Jacobson organ

from Wikipedia, the free encyclopedia
Sagital section 1: Paraseptal cartilage, (Cartilago paraseptalis); 2: Opening to Jacobson's organ, in which a probe was advanced ; 3: tuberculum septi nasi; 4: Nasopalatine duct (Stenson's duct); 5: The mouth of the sphenoid sinus (sinus sphenoidalis) in the recessus sphenoethmoidalis 6: Sinus frontalis .
Positional relationship of the vomeronasal organ (Jacobson) in a human embryo about 3 cm in size. Schematic representation of the nasal and oral cavities with a section in the frontal plane .
The Jacobson's organ is shown here as a two-sided system in the nasal septum adjacent to the septal cartilage ( cartilage septi nasi specified, blue); Below is a paraseptal strip of cartilage on both sides ( Cartilago vomeronasalis , also blue).

The Jacobson organ , Jacobson organ or vomeronasal organ ( Latin organum vomeronasale ) is an olfactory organ developed in many vertebrates , which, like the olfactory mucosa , is assigned to the olfactory system . The organ is named after the Danish surgeon Ludwig Levin Jacobson (1783–1843), who rediscovered the organ first described by Frederik Ruysch in snakes in 1809.

The sense of smell is not only important in order to suspect, locate, check and be able to find food sources, to smell sources of danger and to notice territorial markings, but also to distinguish between individuals, to recognize their affiliation and to help in the search for a sexual partner . In addition to the olfactory region, vertebrates usually have a vomeronasal organ (VNO) developed differently for this specific olfactory or pheromone perception in the nasal cavity .

Scheme of the Jacobson organ in a reptile.

morphology

In mammals, the vomeronasal or Jacobson's organ consists of tiny indentations (diameter between 0.2 and 2 millimeters ) on both sides of the nasal septum ( septum nasi ) approximately opposite the inferior turbinate ( concha nasalis inferior ) and lies below the nasal septum cartilage ( cartilage septi nasi ) as above the adjacent ploughshare ( vomer ).

A detailed examination of the histological ultrastructure of the Jacobson organ was reported by David Taylor Moran, Bruce W. Jafek and J. Carter Rowley (1991).

Oral cavity of a house cat ( Felis silvestris catus ). The access to the Jacobson organ ( ductus incisivus ) is visible as a small papilla ( papilla incisiva ) palatally behind the incisors .

These hose-like indentations are generally not reached by the air we breathe. On the side of their openings, in the contact area with Jacobson's cartilage (Paraseptal cartilage , Cartilago paraseptalis ), there is a cavernous vein network, occasionally with muscle cells, so that the liquid mucus on the nasal mucous membrane, including the dissolved molecules, is pressed out of the epithelial tubes or sucked into the lumen through the expansion or narrowing of vessels can be.

In lower fish, the organ is sensory innervated by its own nerve, the paired terminal nerve , which has already been closely connected to the olfactory nerve ( N. olfactorius ) in the disturbance . It seems to be an evolutionarily old system that serves to find a sexual partner or to fertilize the eggs ( roe ), evolved from the olfactory pits of fish.

Flehmen of a tapir ( Tapirus indicus )

In addition to the vomeronasal duct (VND), the Jacobson or vomeronasal organ (VNO) also includes associated glands and special blood vessels (in Feloidea a rete mirabile arteria maxillaris ) - which, as venae vomeronasales , can operate a pumping mechanism by swelling and decongesting - as well as a special one Nerve. The vomeronasal duct is only a millimeter-wide depression in the septal mucosa of the anterior third of the septum, equipped with receptor cells, the axons of which then combine to form the vomeronasalis nerve and thus pull to the olfactory bulb (accessory) of the olfactory brain .

In humans , the organ is said to arise in the fetal stage, but to regress before birth. A vomeronasal nerve can be detected in histological examinations up to the eighth embryonic month, after that no more. Evidence of a vomeronasal duct in adults varies, according to the literature, between 25 and 100 percent. This is attributed to the various conditions and techniques used to detect this anatomical structure: anterior rhinoscopy or endoscopic techniques, swelling of the nasal mucous membrane , misinterpretations and confusion with other types of morphological structures, such as the nasopalatine duct , but also the significantly different size of a duct between individuals vomeronasalis.

