TYPE Review

PUBLISHED 24 January 2024

DOI 10.3389/fnbeh.2023.1330460

Neurophysiology of male sexual

OPEN ACCESS arousal—Behavioral perspective
EDITED BY
Fabrizio Sanna,
University of Cagliari, Italy
Wiktor Bogacki-Rychlik, Katarzyna Gaw˛
eda and Michal Bialy*
REVIEWED BY Department of Experimental and Clinical Physiology, Laboratory of Centre for Preclinical Research,
Gabriela Rodriguez-Manzo, Medical University of Warsaw, Warsaw, Poland
National Polytechnic Institute of Mexico
(CINVESTAV), Mexico
Pekka Santtila,
New York University Shanghai, China
Wendy Portillo, In the presented review, we analyzed the physiology of male sexual arousal
National Autonomous University of Mexico, and its relation to the motivational aspects of this behavior. We highlighted
Mexico
the distinction between these processes based on observable physiological
*CORRESPONDENCE
Michal Bialy and behavioral parameters. Thus, we proposed the experimentally applicable
[Link]@[Link]; differentiation between sexual arousal (SA) and sexual motivation (SM). We
michalbialy@[Link]
propose to define sexual arousal as an overall autonomic nervous system
RECEIVED 30 October 2023 response leading to penile erection, triggered selectively by specific sexual
ACCEPTED 15 December 2023 cues. These autonomic processes include both spinal and supraspinal neuronal
PUBLISHED 24 January 2024
networks, activated by sensory pathways including information from sexual
CITATION
Bogacki-Rychlik W, Gaw˛ eda K and Bialy M partner and sexual context, as well as external and internal genital organs.
(2024) Neurophysiology of male sexual To avoid misinterpretation of experimental data, we also propose to precise
arousal—Behavioral perspective.
Front. Behav. Neurosci. 17:1330460.
the term “sexual motivation” as all actions performed by the individual that
doi: 10.3389/fnbeh.2023.1330460 increase the probability of sexual interactions or increase the probability of
COPYRIGHT exposition to sexual context cues. Neuronal structures such as the amygdala,
© 2024 Bogacki-Rychlik, Gaw˛ eda and Bialy. bed nucleus of stria terminalis, hypothalamus, nucleus raphe, periaqueductal
This is an open-access article distributed
under the terms of the Creative Commons gray, and nucleus paragigantocellularis play crucial roles in controlling the level
Attribution License (CC BY). The use, of arousal and regulating peripheral responses via specific autonomic effectors.
distribution or reproduction in other forums
is permitted, provided the original author(s) On the highest level of CNS, the activity of cortical structures involved in the
and the copyright owner(s) are credited and regulation of the autonomic nervous system, such as the insula and anterior
that the original publication in this journal is
cited, in accordance with accepted academic
cingulate cortex, can visualize an elevated level of SA in both animal and human
practice. No use, distribution or reproduction brains. From a preclinical perspective, we underlie the usefulness of the non-
is permitted which does not comply with
contact erection test (NCE) procedure in understanding factors influencing
these terms.
sexual arousal, including studies of sexual preference in animal models. Taken
together results obtained by different methods, we wanted to focus attention on
neurophysiological aspects that are distinctly related to sexual arousal and can
be used as an objective parameter, leading to higher translational transparency
between basic, preclinical, and clinical studies.

KEYWORDS

sexual arousal, sexual behavior, sexual motivation, sexual preference, male

Introduction
Understanding the mechanisms that control particular types of behavior
requires appropriate models. In the case of sexual behavior, the evolutionary
conservative character of regulatory loops qualifies animal models as
homologically reliable (Pfaus et al., 2003). This phylogenetic continuance
is visible in similar across-species hormonal regulation (mainly steroid sex
hormones) and neuroanatomical correspondence. To this day, the most studied

