εH Neuroanatomy

Neuroanatomy and Behaviour, 2021, 3(1), e17.

Behaviour
and ISSN: 2652-1768
doi: 10.35430/nab.2021.e17

DATA BRIEF

Orexin neuron activity in mating mice - a pilot study

Denis Burdakov1,2,3 and Mahesh M. Karnani4,5,*
1
Laboratory of Neurobehavioural Dynamics, Institute for Neuroscience, Department of Health Sciences and
Technology, ETH Zürich, Zürich, Switzerland
2
The Francis Crick Institute, London, UK
3
Institute of Psychiatry, Psychology & Neuroscience, King’s College London, UK
4
Neuroscience Center Zürich (ZNZ), ETH Zürich and University of Zürich, Zürich, Switzerland
5
Saints-Pères Paris Institute for the Neurosciences, Université de Paris, France
* mahesh.karnani@parisdescartes.fr

Abstract
Mating behaviours affect hypothalamic orexin/hypocretin neurons and vice versa. However, activity of orexin neurons has not
been recorded during mating before. We report an anecdotal dataset of freely-moving miniature microscope recordings of
orexin neuron activity during mating behaviours, as well as an oral sexual encounter previously undocumented in mice. Across
the orexin neuron population in the male, firing rates were maximally diverse during ejaculation, similarly diverse though
weaker during intromission, and inverse to this during anterior thrusting. In the female mouse, orexin neurons tended to
decrease firing during intromission after a transient increase. We provide this brief dataset for re-use, to enable further studies
of these rare behaviours with challenging surgical preparations.

Related Objects: Dataset - https://doi.org/10.5281/zenodo.4813116; Video - https://doi.org/10.5281/zenodo.4820456

Key words: Lateral hypothalamus; Mating; Orexin; Hypocretin; Calcium imaging; Miniscope; Freely moving recording; Mouse
fellatio

Introduction Animal mating behaviours can be diverse, including in some

species oral sexual encounters [12–15], which have not been doc-
The lateral hypothalamic area (LH) and its orexin/hypocretin umented in mice as far as we know. This lack of documentation
(orexin) neurons are primarily known as vital controllers of would imply that the repertoire of rodent mating behaviours is
arousal, feeding and metabolism [1,2]. However, early studies limited to a canonical cycle of chasing, anogenital investigation,
also showed that electrical stimulation of the LH rapidly and re- mounting and intromission [9,16,17]. One of the recordings here
versibly drives mating in the male rat [3]. In line with this, the contains a type of oral sexual encounter, similar to fellatio, which
orexin neurons integrate external and internal sensory informa- we refer to as anterior thrusting (AT). Due to the low frequency of
tion and output to most of the brain as well as the autonomic ner- these behaviours (mating was observed in 6/22 recordings in two
vous system, and affect sex hormone secretion [1,4]. C-fos assays, animals, and 0/13 recordings in three other animals) the findings
pharmacological experiments and cell-specific lesion data suggest are provided here as anecdotal evidence and data are provided for
that orexin neurons in male rats are activated during mating, and re-use. A subset of the presented recordings have been analyzed
orexin peptides and normal activity of orexin neurons facilitate in a different way in our previous paper [18].
mating initiation, performance and mating-induced place prefer-
ence in male rats [5–8]. However, orexin neuron activity has not
been recorded directly during mating, and therefore its roles in Materials and Methods
the appetitive and consummatory sequences of copulation are un-
known. The activity of orexin neurons during mating is of partic- Animals
ular interest as it has been suggested that a common side-effect
of selective serotonin reuptake inhibitors (SSRIs), inhibition of Animal handling and experimentation was approved by the UK
mating, is mediated by increased serotonergic inhibition of orexin government (Home Office) and by Institutional Animal Welfare
neurons [5,9–11]. Ethical Review Panel. The recorded male was 86, the female was