In humans, the cartilage strip Cartilago vomeronasalis is a remnant of the vomeronasal organ . It lies behind the anterior nasal spine , the anterior spinous process of the nose, which can be felt when the finger is moved upwards over the philtrum from the point of symmetry of the upper lip . The function of a Jacobson organ in humans is controversial in specialist circles.

In most mammals there is access from the nasal and oral cavities to the Jacobson organ via the incisive duct - a pair of oblique-vertical, rostroventrally oriented connecting ducts between the oral and nasal cavities . It is also known as the Stenson's passage (see Nils Stensen ) of the nasopalatine duct and is an existing passage in many vertebrates. It enables a connection between the hard palate (oral cavity) and the nasal cavity. This allows olfactory stimulants to reach the vomeronasal organ. This duct is generally obliterated in humans, but cases of a persistent nasopalatine duct have been described.

Fine construction and function

The vomeronasal organ (VNO) is a chemoreceptive organ that is embedded in a cartilage capsule and covered by an olfactory epithelium in which the endings of its sensory cells are located. The sensory cells of the vomeronasal organ are like those of the regio olfactoria at the same time nerve cells (primary sensory cells ) and form action potentials in response to excitations.

Schematic representation of the seven membrane-spanning parts of a G-protein-coupled receptor, similar to the described V1R and V2R receptors.

Vomeronasal neurons carry two different main types of receptors in their membrane, which - like the other olfactory receptors - belong to the group of seven-fold membrane -spanning (heptahelical) receptors that are coupled to G proteins ( G protein-coupled receptor ). However, the two families of vomeronasal organ receptors (VR) differ from the usual olfactory sensory neurons (OSN) both in terms of their molecular structure and in terms of their signal transduction components. While V2 receptors (V2R) are coupled to the G protein Gαo in many mammals , V1 receptors (V1R), which respond to pheromones , couple to the G protein Gαi2 . A vomeronasal receptor gene for a V1 receptor that is expressed in the olfactory epithelium could also be detected in the human genome.

In most mammals, the receptor cells of the vomeronasal organ conduct their signals to the accessory olfactory bulb . From there, there are connections to the medial parts of the amygdala , more precisely its corticomedial part (nucleus corticalis), and also to other core areas of the limbic system and the hypothalamus .

This accessory olfactory bulb is an additional, independent neuronal complex, usually posterior - dorsal (back and top) on the actual olfactory bulb . The accessory olfactory bulb has a laminar structure and consists of five layers of nerve cells: (1) the vomeronasal nerve cell layer, (2) the glomerular layer (GL), (3) the mitral cell layer, (4) the layer of the lateral olfactory tract (LOT) and ( 5) the granular cell layer .

meaning

Olfactory sensory cells are neurons that take up specific stimuli by means of special olfactory receptors and pass on excitation via their neurites to neurons in the olfactory bulb ( olfactory bulb or accessory olfactory bulb ). Their signals are then passed on to other regions of the brain, where further integration can take place. In contrast to the receptor cells of the olfactory sensory neurons in the olfactory epithelium of the olfactory region , those of the vomeronasal organ are not occupied by cilia . The olfactory receptors (OR) expressed in the olfactory epithelium by the first olfactory sensory neurons (OSN) also differ significantly from the two basic receptor types (V1R and V2R) of the vomeronasal organ in various vertebrates.

Ano-genital control between dogs (Canidae)
A male lion (Panthera leo) marks his territory

The olfactory receptors of the usual olfactory mucous membrane in the roof of the nose are primarily used to perceive odors ( odorants ) from the environment, which can, for example, indicate food sources, dangers (burning smell) or predators , with some odorous substances also being detected by the trigeminal nerve . The vomeronasal organ, on the other hand, is primarily responsible for those odor signals with which animals communicate internally. For example, an individual can use pheromones or other specific odorants to signal their reproductive situation or their hierarchical position to another individual of the same species .

In contrast to olfactory sensory neurons in the usual olfactory epithelium, the olfactory cells of the VNO hardly adapt; in other words, they retain their sensitivity to odorants even when exposed. Furthermore , the approximately 250 different subtypes of olfactory cells of the VNO do not each converge on a neuronal interface, but are distributed over about ten to thirty neuronal complexes of the olfactory glomeruli in the accessory olfactory bulb . A mitral cell can therefore be excited by several types of receptors.

