a TAAR can vary across species. For example, TAAR5 mediates attraction to
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through the penis into the female reproductive tract during copulation. Ejaculation usually occurs after only one intromission in humans,
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Sexual maturity and thus the earliest age at which mammals can reproduce varies dramatically across species. Members of the rodent family
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2183:"A positive relationship between baculum length and prolonged intromission patterns in mammals." 动物学报 50.4 (2004): 490-503.
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seen in most mammals, but they do have mammary glands. The young lick the milk from a mammary patch on the mother's belly.
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1477:"Sperm Use Heat Sensors To Find The Egg; Weizmann Institute Research Contributes To Understanding Of Human Fertilization"
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1751: • Rats show a similar pattern of brain activation to humans in response to cues related to sexual reward
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Georgiadis JR, Kringelbach ML, Pfaus JG (September 2012). "Sex for fun: a synthesis of human and animal neurobiology".
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2067:"Timberol® Inhibits TAAR5-Mediated Responses to Trimethylamine and Influences the Olfactory Threshold in Humans"
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948:: motivation − excitement; consummation − plateau and orgasm; satiety − refraction. Sexual learning (a form of
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matures one ovum which is released and sent down the oviduct. If not fertilized, this egg is released through
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A review of studies involving non-human animals indicated that TAARs in the olfactory epithelium can mediate
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Marsupials' reproductive systems differ markedly from those of placental mammals, though it is probably the
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Figure 2: Table of ligands, expression patterns, and species-specific behavioral responses for each TAAR
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Iowa State
University Biology Dept. Discoveries about Marsupial Reproduction Anna King 2001. webpage
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in gonads (testicles in males and ovaries in females). Sperm are produced by the process of
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Primate sexuality: Comparative studies of the
Prosimians, Monkeys, Apes, and Human Beings
216:, but they can be primarily housed within the abdomen in other animals. For instance, a
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Philosophical
Transactions of the Royal Society of London. Series B: Biological Sciences
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can be quite taxing on the female, who is required to ingest slightly higher levels of
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1557:"The development of the external features of the platypus (Ornithorhynchus anatinus)"
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Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans
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on its own shortly after birth. Not long after, the placenta is passed as well.
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Proceedings of the
National Academy of Sciences of the United States of America
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in humans, is the period of time during which the fetus develops, dividing via
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of a platypus are more like those of a chicken than those of a therian mammal.
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except when mating. Having the testicles outside the abdomen best facilitates
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The mammalian female reproductive system contains three main divisions: the
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Size matters: the (negative) allometry of copulatory duration in mammals
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The mammalian male reproductive system contains two main divisions, the
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trimethylamine aversion has not been examined in published research.
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Animals, including mammals, produce gametes (sperm and egg) through
2165:." Biological Journal of the Linnean Society 87.2 (2006): 185-193.
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Wallrabenstein I, Singer M, Panten J, Hatt H, Gisselmann G (2015).
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inside the female. During this time, the fetus receives all of its
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itself in the wall of the uterus, where it begins the processes of
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take even longer, with the longest duration being recorded for the
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can reach sexual maturity in 1–2 months, e.g. the Norway lemming (
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897:"), reward consummation also known as pleasure ("liking"), and
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condition found in viviparous mammals, including non-placental
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1371:"Heat mutagenesis in bacteriophage T4: The transition pathway"
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different from that of most other mammals. In particular, the
1770:"Neuronal reward and decision signals: from theories to data"
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Manger, Paul R.; Hall, Leslie S.; Pettigrew, John D. (1998).
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Sperm Competition and the Evolution of Animal Mating systems
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in mice and aversion to trimethylamine in rats. In humans, h
318:, the egg selects sperm with which to merge; this is termed
158:), which reaches maturity at an age of only about 23 years.
2197:
Hormones and Reproduction of Vertebrates, Volume 5: Mammals
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and oxygenated blood from the female, filtered through the
245:. The sperm cells are motile and they swim using tail-like
2033:. International Union of Basic and Clinical Pharmacology.