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animal models of sexual behavior are rats (Rattus norvegicus), and During sexual activity, emotions are also expressed. As always,
thus, in this paper, if there is no additional mention of other species, subjective and objective parts of emotional responsiveness are
the presented data refers to rats. Above its translational value, basic challenging during their separation. Previously, it has been
and preclinical studies of rat sexual behavior give insight into many described that detailed ultrasonic vocalization analysis could serve
aspects of general neurobiology, physiology, and pharmacology by as an additional parameter depicting these components (Barfield
providing reliable and repeatable parameters. et al., 1979; Bogacki-Rychlik et al., 2021, 2022).
Despite its conservative and highly innate profile, neurons To express the sexual behavior repertoire, there is a need
involved in sexual behavior are subject to neuronal plasticity to activate vast regions of the CNS. Thus, the major condition
processes (Hull and Dominguez, 2007). These changes are related that enables all other specific reactions is the state of so-called
to perinatal critical periods during brain development (Gorski, general arousal (GA). From a behavioral perspective, if an animal
1985; Baum, 2009). In adult animals, plasticity processes are is awake, responsive, and presents proper locomotor (exploratory)
visible during the acquiring of sexual experience (Bialy et al., activity, we can judge that the level of GA is adequate (similarly to
1992, 2000; Bialy and Kaczmarek, 1996; Dominguez et al., medical examination).
2006; Hull, 2011; Sanna et al., 2017, 2020) adjusting sexual The knowledge accumulated during many years of
performance to environmental conditions. Typically, during the investigation of animal sexual behavior may also be useful
repeated acquisition of experience and condition to the rewarding in preclinical research, as many aspects of physiology and
value of copulation, the sexual interactions become more effective pathophysiology related to age, cardiovascular disease, metabolic
(shorter intromission and ejaculation latencies) (Larsson, 1959; disease, depression, and others affect sexual activity (Bialy et al.,
Dewsbury, 1969). 2019). The study of rat sexual behavior finally matters for
Sexual interaction, especially copulation to ejaculation, understanding human sexuality and behavior (Bancroft, 2008).
activates the reward system. From highly euphoric to merely Despite the general limitation of preclinical transferability, sexual
responsive, the male climax with the ejaculation reflex is one behavior has some more specific restrictions (Le Moëne and
of the most (if not the most) natural-rewarding states. By using Ågmo, 2019), and thus, direct translation of experimental data is
conditioned place preference tests (CPP), there is a possibility to challenging.
depict the rewarding state of particular phases of sexual activity Furthermore, this problem is enhanced by the non-coherent
(Tenk et al., 2009). These rewarding properties can also be used to terminology used by field experts, which implicates a broader
instrumentalize reaction and, by that, measure the motivational misunderstanding when the application of models is trying.
aspect more than the level of sexual arousal (SA) (Whalen, 1961; In this review, we focused on the definition of sexual arousal,
Hård and Larsson, 1968). Inversely, conditioning processes can methods to investigate this state, and its relation to sexual
also inhibit sexual interactions when punishment processes are motivation and GA.
applied (Beach et al., 1956). We propose to define male sexual arousal as a set of autonomic
Regulation of mating behavior is strictly related to hormonal nervous system (ANS) reactions triggered by specific sensory
activity, with the HPA-axis and gonadal sex steroids as the center of inputs, analyzed on different levels of CNS, leading to the
the regulatory mechanism (Awoniyi et al., 1993; Babaei-Balderlou occurrence of penile erection and potentially ejaculation. This
and Khazali, 2016). Sexual behavior has opened the possibility definition emphasizes the importance of physiological reflexes
of identifying sexual-related neuronal loops by testing their rather than subjective (introspective) psychological reactions. It
responsiveness to steroids (see for review: Sachs and Meisel, 1988). also gives the conceptual scaffold to analyze arousal by detecting
As it turned out, neurons containing receptors for sex hormones autonomic parameters and separating them from less specific
themselves do not transfer crucial information for activation of motivational components.
copulation (for instance, odor cues) but instead participate in local We hope that this terminological clarification, together with
loops that modulate and amplify such information (Wood and
reminders of fundamental experiments, will be helpful during
Newman, 1995; Maras and Petrulis, 2010; Petrulis, 2013; Gresham
experiments with sexual behavior.
et al., 2016; Nakamura et al., 2022).
Activation of sexual responses requests specific cues called
attractants. The reception of sexual signals, transduction to CNS,
and consequential decoding and integration of information are Male sexual arousal—Behavioral
maintained by various sensory channels. The main olfactory and perspective
vomeronasal pathways (Mier Quesada et al., 2023) are dominants
in rats (and other rodents), with accessory information conveyed by The first studies of sexual behavior started in the XIX century,
visual, acoustic, and touch-related fibers. Such sensory stimulation focusing on amphibians, mostly frogs (see Aronson and Noble,
gives a chance to tag crucial for sexual arousability neuronal nuclei, 1945). The amphibian spinal cord has far-reaching autonomy
and stimulate them artificially (for instance: optogenetic methods) from the higher levels of the central nervous system and can
(Kunkhyen et al., 2017). independently regulate animal activity. One of the elements of male
As a result of the activation of the sensory system, the effectoric copulation–the clasp response can be clearly visible in a frog, which
activation takes place. The autonomic nervous system prepares has a preserved spinal cord only.
peripheral tissue to initiate and consequently maintain copulation Tarchanoff (1887) found that clasp response is initiated by
patterns until ejaculation. In the case of male sexual behavior, stimulation of skin receptors, and its strength is proportional to
the most evident peripheral indicator of sexual arousal state is an the filling level of seminal vesicles (SVs). When a male, with
occurrence of penile erection. spared only a spinal cord had filled on seminal vesicles, clasp