In mammals, the origin of the pheromones lies mostly in the so-called apocrine sweat glands ( Glandulae sudoriferae apocrinae ). These scent glands are mainly found in certain areas of the skin - such as the armpits or armpit hair , the nipples ( glandulae areolares ), the perianal - and the genital region (such as copulins ) and less densely on the face, on the scalp and abdomen . In contrast to eccrine, apocrine sweat glands each open into a hair follicle . Both are innervated by the sympathetic nervous system, but not by the same messenger substances; the eccrine sweat glands have cholinergic receptors and the apocrine sweat glands adrenergic receptors.

The sensory cells of the Jacobson organ specialize in the olfactory perception of certain substances, in mammals especially on pheromones . Metabolites of sex hormones probably also act as pheromones, such as 16- androsterone or its alcoholic derivative androsterol or its ketone androstenone , which can also be produced by microbial reactions. Via such degradation products, sex hormones could in turn have an influence on different hypothalamic-pituitary-gonadic regulated processes via the surrounding medium.

Various studies suggest an interaction between the respective body odor , one's own MHC complex and genetic variations in the immune system ( genetic matching ) for the perception of odors in connection with the choice of partner .

Some animals have developed special movement patterns for particularly intense odor perception via the vomeronasal organ. In many mammals the flehmen can be observed. Tonguing is the stretching of the (ambivalent) tongue in snakes and many lizards , which thereby absorb odorous substances and present them to this organ in a different way.

The vertebrate pheromones can be divided into different classes depending on the effect they develop on the Jacobson organ, such as

  • Sex attractants,
  • Aggregation pheromones,
  • Dispersion pheromones,
  • Alarm pheromones,
  • Trace pheromones,
  • Territorial marker pheromones,
  • Rut-stimulating pheromones,
  • Crate identifiers.

Meaning using the example of the house mouse

In tests on laboratory mice ( Mus musculus ), the following characteristic reactions can be demonstrated with regard to the effects of pheromones :

  • Lee Boot Effect: The menstrual cycles of female mice that were kept in the absence of males are slowed down and finally canceled out completely.
  • Whitten effect: the menstrual cycles of the female mice are restarted and synchronized when the female animals are then kept with male mice.
  • Vandenbergh effect: Female animals show sexual maturity earlier if they live together with male animals.
  • Bruce Effect: If a pregnant mouse is kept with a mouse ready to mate, the pregnancy can be interrupted.

These interactions are caused by a number of volatile substances found in the body fluids of mice, particularly mouse urine. Male mice or rodents in general need a functioning vomeronasal organ (VNO) in order to bring efficient courtship behavior towards a female animal, but also to face competitors aggressively. In rodent females, in turn, the vomeronasal organ (VNO) promotes the rut and the effects described above in the presence of the male .

history

In 1703, Frederik Ruysch discovered the vomeronasalis duct on a wounded soldier while treating his facial injury . The Danish anatomist Ludwig Levin Jacobson first described this anatomical structure in 1811. Georges Cuvier also published a paper on this organ almost at the same time. It should be mentioned that Ludwig Levin Jacobson was a student of Cuvier. In 1877 Albert von Kölliker from Würzburg published a monograph entitled Ueber die Jacobsons'schen organs of humans . Von Kölliker was the first to try to prove the presence of the vomeronasal organ in humans, and he was able to provide this evidence histologically in human embryos as well as in adult individuals.