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These behavioral states correlate with the phases of the
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Sexual Selection and the Origins of Human Mating Systems
929:), and gonadal hormone cycles and further influenced by
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kids will stay with their mother until they are weaned.
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Most mammals are viviparous, giving birth to live young
307:, while the uterus is attached to the ovaries via the
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DIXSON33, Alan, N. YHOL T. Jenna, and Matt Anderson.
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1650:Baarends WM, van der Laan R, Grootegoed JA (2001).
1500:"Primate pelvic anatomy and implications for birth"
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involved in the olfactive detection of social cues.
975:that is involved in pheromone detection, including
2194:Norris, David O.; Lopez, Kristin H. (2024-08-08).
528:bones, forcing them to produce undeveloped young.
233:until death. The produced sperm are stored in the
212:. In humans, both of these organs are outside the
283:, which act as the receptacle for the sperm, the
1142:; these TAARs putatively function as a class of
731:. These processes are outlined in the article
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1843:. Oxford University Press, 2nd edition, 2012.
496:, only five species of which exist, all from
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134:) take about a year to reach maturity while
2154:Stallmann, Robert R., and A. H. Harcourt. "
1604:Australian Mammal Society (December 1978).
253:. The sperm follows temperature gradients (
229:produce them continuously from the time of
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1369:Baltz, RH; Bingham, PM; Drake, JW (1976).
2227:. Springer Science & Business Media.
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1652:"DNA repair mechanisms and gametogenesis"
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1088:Dolphin § Reproduction and sexuality
358:, and reabsorbed in other mammals in the
1043:Evolution of descended testes in mammals
168:Reproductive system § Other mammals
88:, and gives birth to an underdeveloped (
2277:Encyclopedia of Behavioral Neuroscience
1191:Preston, Elizabeth (13 February 2024).
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96:, located in the front of the mother's
68:Viviparous mammals are in the subclass
1289:10.1146/annurev.genet.42.110807.091714
80:infraclasses. A marsupial has a short
2037:from the original on 23 February 2014
1268:Marshall Graves, Jennifer A. (2008).
1205:from the original on 13 February 2024
1063:Animal sexual behavior § Mammals
7:
2031:"Trace amine receptor: Introduction"
1078:Carnivora § Reproductive system
489:Monotreme § Reproductive system
180:Carnivora § Reproductive system
2884:Sexual selection in scaled reptiles
2224:Comparative Vertebrate Reproduction
989:pheromones in human sexual behavior
314:If, in this transit, it meets with
146:) require more than 10 years. Some
84:period, typically shorter than its
1120:(TAARs) that are expressed in the
1048:Sexual reproduction § Mammals
550:. During embryonic development, a
462:in humans, should typically begin
411:, which is attached to the fetus'
46:, lay eggs. The monotremes have a
14:
2131:Advances in the Study of Behavior
1485:(Press release). 3 February 2003.
1336:10.3897/natureconservation.5.5734
754:Copulation (zoology) § In mammals
737:homologous recombinational repair
249:to propel themselves towards the
2171:10.1111/j.1095-8312.2006.00566.x
1118:trace amine-associated receptors
873:
864:
853:
844:
833:
824:
813:
804:
793:
784:
773:
764:
752:This section is an excerpt from
702:
687:
672:
657:
645:
630:
618:
603:
589:
575:
72:; those living today are in the
2221:Lombardi, Julian (2012-12-06).
1853:Smith, Robert L. (2012-12-02).
1498:Trevathan, Wenda (2015-03-05).
1351:from the original on 2021-01-26
1319:"Generation length for mammals"
295:, which mainly consists of the
2248:Dixson, Alan F. (2012-01-26).
1880:Dixson, Alan F. (2009-05-15).
1023:in mammal species that do not
1:
1116:In humans and other animals,
991:has not yet been determined.