1
 2 | Neuroanatomy and Behaviour, 2021

Figure 1. A, Example whole cell patch clamp recording while imaging an orexin-GCaMP6s neuron in a brain slice. B, Fluorescence to firing rate relationship indicating
fluorescence recordings reliably report changes in orexin neuron firing rate. C, left, schematic of GCaMP6s delivery to orexin neurons and implantation of GRIN lens to
obtain optical access to the neurons. Right, schematic of recording paradigm, with the subject mouse and an opposite sex conspecific in an enclosure with a glass bottom
and a camera tracking behaviour from below. D, field of view of orexin neurons through the GRIN lens in the male mouse recorded in this figure, color coded same as F,G
and H. E, example frames from behaviour video showing intromission, AT (anterior thrusting) and ejaculation epochs. The diagram shows annotation of male mouse rear
paws and tail with a blue ‘V’ shape and female annotated from front paws to upper abdomen with a pink ‘V’ shape. F, Z-scored ΔF/F0 of the cells recorded during the social
interaction during the 10th recording day for this subject. G, mean fluorescence of each cell during the indicated epochs. ‘Pre’ epoch is from the first time point to when
the animals first meet, ‘post’ epoch is an equal time period after ejaculation. H, variability of the fluorescence signal expressed as the std during the same epochs as in G. I,
correlation coefficients of the population signals across behaviour categories, showing inverse correlation between AT and intromission, and a positive correlation between
intromission and ejaculation. J, mean coefficients of determination for each mating behaviour with the others. The color code for C, F, G and H are based on average firing
rate during intromission as that has the overall highest coefficient of determination to the other behaviours shown in J.

79 and their conspecifics were 35 (female in Fig 1), 46 (different housed in a controlled environment on a reversed 12h light-dark
female used as conspecific for some data in Fig 2E) and 134 (sterile cycle with food and water ad libitum. Conspecifics were co-housed
prm1 male in Fig 2A,B,D) days old at the beginning of the record- in groups of 2-5 same sex littermates.
ings. The recorded male had been in the recording arena with a
female conspecific for 5 sessions (30-60min each, not supervised)
prior to the first session where mating was recorded. It was there- Surgery
fore presumed sexually naïve at the first recording analysed here.
Prior to the second session where mating was recorded (also the Mice were stereotactically injected with AAV1-hORX.GCaMP6s
session with three AT epochs), the male was not entirely sexu- (100-150nl, 2.5×1012 GC/ml, U Penn vector core). Orexin promoter
ally naïve as it had been in 9 sessions with a female, out of which virus expression specificity has been characterized previously [18].
one had confirmed mating. The female conspecific in this record- After anesthesia with isoflurane, the scalp was infiltrated with li-
ing was however sexually naïve. The recorded female had been docaine and cut. Then a craniotomy was drilled at 0.9 mm lateral,
in 12 sessions with a male before the recording shown here. In 3 1.4 mm posterior from Bregma. A pulled borosilicate glass injec-
of the prior sessions the female mated with the same conspecific tion tip was used to inject virus at a depth of 5.4 mm at a rate of
as shown here. In between recordings, recorded mice were single 50 nl/min. The scalp was sutured or glued closed after removal of
 | 3