The term pheromone was introduced in 1950 for a secreted substance that causes a specific response via the olfactory system in another animal.

literature

  • M. Atzmüller, K. Grammer: Biology of the smell: The meaning of pheromones for behavior and reproduction. In: Speculum. 18th year, 1/2000 ( PDF; 492 kB ).
  • M. Atzmüller, K. Grammer (Hrsg.): Biology of the smell: The meaning of pheromones for behavior and reproduction. In: Speculum - Journal of Gynecology and Obstetrics. 2000, Volume 18, No. 1, (edition for Switzerland), pp. 12–18 ( full text as PDF file ).
  • Marcello Aspria: Sex Smells: Odor, Sexuality, and the Erotic Imaginary. On: ScentedPages.com January 15, 2009.
  • R. Besli, C. Saylam, A. Veral, B. Karl, C. Ozek: The existence of the vomeronasal organ in human beings. In: J Craniofac Surg. Volume 15, 2004, pp. 730-735.
  • Irving Bieber: Olfaction in Sexual Development and Adult Sexual Organization. In: American Journal of Psychotherapy. Volume 13, 1963, pp. 851-859.
  • Konrad Botzenhart, Hans E. Müller, Otfried Strubelt: Indoor air pollution. expert publishing house, 2001.
  • Maria de Fátima Pereira de Carvalho, Adriana Leal Alves, Mirna Duarte Barros: Study on the Morphology and Frequency of the Vomeronasal Organ in Humans. In: Int. J. Morphol. Volume 26, No. 2, 2008, pp. 283-288. ( PDF; 299 kB )
  • FJ Cole: Jacobson's Organ. In: Nature . Volume 168, September 29, 1951, pp. 529-530, doi: 10.1038 / 168529b0 .
  • Ludwig Levin Jacobson: Description of Anatomique d'un organs Observé dans les Mammifères. In: Ann. d. Mus. d'hist. nat. Paris. Volume 18, 1811, p. 412. The same: Anatomisk Beskrivelse over et nyt Organ i Huusdyrenes Næse. Veterinair-Selskabets scrifter. Volume 2, 1813, p. 209.
  • Gerhard Heldmaier, Gerhard Neuweiler: Comparative animal physiology: Neuro- and sensory physiology. Volume 1, Springer-Verlag, Berlin, ISBN 3-540-44283-9 , p. 121.
  • Willem J. Hillenius, Susan J. Rehorek: From the eye to the nose: Ancient orbital to vomeronasal communication in tetrapods? In: Chemical Signals in Vertebrates. Volume 10, 2005, pp. 228-241.
  • Kevin R. Kelliher, Scott R. Wersinger: Regulation of the Sexual Behavior and Reproductive Physiology of the Laboratory Mouse: Effects and Neural Mechanisms. ( Memento from September 15, 2012 in the Internet Archive ) In: ILAR Journal. (PDF; 808 kB)
  • M. Knecht, M. Witt, N. Abolmaali, KB Hüttenbrink, T. Hummel: The vomeronasal organ of humans. In: The neurologist. Volume 74, No. 10, 2003, pp. 858-862, doi: 10.1007 / s00115-003-1573-7 .
  • Albert von Kölliker: About Jacobson's organs in humans. In: F. von Rinecker: Festschrift. Wilhelm Engelmann. Leipzig 1877.
  • Friedrich Merkel: About the Jacobson organ of the adult and the papilla palatina. From the Anatomical Institute in Göttingen. Festschrift for the fiftieth medical anniversary of Privy Councilor A. von Kölliker. Bergmann, Wiesbaden 1892.
  • Desider v. Navratil: About the Jacobsonian organ of the vertebrates. In: Journal of Anatomy and History of Development . Volume 81, Issue 5-6, December 15, 1926, pp. 648-656.
  • Taymour Mostafa, Ghada El Khouly, Ashraf Hassan: Pheromones in sex and reproduction: Do they have a role in humans? In: Journal of Advanced Research. Volume 3, 2012, pp. 1-9, doi: 10.1016 / j.jare.2011.03.003 .
  • Christine Schaefer: About the JACOBSON organ (Organum vomeronasale) of the house guinea pig (Cavia aperea porcellus LINNE, 1758). Dissertation . Institute for Veterinary Anatomy of the Department of Veterinary Medicine at the Free University of Berlin, 1997. (PDF; 2.1 MB)
  • Sabine Schultz: Investigations on the vomeronasal organ of humans with electrophysiological and psychophysiological methods. Dissertation, Ruhr University, Bochum 2008, [2]
  • Tristram D. Wyatt: Pheromones and Animal Behavior: Communication by Smell and Taste. Cambridge University Press, Cambridge 2003, ISBN 0-521-48526-6 .
  • J. Verhaeghe, R. Gheysen, P. Enzlin: Pheromones and their effect on women's mood and sexuality. In: Facts, views & vision in ObGyn. Volume 5, number 3, 2013, pp. 189-195, PMID 24753944 , PMC 3987372 (free full text) (review).
  • E. Zuckerkandl : The Jacobson organ. In: Result. Anat. Development Volume 18, 1910, pp. 801-843.