562:forms, although it lacks the
538:Marsupial reproductive system
287:, which produce the female's
108:Maturity and reproductive age
2319:10.1016/0010-7824(73)90023-1
2092:10.1371/journal.pone.0144704
1970:Liberles SD (October 2015).
1610:. Australian Mammal Society.
554:forms in all marsupials. In
176:Rut (mammalian reproduction)
2559:semelparity and iteroparity
2128:Naguib, Marc (2020-04-19).
1157:behavioral responses to an
1098:Domestic sheep reproduction
946:human sexual response cycle
3201:
2303:"Coitus-induced ovulation"
1988:10.1016/j.conb.2015.01.001
1786:10.1152/physrev.00023.2014
1073:Equine reproductive system
1038:Mammal reproductive system
883:Mating postures of mammals
751:
535:
486:
450:
381:
257:) and chemical gradients (
165:
2650:Non-reproductive behavior
2301:Jöchle, Wolfgang (1973).
1607:Australian Mammal Society
1277:Annual Review of Genetics
1239:10.1007/s10577-007-1185-3
1093:Llama § Reproduction
909:also known as "wanting";
727:and eggs are produced by
184:Human reproductive system
2280:. Elsevier. 2010-06-03.
1912:Nature Reviews. Genetics
1053:Sex organ § Mammals
560:chorioallantoic placenta
552:choriovitelline placenta
172:Sex organ § Mammals
48:sex determination system
3180:Reproduction in mammals
2368:Animal sexual behaviour
1724:10.1038/nrurol.2012.151
1712:Nature Reviews. Urology
1669:10.1530/reprod/121.1.31
971:Most mammals possess a
504:, are mammals that lay
427:. In addition, certain
2517:traumatic insemination
1573:10.1098/rstb.1998.0270
1516:10.1098/rstb.2014.0065
1396:10.1073/pnas.73.4.1269
379:
374:The initial stages of
261:) to locate the ovum.
226:temperature regulation
196:Male placental mammals
27:
2645:Interspecies breeding
1976:Curr. Opin. Neurobiol
1774:Physiological Reviews
536:Further information:
487:Further information:
373:
166:Further information:
22:
3159:Short-beaked echidna
2899:side-blotched lizard
2416:sexual ornamentation
1138:, including certain
1122:olfactory epithelium
985:olfactory epithelium
960:intromission in the
950:associative learning
941:) in some mammals.
354:in humans and other
2986:Homosexual behavior
2951:Homosexual behavior
2806:Spawning strategies
2606:Bateman's principle
2426:sexy son hypothesis
2404:hormonal motivation
2399:reproductive system
2389:Sexual reproduction
2083:2015PLoSO..1044704W
1567:(1372): 1115–1125.
1387:1976PNAS...73.1269B
1323:Nature Conservation
1227:Chromosome Research
1144:pheromone receptors
1126:olfactory receptors
1083:Canine reproduction
1068:Pregnancy (mammals)
625:Female domestic cat
477:obstetrical dilemma
388:Pregnancy (mammals)
376:human embryogenesis
162:Reproductive system
122:) in 39 days. Many
2934:Breeding behaviour
2655:Fisher's principle
2480:sexual intercourse
2421:handicap principle
2161:2022-04-20 at the
2134:. Academic Press.
1768:Schultz W (2015).
1626:2012-09-05 at the
1510:(1663): 20140065.
1443:10.1038/nm0203-149
1198:The New York Times
1058:Human reproduction
981:pheromone receptor
939:lordosis behaviour
516:. Like marsupial "
380:
156:Balaena mysticetus
28:
3167:
3166:
2981:Lordosis behavior
2861:Frog reproduction
2811:Polyandry in fish
2591:Sexual dimorphism
2512:sperm competition
2431:Fisherian runaway
2411:Courtship display
2287:978-0-08-045396-5
2261:978-0-19-954464-6
2234:978-1-4615-4937-6
2207:978-0-443-15985-5
2141:978-0-12-820726-0
1893:978-0-19-156973-9
1866:978-0-323-14313-4
973:vomeronasal organ
927:sexual attraction
269:Female placentals
189:Placental mammals
3192:
3137:ringtailed lemur
2996:African wild dog
2969:Sexual selection
2929:Sexual selection
2849:Sexual selection
2490:pseudocopulation
2384:Sexual selection
2361:
2354:
2347:
2338:
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2119:
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2104:
2094:
2077:(12): e0144704.