the injection tip. Animals received 5 mg/kg carprofen injections was placed close to the recorded cell. Then 10s current steps of
for two days as post-operative pain medication. varying size were injected intracellularly to elicit varying firing fre-
Two weeks after the virus injection, a custom-made alu- quencies. The 0.3 mm brain slices were bathed in continuously
minium head plate was attached to the skull with dental cement bubbled (95% O2 and 5% CO2 ) ACSF, containing (in mM): 126
(Metabond) and skull screws. A 0.8 mm diameter hole was drilled NaCl, 3 KCl, 2 MgSO4 , 2 CaCl2 , 1.1 NaH2 PO4 , 26 NaHCO3 , 0.1 pyru-
at the same position as in virus injection surgery and a 0.39 NA, vic acid, 0.5 L-glutamine, 0.4 ascorbic acid and 25 D-glucose. In-
7.3 mm long, 0.6 mm diameter cylindrical graded refractive in- tracellular solution contained (in mM): 130 K-gluconate, 5 NaCl,
dex (GRIN) lens (Grintech) was lowered slowly (150 μm/min) to 2 MgSO2 , 10 HEPES, 0.1 EGTA, 4 Mg-ATP, 0.4 Na-GTP, 2 pyruvic
a depth of 5.1 mm with a micromanipulator. The GRIN lens was acid, 0.1 Alexa-594, 0.1% biocytin, and 10 mM KOH (to set pH to
then cemented onto the skull. Thereafter the implant was coated 7.3).
with black dental cement and painted with several layers of black
nail polish to protect from light contamination.
Histology
Animals received one dose of 0.6mg/kg dexamethasone as anti-
inflammatory medication along with 5mg/kg carprofen injections
Following standard perfusions as described in [18], 0.1 mm sec-
for post-operative pain management for two days. After a mini-
tions were cut on a vibratome. Sections were blocked in PBS
mum of two weeks of recovery, mice were trained for freely mov-
with 0.3% Triton X-100 and 1% bovine serum albumin (blocking
ing miniature endoscope recordings under an incremental regi-
solution) for 1 h, incubated with goat anti-orexin (1:1000; Cat#
men of wearing the head-mounted miniature microscope (~2g
8072, RRID: AB_653601, Santa Cruz) over-night, washed, incu-
weight) in the test arena (Figure 1C).
bated with Alexa 647 conjugated donkey anti-goat (1:1000; Cat# A-
21447, RRID: AB_2535864, Invitrogen/Thermo Fisher Scientific)
Freely moving Ca2+ imaging for 3.5 h, washed and mounted. Antibodies were applied in block-
ing solution. Confocal micrographs were acquired on a Nikon A1
Ca2+ imaging was performed with a miniaturized head-mounted and merged in imageJ.
microscope (Inscopix) which was attached to the animal’s head,
without anaesthesia shortly before recording. The animal was
Statistics
then placed into the arena with a plexiglass floor. The animal
had already spent time in the same arena leaving scent markings. All data are shown as mean ± std unless stated otherwise.
Recording was begun after approximately 10 minutes of initial ex-
ploration of the familiar arena. Each recording consisted of obser-
vation of the subject and conspecific with food and water available Results
ad libitum. Sexual receptivity was not induced with hormones,
females were not ovariectomized and estrous was not confirmed Orexin neuron activity during intromission and AT in the
with vaginal cytology. The arena was lit with red LED lighting and
male mouse, and during intromission in the female
was covered from view in a dark, quiet room with a silent fume ex-
tractor pipe above the arena to eliminate background odors. Ca2+ We recorded orexin neuron firing rates using the GCaMP6s Ca2+
imaging frame rate was 10 frames/s. Behaviour was tracked with sensor [20] which was delivered in a viral vector under the orexin
a CCD camera (Lumenera Infinity) from below the arena at 65 promoter yielding GCaMP6s expression with 97.4 ± 1.0 % speci-
frames/s. An LED facing the behaviour camera and blinking in re- ficity and 65.8 ± 3.7 % penetrance in orexin neurons [18]. Simulta-
sponse to each Ca2+ imaging frame was used to synchronize Ca2+ neous electrical and optical recordings in brain slices showed that
imaging and behaviour videos. Behaviour analysis was performed GCaMP6s fluorescence faithfully reported spiking activity (Figure
in ImageJ by manually annotating behaviours. The full datasets 1A,B). Using implanted graded refractive index (GRIN, Figure 1C)
are available for free re-use under a public domain dedication [19]. lenses and a miniature fluorescence microscope mounted on the
skull, we recorded activity of orexin neuron populations in freely
behaving mice as they navigated a familiar 24x24cm arena that
Ca2+ imaging analysis
contained food pellets, water and an opposite sex conspecific (Fig-
ure 1C-F). Some of these datasets were analyzed in less detail in
Ca2+ imaging data were preprocessed for dropped frames and pix-
Figure 1 of our previous article [18], and here we report in more de-
els, downsampled by a factor of three spatially, then motion cor-
tail on the mating behaviours observed in a limited subset of those,
rected in Mosaic (Inscopix) and saved as 16bit TIFF stacks. Further
and previously undocumented recordings.
processing was performed in Matlab with custom routines. Re-
gions of interest (ROIs) were drawn manually around cells and av- The firing rate changes in male orexin neurons were large
erage fluorescence was extracted across all pixels within each ROI during mating behaviours, reaching, in different cells, peak and
as well as a neuropil ‘halo’ around each ROI consisting of the third through values during ejaculation corresponding to the extremes
to the sixth nearest pixels outside the outline that did not contain of their 0 to 30 Hz firing range (Figure 1B,F,G). The population fir-
other ROIs. These signals were lowpass filtered with a moving av- ing pattern of orexin neurons during AT was inverted compared to
erage of three frames. Neural signals were calculated by subtract- that during intromission (Figure 1G), and firing was less variable
ing the neuropil ‘halo’ signal from each ROI specific signal. This during AT (Figure 1H) than other mating behaviours. Correlation
signal was then corrected for photobleaching by computing ΔF/F0 coefficients of the population firing rate vector across behaviours
using the mean over a 120 s moving window as F0 and z-scored. revealed that intromission and ejaculation were highly correlated
and both were inversely correlated with AT (Figure 1I). Intromis-
sion had the strongest average of coefficients of determination
Patch-clamp recordings (R2 ) with other mating behaviours (Figure 1J), and was therefore
used to sort neurons in plots and color code.
Correspondence between firing rate and GCaMP6 fluorescence AT was observed in the last one out of ten recordings in one
was recorded in whole cell mode as described in [18]. Briefly, acute male over an 86 day period. At this point the male was not sexually
slices were cut from animals injected with AAV1-hORX.GCaMP6s naïve, though the female was (see Methods). This behaviour con-
as above. A whole-cell recording was established and the same sisted of the male approaching the female’s head from the front or
miniature microscope and GRIN lens as in the in vivo recordings, side, rearing, using front paws to push down the female’s head fol-
 4 | Neuroanatomy and Behaviour, 2021