Web links

Individual evidence

  1. ↑ Neurobiology. Chemoreception. Free University of Berlin ( Memento from July 14, 2014 in the Internet Archive ) (PDF; 12.0 MB)
  2. ↑ Roberto Tirindelli, Michele DiBattista, Simone Pifferi, Anna Menini: From Pheromones to Behavior. In: Physiol Rev . Volume 89, No. 3, July 2009, pp. 921-956. doi: 10.1152 / physrev.00037.2008 .
  3. ↑ Cornelia Menke: Morphology and topographical anatomy of the nose, the nasal cavity and the paranasal sinuses and associated structures in the European mouflon (Ovis gmelini musimon PALLAS 1811). Dissertation . University of Veterinary Medicine Hannover, 2003, p. 35. (PDF; 3.0 MB)
  4. ↑ David Taylor Moran, Bruce W Jafek, J Carter Rowley: The Ultrastructure of the Human Olfactory Mucosa. In: David G. Laing, Richard L. Doty, Winrich Breipohl (eds.): The Human Sense of Smell. Springer, Berlin / Heidelberg 1991, ISBN 3-642-76225-5 , pp. 3-28 III
  5. ^ Luis R. Saraiva, Sigrun I. Korsching: A novel olfactory receptor gene family in teleost fish. ( Memento of the original from October 23, 2014 in the Internet Archive ) Info: The archive link was automatically inserted and not yet checked. Please check the original and archive link according to the instructions and then remove this notice. In: Genome Res. 17, 2007, pp. 1448-1457. (PDF; 1.4 MB) @1@ 2Template: Webachiv / IABot / www.genetik.uni-koeln.de
  6. ↑ Katharina Simon: Investigations into the fine structure of the vomeronasal organ in pigs - an immunohistochemical and scanning electron microscope study. Inaugural dissertation . Ludwig-Maximilians-University Munich. VVB Laufersweiler Verlag, 2008. (PDF; 9.6 MB)
  7. ^ I. Salazar, P. Sánchez Quinteiro, JM Cifuentes, P. Fernández, M. Lombardero: Distribution of the arterial supply to the vomeronasal organ in the cat. In: The Anatomical Record . (Anat Rec.) Jan 1997, Vol. 247, No. 1, pp. 129-136, PMID 8986310 .
  8. ↑ a b M. Knecht, M. Witt, N. Abolmaali, KB Hüttenbrink, T. Hummel: Das vomeronasaleorgan des Menschen. In: The neurologist. Volume 74, No. 10, 2003, pp. 858-862, doi: 10.1007 / s00115-003-1573-7 .
  9. ↑ Thomas von Arx, Michael M. Bornstein: The open ductus nasopalatinus. A rare malformation and diagnostic trap. In: Switzerland Monthly Dental Medicine. Volume 119, 4/2009.
  10. ↑ KP Bhatnagar, TD Smith: The human vomeronasal organ. III. Postnatal development from infancy to the ninth decade. In: Journal of anatomy. Volume 199, Pt 3, September 2001, pp. 289-302, ISSN  0021-8782 , PMID 11554506 , PMC 1468331 (free full text).
  11. ↑ Thomas von Arx, Michael M. Bornstein: The open ductus nasopalatinus. A rare malformation and diagnostic trap. In: Switzerland Monthly Dental Medicine. Volume 1194, 2009, pp. 379-384 [1] .
  12. ↑ L. Buck, R. Axel: A novel multigene family may encode odorant receptors: A molecular basis for odor recognition. In: Cell. Volume 65, 1991, pp. 175-187.
  13. ↑ Rodrigo Suárez, Pedro Fernández-Aburto, Paul R. Manger, Jorge Mpodozis: Deterioration of the Gαo Vomeronasal Pathway in Sexually Dimorphic Mammals. In: plos one. October 19, 2011.
  14. ^ Metalife. Overview for receptor family V1R ( Memento of the original from June 26, 2014 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. (English)  @1@ 2Template: Webachiv / IABot / www.metalife.com