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2018:
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1640:Zalambdalestidae
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1398:
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1307:
1305:
1299:. Archived from
1274:
1265:
1259:
1258:
1221:
1215:
1214:
1212:
1210:
1188:
1171:
1114:
1021:induce ovulation
983:proteins in the
877:
868:
857:
848:
837:
828:
817:
808:
797:
788:
777:
768:
706:
691:
676:
664:Male and female
661:
649:
634:
622:
607:
593:
579:
558:, an additional
433:gestation period
419:. This drain of
271:
270:
220:is covered by a
214:abdominal cavity
142:) and dolphins (
3200:
3199:
3195:
3194:
3193:
3191:
3190:
3189:
3170:
3169:
3168:
3163:
3142:sexual swelling
3038:European badger
2955:
2915:
2870:
2835:
2792:
2659:
2628:Sexual conflict
2569:hermaphroditism
2370:
2365:
2335:
2334:
2300:
2299:
2295:
2288:
2274:
2273:
2269:
2262:
2247:
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2235:
2220:
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2215:
2208:
2193:
2192:
2188:
2180:
2176:
2163:Wayback Machine
2153:
2149:
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2127:
2126:
2122:
2064:
2063:
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2038:
2029:
2028:
2021:
2014:
1969:
1968:
1951:
1924:10.1038/nrg2480
1909:
1908:
1901:
1894:
1879:
1878:
1874:
1867:
1852:
1851:
1847:
1838:
1834:
1829:
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1767:
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1759:
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1748:
1746:
1709:
1708:
1695:
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1628:Wayback Machine
1619:
1615:
1603:
1602:
1598:
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1497:
1496:
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1475:
1431:Nature Medicine
1427:
1426:
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1303:
1272:
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1190:
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1185:
1180:
1175:
1174:
1148:
1147:
1115:
1111:
1106:
1034:
1029:
1028:
1019:Copulation can
887:
886:
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749:
741:mismatch repair
725:spermatogenesis
717:
710:
707:
698:
692:
683:
677:
668:
662:
653:
650:
641:
635:
626:
623:
614:
608:
599:
594:
585:
580:
564:chorionic villi
540:
534:
491:
485:
472:
455:
449:
439:and 16 days in
390:
382:Main articles:
368:
273:
268:
267:
243:vasa deferentia
231:sexual maturity
198:
191:
186:
164:
110:
52:sex chromosomes
17:
12:
11:
5:
3198:
3196:
3188:
3187:
3182:
3172:
3171:
3165:
3164:
3162:
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3156:
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3139:
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3129:
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3119:
3114:
3106:
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3082:
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2608:
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2482:
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2466:
2465:
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2455:
2450:
2445:
2438:Mating systems
2435:
2434:
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2428:
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2418:
2408:
2407:
2406:
2401:
2396:
2386:
2380:
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2372:
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2366:
2364:
2363:
2356:
2349:
2341:
2333:
2332:
2313:(6): 523–564.
2293:
2286:
2267:
2260:
2254:. OUP Oxford.
2240:
2233:
2213:
2206:
2186:
2174:
2147:
2140:
2120:
2052:
2019:
1949:
1918:(12): 951–63.
1899:
1892:
1886:. OUP Oxford.
1872:
1865:
1845:
1832:
1823:
1813:
1780:(3): 853–951.
1757:
1693:
1642:
1632:
1613:
1596:
1547:
1490:
1488:
1487:
1420:
1381:(4): 1269–73.
1361:
1309:
1306:on 2012-09-04.
1260:
1233:(8): 949–959.