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response was strong. Even after the removal of the heart, liver,
kidneys, or testis, this reflex remained to be properly expressed and
disappeared shortly before death. In more physiological situations,
clasp response disappeared only when SVs were empty and could
be restored by filling seminal vesicles with milk (Tarchanoff,
1887). These results suggest that internal sensory information
from seminal vesicles passed via the autonomic nervous system
is essential to maintain copulation, at least at the level of the
spinal cord, and can be the element of sexual arousal. However,
amphibians copulation differs compared to mammals activity in
many aspects. For example, this group of animals has distinct
genital anatomy adapted to external insemination. Their central
nervous system is composed of relatively few neurons that mostly
occupy ventricular and periventricular regions with poorly visible
neuronal clusters (nuclei) without developed neocortex. Hence,
there are no close homologs with mammalian brain nuclei
(Wilczynski, 2009). Also, the amphibian ANS is poorer developed
in comparison with the mammalian system (Biaggioni et al., 2022).
Nevertheless, even in this group, different aspects of sexual activity
are controlled by various regions of the central nervous system,
creating a relatively complex system (Aronson and Noble, 1945).
In mammals, a more comprehensive analysis of male sexual
activity started from Frank Beach’s research on rat sexual behavior. FIGURE 1
Three primary processes regulating sexual behavior. Although all
Beach (1942, 1956) described that initiation of copulation did not three processes are defined in the text as separate mechanisms,
correlate with other behavioral parameters describing copulatory they are interconnected and affect one another. Sensory cues from
performance. He has proposed to distinguish two independent genital organs, referred to as internal cues, primarily affect sexual
arousal (SA). External cues related to the sexual context have an
mechanisms: the sexual arousal mechanism (SAM) and the impact on all SM, SA, and GA. GA is additionally activated by
intromissive-ejaculatory mechanism (IEM). SAM initiates sexual non-specific cues which induce arousal and wakefulness. SA, sexual
interaction and can be measured by mount latency (time from the arousal; GA, general arousal; SM, sexual motivation.