  15. ↑ The pheromone receptor family: Vomeronasal receptor genes. The analysis of mouse genomes is shown here. 137 receptors were found which can be divided into 12 families. I. Rodriguez, K. Del Punta, A. Rothman, T. Ishii, P. Mombaerts: Multiple new and isolated families within the mouse superfamily of V1r vomeronasal receptors. In: Nat. Neurosci. 5, 2002, pp. 134-140.
  16. ↑ Wendy E. Grus, Peng Shi, Ya-ping Zhang, Jianzhi Zhang: Dramatic variation of the vomeronasal pheromone receptor gene repertoire among five orders of placental and marsupial mammals. In: PNAS. Volume 102, No. 16, April 19, 2005, pp. 5767-5772. doi: 10.1073 / pnas.0501589102 ( PDF; 383 kB ).
  17. ↑ Erica Pantages, Catherine Dulac : A Novel Family of Candidate Pheromone Receptors in Mammals. In: Neuron. Volume 28, No. 3, December 1, 2000, pp. 835-845. doi: 10.1016 / S0896-6273 (00) 00157-4 ( PDF file ).
  18. ^ I. Rodriguez, CA Greer, MY Mok, P. Mombaerts: A putative pheromone receptor gene expressed in human olfactory mucosa. In: Nature Genetics. 26, 2000, pp. 18-19.
  19. ↑ Mahmood F. Bhutta: Sex and the nose: human pheromonal responses. ( Memento of the original from August 22, 2012 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. In: JR Soc Med. Volume 100, No. 6, June 2007, pp. 268-274, doi: 10.1258 / jrsm.100.6.268 . @1@ 2Template: Webachiv / IABot / jrsm.rsmjournals.com
  20. ^ DL Berliner, L. Monti-Bloch, C. Jennings-Ehite, V. Diaz-Sanches: The functionality of the human vomeronasal organ (VNO): evidence for steroid receptors. In: Journal of Steroid Biochemistry and Molecular Biology. Vol. 58, No. 3, June 1996, pp. 259-265, PMID 8836161 .
  21. ^ S. Takami, GD Fernandez, PP Graziadei: The morphology of GABA-immunoreactive neurons in the accessory olfactory bulb of rats. In: Brain Res. Volume 588, 1992, pp. 317-323.
  22. ↑ Illustration of a schematic neuronal circuit of the VNO in a mouse. From: Michael J. Baum, Kevin R. Kelliher: Complementary Roles of the Main and Accessory Olfactory Systems in Mammalian Mate Recognition. In: Annual Review of Physiology. Volume 71, 2009, pp. 141-160.
  23. ↑ Schematic representation of a parasagittal section through part of the head of a mouse. ( Memento of the original from December 8, 2015 in the Internet Archive ) Info: The archive link was automatically inserted and not yet checked. Please check the original and archive link according to the instructions and then remove this notice. Representation of the various olfactory receptors and their projections to the olfactory bulb (AOB; vomeronasal organ, VNO) Ignacio Salazar, Pablo Sánchez Quinteiro: The risk of extrapolation in neuroanatomy: the case of the mammalian vomeronasal system. In: Front. Neuroanate. October 30, 2009. doi: 10.3389 / neuro.05.022.2009 @1@ 2Template: Webachiv / IABot / c431376.r76.cf2.rackcdn.com
  24. ↑ DT Moran, BW Jafek, JC Rowley: Ultra Structure of the human olfactory mucosa. In: DG Laing, RL Doty, W. Breipohl (eds.): The Human Sense of Smell. Springer-Verlag, Berlin 1992, pp. 3-28.
  25. ↑ H. Matsunami, LB Bock: A multigene family encoding a diverse array pheromone receptors in mammals. In: Cell. 90 (4), pp. 775-784.
  26. ↑ Andreas Ziegler: Molecules of the MHC and olfactory receptors: possible significance in the context of reproduction. In: J. FERTIL. REPROD. 4/2003. Krause & Pachernegg Gablitz, pp. 14–18. (PDF; 1.1 MB)