Figure 2. A, Z-scored ΔF/F0 of the cells recorded during the social interaction. B, field of view in the male mouse recorded in this A and C. C, mean fluorescence of each
cell during the indicated epochs in the female recorded in A and B (above plot) and in the male recorded in Figure 1. D, event triggered averages of cell fluorescence across
repetitions of the indicated behavioural events in the female. Mean across cells shown below the cell rasters. E, same as D but from the male recorded in Fig1, including also
anterior thrusting, ejaculation, tumescence and detumescence of the penis. F, GRIN lens placements (* indicates the cylindrical lesion left by the GRIN lens) in the lateral
hypothalamus in the female (above) and male (middle) along with an atlas schematic of the coronal plane (bottom).
 | 5

lowed by erection and multiple rapid thrusts of the penis into the cence (which coincided with thrusting) the male orexin neurons
female’s mouth (Movie 1, Figure 1E). AT occurred in three bouts tended to decrease activity, suggesting that overall orexin neuron
each lasting 2.4, 4.6 and 1.1 s and containing an estimated 21, 13 and activity specifically increases locomotion [18] rather than other ac-
15 thrusts toward the mouth and 5, 4 and 9 estimated insertions tion patterns. Additionally orexin neurons may have a role in or-
of the penis into the female’s mouth. The bouts ended with the fe- chestrating autonomic nervous system outflow and CNS reward
male rearing away or turning to the side, penile detumescence and processing during ejaculation, as, during ejaculation, they express
either the male or female moving away while the other remained even more strongly their intromission firing rates (Figure 1F,G).
in place, apart from the third bout which ended in the male chasing This is in line with previous work showing decreased conditioned
the female and intromission. The multiple thrusts into the mouth place preference for mating behaviour in male mice upon orexin
and the coordinated end without signs of aggression, stress or es- neuron ablation [8]. AT lasted 2.7 ± 1.8 s while intromission lasted
cape suggest that the behaviour was not aversive. The bouts oc- 35.6 ± 22.9 s (p=0.048 with Student’s t-test), suggesting an early
curred during a 64.5 s interval between two bouts of intromission. switch to other behaviour patterns from AT. The inversion of the
The delay from stop of first intromission to start of first AT was 22.5 male intromission firing pattern during AT (Figure 1G,I,2E) could
s. The intervals between AT bouts were 11.9 and 11.4 s and intromis- be part of a mechanism for switching of action plans. As orexin
sion started 10.9 s after the third AT bout. Intromission before AT neurons promote wakefulness, their activity during mating may
lasted 41.6 s, whereas after the AT intromission lead to ejaculation critically support arousal while the CNS is engaged in coordinating
in 31.5 s, which was evident as spasming followed by falling to the autonomic nervous system outflow. This is consistent with occa-
side and prolonged immobility (Figure 1E). In another recording sional reports of loss of consciousness during sex - orgasmolepsy
29 days prior, intromission also occurred but bouts were shorter - in narcoleptic patients, who typically lack orexin signaling [22].
(29.7 ± 26.8 s, n=4). In this recording there was one apparent at-
tempted AT interrupted by the female rearing away. Total cumu-
lative time of intromission before ejaculation was 73.1 s (across 2 Declarations
bouts) with AT and 101.9 s (across 4 bouts) without AT. These data
would suggest that mouse AT shortens mating duration by hasten- Acknowledgements
ing ejaculation.
We also captured mating in a female mouse during orexin neu- We thank Dr Edward Bracey for useful comments.
ron imaging (Figure 2A-D). These recordings did not contain AT
bouts. As a contrast to the male firing patterns, female orexin neu-
rons were predominantly inhibited during intromission after an Funding
initial peak in their activity (Figure 2C-E). We additionally aligned
the male neural activity to erections. On average, orexin neuron This project has received funding from the European Union’s Hori-
activity started to decrease upon onset of tumescence of the penis, zon 2020 research and innovation programme under the Marie
and onset of detumescence coincided with the through of orexin Skłodowska-Curie grant agreement DRIVOME (grant agreement
neuron activity (Figure 2E). These data suggest that overall, cop- 701986). The experiments in this study were supported by the
ulation coincides with a decrease of orexin neuron firing in both Francis Crick Institute which receives its core funding from Can-
sexes. However, both sexes had heterogeneity within the neural cer Research UK (FC001055), the UK Medical Research Council
population, and in particular the male had several orexin neurons (FC001055), and the Wellcome Trust (FC001055).
that increased activity during tumescence, intromission and ejac-
ulation (Figure 2E). GRIN lens placement was confirmed to be in
Conflict of Interest Declaration
the lateral hypothalamus (Figure 2F).