introduction of receptive female to first attempt to copulation—

mounting). Consequently, IEM regulates copulation and can be
measured by ejaculation latency (time from first intromission - In contrast, touch-based (ex-copula) erection depends on
to ejaculation). Future factor analysis of all sexual parameters androgens - dihydrotestosterone or testosterone treatment
distinguished five independent groups of behavioral parameters: (Meisel et al., 1984).
(1) anticipatory behavior (measured by level changes in the - Also, erections during non-contact tests after castration
bi-level cage), (2) initiation of copulation, (3) copulatory efficiency depend on androgens (dihydrotestosterone or testosterone)
(measured by ejaculation latency), (4) a number of intromissions given peripherally (Manzo et al., 1999).
required to achieve ejaculation, and (5) intromission (hit) ratio
(number of intromissions to all attempt copulation). Thus, every We can see that different observable aspects of male
group reflects the activity of different neuronal regulatory loops sexual behavior have even more complex inner regulatory
(Sachs, 1978; Dewsbury, 1979; Pfaus et al., 1990). composition. This complexity emerges from the parallel and
It’s important to notice that these statistically defined subsequent processing of different neuronal modules. Some of
independent factors (groups) do not enable us to fully separate them are specific, and others are not for sexual reactions.
SM from SA (Bialy et al., 2019). However, it is now concluded During sexual behavior experiments, we can thus observe that
that initiation of copulation measured by mount latency describes particular behavioral components are more associated with some
sexual motivation rather than sexual arousal (Hull et al., 2002). combination (Figure 1) of GA, SA, and SM (Bialy et al., 2019).
Sachs (2000) pointed out another important factor indicating Distinctions between sexual arousal and motivation (Sachs,
the complexity of male sexual behavior: the diversity of erection 2000, 2007) and between general arousal and sexual arousal
mechanisms. This diversity was visible not only by changes in (Bancroft, 1999; Pfaff et al., 2002; Heiman and Pfaff, 2011) were
behavioral contexts but also by various reactions to hormonal generally accepted during discussions on sexual arousal in the
stimulation required for erection restoration after castration. To be Hormones and Behavior journal [2011, vol. 59(5)].
precise: What do we exactly mean by the mention of male sexual
arousal? In the literature, we can come across many terms that refer
- The occurrence of erection obligated to restore initiation to sexual arousal. Especially when describing human sexuality, the
of copulation (intromission latency; the time between terminology differs significantly from that used in animal studies.
the introduction of female to the first intromission) It’s owing to the elaborate description of emotional states proposed
depends on estrogen (estradiol or testosterone therapy) by psychologists and sexologists (Janssen, 2011). For instance,
(Meisel et al., 1984). the term “sexual desires” may not be entirely proper in animal