  27. ^ E. Biasi, L. Silvotti, R. Tirindelli: Pheromone detection in rodents. In: Neuroreport. 12 (14), pp. A81-A84.
  28. ↑ also ovulation copulins, which are mixtures of volatile, short-chain fatty acids , which in the female vaginal cycle-dependent occur
  29. ↑ Hans-Rudolf Tinneberg, Michael Kirschbaum, F. Oehmke (eds.): Gießener Gynäkologische Furtherbildung 2003: 23rd further training course for doctors in gynecology and obstetrics. Springer-Verlag, Berlin / Heidelberg / New York 2013, ISBN 978-3-662-07492-3 , p. 151.
  30. ↑ Th. Boyd: On clitoris and preputial glands, especially in man and in some animals. In: Archives for Gynecology. 89, 1909, pp. 581-595. doi: 10.1007 / BF01929547 .
  31. ↑ Benninghoff: Macroscopic and microscopic anatomy of humans. Volume 3, Urban & Schwarzenberg, Munich / Vienna / Baltimore 1985, ISBN 3-541-00264-6 , p. 583 ff.
  32. ^ Martin Witt, Witold Woźniak: Structure and function of the vomeronasal organ. In: Advances in oto-rhino-laryngology. 2006, p. 63. doi: 10.1159 / 000093751
  33. ↑ C. Eizaguirre, TL Lenz, M. Kalbe, M. Milinski: Rapid and adaptive evolution of MHC genes under parasite selection in experimental vertebrate populations. In: Nature Communications . Volume 3, 2012, p. 621. doi: 10.1038 / ncomms1632
  34. ↑ Nicola von Lutterotti: Immune system determines choice of perfume, The fragrance of defense. on: faz.net , January 23, 2013.
  35. ↑ The smell of genes - what really matters when choosing a partner. In: Curious about science. Fall 2006, pp. 1-4. (PDF; 359 kB)
  36. ↑ A rhinoceros bull pleading
  37. ↑ AL Cerda-Molina, L. Hernández-López, CE de la O, R. Chavira-Ramírez, R. Mondragón-Ceballos: Changes in Men's Salivary Testosterone and Cortisol Levels, and in Sexual Desire after Smelling Female Axillary and Vulvar Scents. In: Frontiers in endocrinology. Volume 4, 2013, p. 159, doi: 10.3389 / fendo.2013.00159 . PMID 24194730 , PMC 3809382 (free full text).
  38. ↑ Michael Weidlitsch: Copulins and their effects on the testosterone balance of athletes. Diploma thesis University of Vienna, Vienna 2008, pp. 51–52.
  39. ↑ Mahmood F. Bhutta: Sex and the nose: human pheromonal responses. In: JR Soc Med. 100, 2007, pp. 268-274.
  40. ^ S. Van der Lee, LM Boot: Spontaneous pseudopregnancy in mice. In: Acta Physiol Pharm Neerland. 4, 1955, pp. 422-444.
  41. ↑ WK Whitten: Ocuurence of anestrus in mice caged in groups. In: J Endocrinol. 18, 1959, pp. 102-107.
  42. ↑ JG Vandenbergh, JM Whitsett, JR Lonabard: Partial isolation of a pheromone accelerating puberty in female mice. In: J Reprod Fertil. 43, 1975, pp. 515-523.
  43. ↑ HM Bruce: A block to pregnancy in the mouse caused by proximity of strange males. In: J Reprod Fertil. 1, 1960, pp. 96-103.
  44. ↑ C. Dulac , AT Torello: Molecular detection of pheromone signals in mammals: from genes to behavior. In: Nature Rev Neurosci. 14, 2003, pp. 551-562.
  45. ↑ Didier TROTIER, Kjell B. Døving: Anatomical Description of a New organ in the Nose of Domesticated Animals' by Ludvig Jacobson (1813). In: Chem. Senses. 23, 1998, pp. 743-754. (PDF; 3.2 MB)
  46. ↑ Georges Cuvier: Description anatomique d'un organ obsen / 6 dans les mammifieres. In: Ann. Mus. Hist. Nat. 18, 1811, pp. 412-424.
  47. ↑ P. Karlson, M. Luscher: Pheromones': a new term for a class of biologically active substances. In: Nature. 183, 1959, pp. 55-56.