The authors declare no conflicts of interest.

Discussion
Editorial Notes
This pilot study is an anecdotal report as it consists of two record-
ings in a male and a female mouse. The data however indicate
History
that orexin neuron activity is strongly and heterogeneously mod-
ulated during mating behaviours, and suggest that mouse mat-
• Received: 2020-11-19
ing behaviours can exhibit more diversity than documented previ-
• Revisions Requested: 2020-12-14
ously. Though AT seemed non-aversive to the female, it is possi-
• Revised: 2021-05-26
ble that the size of the arena limited the options to retreat from this
• Accepted: 2021-05-26
behaviour, as female mice pace mating behaviours when given the
• Published: 2021-06-02
opportunity to retreat [21]. We should note that the head mounted
miniature microscope is ~3 cm high and thus may affect mating
behaviours, such as AT when the female is recorded. Another po- Editorial Checks
tential caveat is that the GRIN lens implantation entails lesioning
brain tissue in somatosensory neocortex, dorsal hippocampus and • Plagiarism: Plagiarism detection software found no evidence
thalamus in one hemisphere, as these structures stand in the path of plagiarism.
of the lens. • References: Zotero did not identify any references in the Re-
These recordings suggest that sub-second firing rate changes tractionWatch database.
in orexin neurons have a role in mating behaviours. What could
this role be? We recently reported that orexin neurons are involved
in locomotion and sensorimotor processing [18]. The increased Peer Review
orexin neuron firing during chasing (Figure 2D,E) is likely a part
of that. Orexin neurons may have a role in supporting the motor This paper followed a standard single-blind review process.
outflow underlying intromission as the female who typically re- For the benefit of readers, reviewers are asked to write a public
mains stationary has overall more inhibited orexin neurons than summary of their review to highlight the key strengths and weak-
the male, who is thrusting (Figure 2C-E). During penile tumes- nesses of the paper. Signing of reviews is optional.
 6 | Neuroanatomy and Behaviour, 2021

Reviewer 1 (Anonymous) Reproduction. Amsterdam: Academic Press; 2015. doi:

10.1016/C2011-1-07288-0.
In this pilot study the authors provide preliminary two-photon cal- 10. Lorrain DS, Matuszewich L, Friedman RD, Hull EM. Extracel-
cium recordings from orexin neurons in the lateral hypothalamus lular serotonin in the lateral hypothalamic area is increased
in freely behaving mice (one male and one female mouse) during during the postejaculatory interval and impairs copulation in
copulation and non-copulatory behaviours. male rats. The Journal of Neuroscience. 1997;17(23):9361–
There are several commendable aspects to this pilot study: 9366. doi: 10.1523/jneurosci.17-23-09361.1997.
1) In-vivo recordings from orexin neurons in mice during cop- 11. Li Y, Gao XB, Sakurai T, van den Pol AN. Hypocretin/orexin ex-
ulation has not yet been reported. cites hypocretin neurons via a local glutamate neuron—A po-
2) The inclusion of both male and female animals, and a com- tential mechanism for orchestrating the hypothalamic arousal
parison of neural activity during the phases of copulation. system. Neuron. 2002;36(6):1169–1181. doi: 10.1016/s0896-
The authors demonstrate a wide variation in orexin neuron ac- 6273(02)01132-7.
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tantly, show preliminary evidence of increased inhibition during latio by fruit bats prolongs copulation time. PLoS ONE.
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male Bonin Flying Foxes Pteropus pselaphon. PLOS ONE.
3) The authors report the recordings for each neuron in the vi-
2016;11(11):e0166024. doi: 10.1371/journal.pone.0166024.
sual plane of the GRIN lens.
14. Maruthupandian J, Marimuthu G. Cunnilingus apparently in-
4) These pilot data are interesting, and hopefully motivate the
creases duration of copulation in the Indian Flying Fox, Ptero-
authors to pursue this line of inquiry with an adequately powered
pus giganteus. PLoS ONE. 2013;8(3):e59743. doi: 10.1371/jour-
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nal.pone.0059743.
The authors report a previously unreported behaviour resem-
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bling fellatio in mice. However, the testing conditions and sex-
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ual experience status of the mice makes it difficult to determine
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whether this behaviour is truly part of a male mouse’s pattern of
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168. doi: 10.1016/j.conb.2019.12.001.
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