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studies. Sachs (2007) tried to unify terminology that would allow Concomitantly, the level of sexual motivation and sexual arousal
achieving greater translational of animal studies (by comparison stays low. Only later, the occurrence of non-contact erection
with sexological descriptions). He indicates that penile erection is indicates an elevated level of SA. However, there is still lack of
accepted as the golden parameter that measures elevated sexual approaching to the female. In the last phase of PEI, the male starts
arousal in males. In the case of psychogenic erections, he proposed to approach and investigate the female, which depicts the elevation
that two conditions must be met: (1) the occurrence of penile of SM and initiation of copulation (Sachs and Bialy, 2000; Bialy
erection and (2) sexual context. He also proposed (Sachs, 2000, et al., 2019). The initiation of copulation ends the PEI. Currently,
2007) to distinguish between two processes: sexual arousal and all processes GA, SA, and SM are back on the high level (Figure 2).
sexual motivation. Likewise, the dissociation between SA and SM could be
Motivation is a process “that causes an organism to seek a observed during the first initiation phase after the anticipatory
goal,” and sexual motivation is “impetus arising from internal or phase (the time between the introduction of the female to the first
external stimulation to seek out or create occasion for engaging mount). At that time, some males also can display non-contact
in sexual behavior.” Motivation manifests itself through the rate of erection. It’s observed more frequently in sexually naïve males, and
executing a response that changes the probability of the occurrence this type of erection can be visible without subsequent copulation
of an unconditioned stimulus (or the signal stimulus of an (personal observation). In humans, dissociation between sexual
instinctive behavior) or the conditioned stimulus (in second-order arousal and sexual motivation can be observed in asexual men.
reinforcement). Manifestation of motivation level is conditioning They tend to avoid sexual intercourse with partners, although the
type II also, named as instrumental reaction (Konorski and Miller, frequency of masturbation is on the average level of the male
1937; Konorski, 1967) or alternatively “operant behavior” proposed population (Portillo and Paredes, 2019). Furthermore, we can
by Skinner (1938). Thus, measurements of the instrumentalized distinguish, during behavioral tests, the motivational component
reaction (including frequency, latencies, and persistency) could by measuring only the approach behavior (Le Moëne and Ågmo,
serve as a reflection of motivation level. In laboratory research 2019). In this phase, a male presented an oriented reaction
of rats’ sexual behavior, different responses can be used, but toward the sexual-related cues. In other words, his reactions
usually, there are bar-pressing or runway, rewarded by contact to lead to greater sensory stimulation. During approach behavior,
mountings, intromission, or ejaculation (Beck, 1971, 1974, 1997; the autonomic parameters related to sexual arousal could be
Everitt et al., 1987, 1989; Beck et al., 2002). absent (including erection). Also, after the approaching phase,
These motivational regulations were profoundly studied the initiation of copulation and/or copulatory phase could not
using the pharmacological, electrophysiological, and optogenetic be observable. Describing this phenomenon in the context of
methods and are related mainly to the activity of the dopaminergic classic sexual behavior parameters we can notice: parameters of the
system of the ventral striatum. Although it seems to be non-specific instrumentalization of reaction, but improper latency and number
for sexual behavior, maintaining the intrinsic system of processing of intromission, and consequently often lack of ejaculation (we
all seeking behavior related to conditioned or unconditioned discuss these changes in Bialy et al., 2019). We can see that if there
stimuli. Reciprocal connections of the mesolimbic pathway with is a possibility to observe the separation of motivational and SA-
more sexual-specific structures such as MPOA enable channeling related parameters, there are at least two separate interconnected
motivational (reward-related) response and initiate sexual activity neuronal loops. Physiologically, one of them leads to a greater
(Everitt et al., 1989). By using the anticipatory paradigm (Bogacki- probability of object-related interaction (motivational aspect), and
Rychlik et al., 2021), mentioned before second-order conditioning the other controls the sexual-performance autonomic reactions
or pharmacological modulation (Rodríguez-Manzo and Canseco- (sexual arousal aspect).
Alba, 2023) of the ventral striatum, there is a possibility to partially General arousal and sexual arousal are closely interlinked
separate the motivational component from the sexual arousal processes as the reduction of GA (pharmacologically) decreases
activation. SA, and elevated sexual arousal increases general arousal (Pfaff,
In a more natural paradigm, dissociation between general 2017). On the other hand, not every rise in general arousal
arousal, sexual arousal, and sexual motivation can be visible, for automatically increases sexual arousal. From a neurophysiological
instance, during the post-ejaculatory period (PEI, see Figure 2) point of view, GA is a result of the activation of an ascending
based on results obtained from Sachs and Bialy (2000). About
half a minute after the first ejaculation (and slightly later after
followings), male 22-kHz vocalizations can usually be observed
simultaneously with natural immobility (Barfield and Geyer, 1972;
Sachs and Barfield, 1976). Such 22-kHz vocalization reflects a
relaxation state (Bialy et al., 2016), which occurs after a rapid
decrease of arousal (Bell, 1974), at the time when sleep-like spindles
in EEG can be recorded (Kurtz and Adler, 1973; Barfield and
Geyer, 1975). Taking together all these data, we can assume that FIGURE 2
about 1 min after ejaculation, general arousal, sexual arousal, and The GA, SA, and SM activation time sequence during the
sexual motivation are at a very low level. As the post-ejaculatory post-ejaculatory interval. Immobilization reflects a low level of GA,
SA, and SM. The occurrence of exploration indicates activation of
period continues, a rise in locomotion activity is observable. This
GA.; First NCE (non-contact erection) is associated with an increase
locomotion is manifested as exploratory behavior, focused on the in SA; The build-up in SM leads to the manifestation of approach
experimental chamber but not on the approach to the female. This behavior.
movement is mainly related to an enhanced level of general arousal.