1216:
1182:
1181:
1179:
1176:
1173:
1172:
1163:trimethylamine
1108:
1107:
1105:
1102:
1101:
1100:
1095:
1090:
1085:
1080:
1075:
1070:
1065:
1060:
1055:
1050:
1045:
1040:
1033:
1030:
1027:spontaneously.
977:sex pheromones
935:motor reflexes
931:sex pheromones
915:sexual arousal
911:norepinephrine
882:
881:
872:
871:
863:
862:
861:
852:
851:
843:
842:
841:
832:
831:
823:
822:
821:
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811:
803:
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792:
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783:
782:
781:
772:
771:
763:
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684:
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671:
669:
663:
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654:
651:
644:
642:
636:
629:
627:
624:
617:
615:
609:
602:
600:
595:
588:
586:
581:
574:
533:
530:
520:", monotreme "
484:
481:
471:
468:
451:Main article:
448:
445:
417:umbilical cord
367:
364:
272:
265:
197:
194:
190:
187:
163:
160:
130:) and bovids (
128:family Canidae
109:
106:
59:mammary glands
15:
13:
10:
9:
6:
4:
3:
2:
3197:
3186:
3183:
3181:
3178:
3177:
3175:
3160:
3157:
3155:
3152:
3150:
3147:
3143:
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3138:
3135:
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3130:
3128:
3125:
3123:
3120:
3118:
3115:
3113:
3110:
3109:
3107:
3103:
3100:
3099:
3098:
3095:
3091:
3088:
3087:
3086:
3083:
3081:
3080:Spotted hyena
3078:
3076:
3073:
3071:
3068:
3064:
3061:
3059:
3056:
3054:
3051:
3049:
3046:
3045:
3044:
3041:
3039:
3036:
3034:
3031:
3029:
3026:
3022:
3019:
3017:
3014:
3012:
3009:
3007:
3004:
3002:
2999:
2997:
2994:
2993:
2992:
2989:
2987:
2984:
2982:
2979:
2975:
2972:
2971:
2970:
2967:
2966:
2964:
2962:
2958:
2952:
2949:
2945:
2942:
2940:
2937:
2936:
2935:
2932:
2930:
2927:
2926:
2924:
2922:
2918:
2912:
2909:
2907:
2904:
2900:
2897:
2895:
2892:
2890:
2887:
2886:
2885:
2882:
2881:
2879:
2877:
2873:
2867:
2864:
2862:
2859:
2855:
2852:
2851:
2850:
2847:
2846:
2844:
2842:
2838:
2832:
2829:
2827:
2824:
2822:
2819:
2817:
2814:
2812:
2809:
2807:
2804:
2803:
2801:
2799:
2795:
2787:
2786:penis fencing
2784:
2782:
2779:
2777:
2774:
2773:
2771:
2769:
2766:
2762:
2759:
2757:
2756:apophallation
2754:
2753:
2752:
2749:
2747:
2744:
2740:
2737:
2735:
2732:
2730:
2727:
2726:
2725:
2722:
2718:
2715:
2714:
2713:
2710:
2704:
2701:
2700:
2699:
2696:
2694:
2691:
2689:
2686:
2684:
2681:
2679:
2676:
2675:
2674:
2671:
2670:
2668:
2666:
2665:Invertebrates
2662:
2656:
2653:
2651:
2648:
2646:
2643:
2639:
2636:
2634:
2631:
2630:
2629:
2626:
2622:
2619:
2617:
2614:
2612:
2609:
2607:
2604:
2602:
2599:
2597:
2594:
2593:
2592:
2589:
2585:
2582:
2580:
2577:
2575:
2572:
2570:
2567:
2565:
2564:opportunistic
2562:
2560:
2557:
2555:
2552:
2550:
2547:
2545:
2542:
2540:
2537:
2535:
2532:
2531:
2530:
2527:
2523:
2522:penile spines
2520:
2518:
2515:
2513:
2510:
2508:
2505:
2503:
2500:
2499:
2498:
2497:Fertilisation
2495:
2491:
2488:
2486:
2485:Pelvic thrust
2483:
2481:
2478:
2476:
2473:
2472:
2471:
2468:
2464:
2461:
2459:
2456:
2454:
2453:mate guarding
2451:
2449:
2446:
2444:
2441:
2440:
2439:
2436:
2432:
2429:
2427:
2424:
2422:
2419:
2417:
2414:
2413:
2412:
2409:
2405:
2402:
2400:
2397:
2395:
2392:
2391:
2390:
2387:
2385:
2382:
2381:
2379:
2377:
2373:
2369:
2362:
2357:
2355:
2350:
2348:
2343:
2342:
2339:
2328:
2324:
2320:
2316:
2312:
2308:
2307:Contraception
2304:
2297:
2294:
2289:
2283:
2279:
2278:
2271:
2268:
2263:
2257:
2253:
2252:
2244:
2241:
2236:
2230:
2226:
2225:
2217:
2214:
2209:
2203:
2199:
2198:
2190:
2187:
2184:
2178:
2175:
2172:
2168:
2164:
2160:
2157:
2151:
2148:
2143:
2137:
2133:
2132:
2124:
2121:
2117:
2112:
2108:
2103:
2098:
2093:
2088:
2084:
2080:
2076:
2072:
2068:
2061:
2059:
2057:
2053:
2049:
2036:
2032:
2026:
2024:
2020:
2017:
2012:
2007:
2003:
1998:
1993:
1989:
1985:
1981:
1977:
1973:
1966:
1964:
1962:
1960:
1958:
1956:
1954:
1950:
1946:
1941:
1937:
1933:
1929:
1925:
1921:
1917:
1913:
1906:
1904:
1900:
1895:
1889:
1885:
1884:
1876:
1873:
1868:
1862:
1858:
1857:
1849:
1846:
1842:
1836:
1833:
1827:
1824:
1817:
1814:
1810:
1805:
1801:
1796:
1791:
1787:
1783:
1779:
1775:
1771:
1764:
1762:
1758:
1754:
1741:
1737:
1733:
1729:
1725:
1721:
1718:(9): 486–98.
1717:
1713:
1706:
1704:
1702:
1700:
1698:
1694:
1689:
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1478:
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1437:(2): 149–50.
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1018:
1016:
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1005:
1002:Male mammals
1000:
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992:
990:
986:
982:
978:
974:
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967:
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959:
953:
951:
947:
942:
940:
936:
932:
928:
924:
923:melanocortins
920:
916:
912:
908:
907:sexual desire
904:
900:
896:
893:motivation ("
892:
876:
867:
856:
847:
836:
827:
816:
807:
796:
787:
776:
767:
755:
746:
744:
742:
738:
734:
733:gametogenesis
730:
726:
722:
715:Gametogenesis
714:
709:Female rabbit
705:
700:
697:
690:
685:
682:
675:
670:
667:
666:spotted hyena
660:
655:
648:
643:
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628:
621:
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598:
592:
587:
584:
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571:
569:
565:
561:
557:
553:
549:
545:
544:plesiomorphic
539:
531:
529:
527:
523:
519:
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511:
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377:
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365:
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361:
357:
353:
349:
345:
341:
335:
333:
332:morphogenesis
329:
328:embryogenesis
325:
321:
320:fertilization
317:
312:
310:
306:
302:
298:
294:
290:
286:
282:
278:
266:
264:
262:
260:
256:
252:
248:
244:
240:
236:
232:
227:
223:
222:penile sheath
219:
215:
211:
207:
203:
195:
193:
188:
185:
181:
177:
173:
169:
161:
159:
157:
153:
152:bowhead whale
149:
145:
141:
137:
133:
129:
125:
121:
120:
119:Lemmus lemmus
115:
107:
105:
103:
99:
95:
91:
87:
86:estrous cycle
83:
79:
75:
71:
66:
64:
60:
55:
53:
49:
45:
41:
37:
33:
25:
21:
3127:olive baboon
3075:Hippopotamus
3063:domestic cat
3011:domestic dog
2960:
2939:golden eagle
2906:Crocodilians
2554:polygynandry
2475:cloacal kiss
2310:
2306:
2296:
2276:
2270:
2250:
2243:
2223:
2216:
2200:. Elsevier.