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neuronal web consisting of, e.g., reticular formation with upper - These loops are regulated by higher neuronal systems, creating
pons and mesencephalon with noradrenergic, serotonergic, and reciprocal interactions at the three levels of the autonomic
cholinergic neurons, hypothalamus with histamine and hypocretin nervous system.
(orexin) neurons and thalamus. Disruption in the hypocretin
system is a suggested mechanism of narcolepsy (Adamantidis et al., Thus, the second assumption is:
2020). Lesions in these areas could eventually lead to coma in 2. The activity of the sexual-related ANS neuronal structures
humans (Silva et al., 2010; Kandel, 2013). The disruption of the can serve as a parameter describing male sexual arousal.
activity of the thalamocortical pathways is a proposed mechanism
of deep-sedative drugs used in anesthesiology (Malekmohammadi
et al., 2019). The activation of GA can also be visualized by the Levels of the autonomic nervous system
arousing effect of the pain stimulus during the Glasgow Coma
Scale examination (Todd et al., 2017). The behavioral reflection of
Mechanism of erection
GA is visible by all non-specific reactions, often with a reflexive
The penile erectile tissue consisting of pair corpora cavernosa
component, which corresponds with a normal state of awakeness.
and corpus spongiosum have a complex sinusoidal blood
In rats, we can describe it by, e.g., total locomotor activity during
system innervated atypically by parasympathetic, sympathetic,
exploration of the cage, number of rearing, and sometimes climbing
and nitrinergic NO fibers (non-adrenergic non-cholinergic fibers)
or digging.
(Giuliano et al., 1995). The rareness of it arises from the fact
The influence of GA during sexual interactions was pictured
that most blood vessels are innervated only by sympathetic
in some experiments with pain stimulation. Results showed that
fibers with dominantly adrenergic receptor expression. Adrenergic
moderate painful shock can promote copulation (approach to
stimulation participates in vasoconstrictive response, typically
female and intromission) (Barfield and Sachs, 1968; Sachs and
maintaining redistribution of blood flow with opposite local
Barfield, 1974). Interestingly, during PEI, moderate pain effectively
mechanisms of paracrine vasodilation as an antagonistic lever.
shortened this phase but only when applied after the end of 22-kHz
In opposition to that, sexual performance requires all autonomic
vocalizations. It was ineffective or even prolonged post-ejaculatory
types of fibers to be active simultaneously, creating probably the
period (PEI) when shocks were applied during 22-kHz emission
most complex autonomic response (Figures 3, 4). It seems that
(Pollak and Sachs, 1975). Therefore, it is possible that GA can
the foremost neurons responsible for the intensity of erectile
stimulate processes that are the most active at that moment. When
performance are pelvic nerves located in the sacral segment of
inhibition during the first part of the post-ejaculatory interval
the spinal cord, which contains all types of necessary fibers. The
dominates, then general arousal primarily enhances inhibitory
parasympathetic stimulation leads to the secretion of acetylcholine
processes, but when the elevation of SA starts to be observed,
from postganglionic fibers, and thus, by activating the M-type
general arousal stimulates sexual arousal and potentially sexual
receptors on the vascular epithelium (endothelium) generates the
motivation. The same problem can be observed in clinics in patients
vasodilatory effect by (nitric oxide) NO production. This effect is
with an enhanced anxiety level, which has an ambiguous effect
mediated simultaneously by the secretion of VIP and adenosine.
on sexual behavior (Janssen and Bancroft, 2007). Anxiety can
Similarly to parasympathetic stimulation, non-adrenergic non-
activate general arousal and, in this way, sexual arousal (Bancroft,
cholinergic fibers directly secrete NO, thereby contributing to
1999), but high anxiety levels can evoke erection dysfunction as
evoke erection in men and rats (Hull et al., 1994; Andersson and
well as premature ejaculation (Janssen and Bancroft, 2007). On
Wagner, 1995; Bialy et al., 1996).
the other hand, anxiety inhibits sexual motivation and initiation
At the same time, sympathetic stimulation increases, triggering
of copulation via neuronal networks in the nucleus accumbens
the cardiovascular system. It leads to positive tropic effects
related to initiation of copulation, activated by PKA (protein
on heart parameters (positive chrono-, ino-, and dromotropic
kinase A), CREB (cAMP response element-binding protein), and
effect), with an increase in heart rate and stroke volume. By
D1 receptor (dopamine receptor type 1) (Barrot et al., 2005;
adrenergic stimulation of the vascular component, this response
Bialy et al., 2010).
constricts most of the body’s blood vessels (including pelvic
arteries) and raises total resistance and blood pressure. In
penile arterioles, in a rest state, adrenergic activation inhibits
Male sexual arousal—Core assumptions the erection in this vasoconstrictive mechanism. However,
parasympathetic and nitrergic coactivation, together with local
It remains to ask what sexual arousal means and whether we mediators, e.g., prostacyclin, reduce the contraction of the
can distinguish specific neuronal loops relevant to it. We need to supply arterioles and provide sufficient fill of the penile corps.
make basic assumptions, which are listed below. Activation of the sympathetic nervous system also determines
1. We should assume that the most evident indicator of sexual another parameter of sexual arousal, which is ejaculation. It does
arousal in males is penile erection. so by innervating organs participating in ejaculation: the vas
If we assume the above implicatively, we have to consider that: deferens, seminal vesicles, prostate, and testicles (Giuliano, 2011;
Clement and Giuliano, 2016).
- The basis of sexual arousal is neuronal loops consisting of The next important mechanism refers to the atypical regulation
the afferent sensory inputs at the level of the spinal cord, of muscle tension in the pelvic floor group. In humans, contraction
creating synapses with efferent autonomic fibers responsible of these muscles blocks blood outflow by pressing on the vein
for erection and ejaculation. system. In rats, the quick muscular response provides rather more