2196:
2189:
2177:
2150:
2130:
2123:
2114:
2074:
2070:
2046:
2039:. Retrieved
2009:
1979:
1975:
1943:
1915:
1911:
1882:
1875:
1859:. Elsevier.
1855:
1848:
1839:Dixson A.F.
1835:
1826:
1816:
1807:
1777:
1773:
1743:
1715:
1711:
1659:
1656:Reproduction
1655:
1645:
1635:
1616:
1606:
1599:
1564:
1560:
1550:
1507:
1503:
1493:
1482:ScienceDaily
1480:
1434:
1430:
1423:
1378:
1374:
1364:
1353:. Retrieved
1326:
1322:
1312:
1301:the original
1280:
1276:
1263:
1230:
1226:
1219:
1207:. Retrieved
1196:
1186:
1128:that detect
1124:function as
1112:
1001:
999:in mammals.
993:
970:
954:
943:
888:
718:
652:Male raccoon
612:domestic cat
541:
492:
473:
470:Human births
456:
396:
391:
360:estrus cycle
352:menstruation
336:
313:
274:
263:
241:through the
235:epididymides
199:
192:
155:
117:
111:
67:
56:
29:
2866:Salamanders
2746:Echinoderms
2729:sea anemone
2712:Cephalopods
2616:cannibalism
2539:promiscuity
2458:mating plug
2448:mating call
2443:mate choice
2041:15 February
1662:(1): 31–9.
1452:11336/66658
1283:: 568–586.
1209:16 February
570:placentas.
464:respiration
255:thermotaxis
239:ejaculation
218:dog's penis
144:Delphinidae
138:(including
78:Placentalia
74:Marsupialia
3174:Categories
2841:Amphibians
2821:Salmon run
2751:Gastropods
2673:Arthropods
2638:intralocus
2633:interlocus
2611:bimaturism
2470:Copulation
2463:lek mating
1355:2021-03-03
1178:References
1151:attractive
1140:pheromones
747:Copulation
681:marsupials
556:bandicoots
548:eutherians
532:Marsupials
502:New Guinea
494:Monotremes
483:Monotremes
356:great apes
291:, and the
259:chemotaxis
114:Cricetidae
40:platypuses
36:viviparous
3185:Fertility
3108:Primates
3097:Pinnipeds
3085:Marsupial
3016:gray wolf
2776:earthworm
2761:love dart
2734:jellyfish
2703:butterfly
2596:anisogamy
2584:synchrony
2574:cuckoldry
2544:polyandry
2394:evolution
2327:0010-7824
1679:1765/9599
1524:0962-8436
1345:1314-3301
1329:: 89–94.
1015:ungulates
1004:ejaculate
966:Hominidae
729:oogenesis
568:eutherian
566:found in
498:Australia
421:nutrients
405:nutrition
397:pregnancy
395:, called
393:Gestation
384:Gestation
366:Gestation
348:oogenesis
206:testicles
102:precocial
94:marsupium
90:altricial
82:gestation
3132:mandrill
3090:kangaroo
3033:Elephant
2944:seabirds
2876:Reptiles
2826:Seahorse
2724:Cnidaria
2688:scorpion
2621:coercion
2579:seasonal
2549:polygyny
2534:monogamy
2507:external
2502:internal
2159:Archived
2111:26684881
2071:PLOS ONE
2035:Archived
2006:25616211
1940:11135361
1932:19002141
1804:26109341
1740:13813765
1732:22926422
1688:11226027
1624:Archived
1542:25602069
1469:36538049
1461:12563318
1349:Archived
1297:18983263
1247:18185981
1203:Archived
1155:aversive
1136:odorants
1130:volatile
1032:See also
919:oxytocin
903:dopamine
597:Stallion
526:epipubic
441:hamsters
437:giraffes
429:vitamins
425:calories
409:placenta
324:implants
309:oviducts
301:clitoris
247:flagella
204:and the
136:primates
44:echidnas
42:and the
3149:Raccoon
3122:gorilla
3058:cheetah
3043:Felidae
3028:Dolphin
3021:red fox
2961:Mammals
2911:Tuatara
2889:lizards
2781:epitoky
2717:octopus
2376:General
2102:4684214
2079:Bibcode
1997:4508243
1982:: 1–7.
1795:4491543
1591:9720109
1582:1692310
1533:4305166
1383:Bibcode
1159:agonist
1025:ovulate
997:baculum
937:(i.e.,
899:satiety
895:wanting
721:meiosis
679:Female
639:raccoon
637:Female
522:puggles
415:via an
413:abdomen
401:mitosis
285:ovaries
210:scrotum
132:Bovidae
98:abdomen
32:mammals
3154:Rodent
3117:bonobo
3102:walrus
3001:coyote
2894:snakes
2831:Sharks
2772:Worms
2768:Sponge
2698:insect
2693:beetle
2683:spider
2601:oogamy
2325:
2284:
2258:
2231:
2204:
2138:
2109:
2099:
2004:
1994:
1938:
1930:
1890:
1863:
1802:
1792:
1738:
1730:
1686:
1589:
1579:
1540:
1530:
1522:
1467:
1459:
1413:
1406:430244
1403:
1343:
1295:
1255:812974
1253:
1245:
1013:, and
1011:canids
962:vagina
891:reward
696:rabbit
510:cloaca
460:infant
344:embryo
340:zygote
305:cervix
281:uterus
277:vagina
237:until
182:, and
148:whales
140:humans
70:Theria
3112:human
3070:Fossa
3053:tiger
3006:dingo
2991:Canid
2921:Birds
2854:frogs
2739:coral
2529:Modes
1936:S2CID
1736:S2CID
1465:S2CID
1304:(PDF)
1273:(PDF)
1251:S2CID
1167:TAAR5
1133:amine
1104:Notes
1007:semen
958:penis
694:Male
610:Male
518:joeys
514:birds
453:Birth
447:Birth
316:sperm
297:labia
293:vulva
202:penis
63:teats
30:Most
3048:lion
2816:Eels
2798:Fish
2678:crab
2323:ISSN
2282:ISBN
2256:ISBN
2229:ISBN
2202:ISBN
2136:ISBN
2107:PMID
2043:2014
2002:PMID
1928:PMID
1888:ISBN
1861:ISBN
1821:1994
1800:PMID
1728:PMID
1684:PMID
1587:PMID
1538:PMID
1520:ISSN
1457:PMID
1415:4797
1411:PMID
1341:ISSN
1293:PMID
1243:PMID
1211:2024
933:and
921:and
739:and
583:Bull
506:eggs
500:and
386:and
342:and
330:and
299:and
279:and
251:ovum
124:dogs
76:and
57:The
34:are
24:Goat
2974:rut
2315:doi
2167:doi
2097:PMC
2087:doi
1992:PMC
1984:doi
1920:doi
1790:PMC
1782:doi
1720:doi
1674:hdl
1664:doi
1660:121
1577:PMC
1569:doi
1565:353
1528:PMC
1512:doi
1508:370
1447:hdl
1439:doi
1401:PMC
1391:doi
1331:doi
1285:doi
1235:doi
1153:or
458:an
289:ova
3176::
2321:.
2309:.
2305:.
2113:.
2105:.
2095:.
2085:.
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2073:.
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2022:^
2008:.
2000:.
1990:.
1980:34
1978:.
1974:.
1952:^
1942:.
1934:.
1926:.
1914:.
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1325:.
1321:.
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1281:42
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1275:.
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1231:15
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2169::
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