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FIGURE 3
Autonomic nervous system (ANS) structures involved in sexual arousal. We pointed out three levels of ANS. Visual cues are more likely to elevate
sexual arousal in humans, while rats rely more on olfactory cues. BNST; bed nucleus of stria terminalis, LSt, lumbar spinothalamic cells, MPOA,
medial preoptic area; PAG, periaqueductal gray; SPFp, parvocellular subparafascicular thalamic nucleus; PB, parabrachial nucleus; NAmb, nucleus
ambiguus; NTS, nucleus tractus solitarius; RVLM, rostral ventrolateral medulla.

inflow than inhibits the blood return. Despite some physiological bulbocavernosus muscle contraction and very quick erection (at the
differences between species, the contraction of ischiocavernosus time less than 0.5 s).
and bulbospongiosus muscles during sexual activity is coordinated If the impulse from the internal genitalia can elevate sexual
by the autonomous pathways on the spinal level (Hull et al., 2002). arousal, is it possible for a male to have an elevated level of
Thus, we can see that the most apparent arousal parameters are sexual arousal without an explicit sexual context, but only as
functions of autonomic neuronal efferent activation (output). So, the effect of activation of these internal receptors? In the frog,
the question is, which receptors and afferent sensory inputs of these such information is crucial for the clasp reflex (Tarchanoff, 1887),
reflexes are necessary to activate the neurons of the sacral segment but innervations of internal organs can differ between groups
of the parasympathetic nervous system? of animals. So-called vacuum activity (“germ. Leerlaufreaktion”)
proposed by Konrad Lorenz, can suggest that information from
Sensory receptors internal organs can induce elevated sexual arousal. This term means
Surprisingly, such receptors are relatively poorly investigated that an animal discharges an elevated level of sexual arousal on an
(Figure 4). Krause corpuscles are one of the receptors detected inappropriate object and in an inadequate situation. This reaction
on the glans penis. They are rapid-adapting low-threshold is sometimes seen in dogs that mount, for example, plush toys.
mechanoreceptors, sensitive to light touch and vibration (optimum This mentioned hypothesis is an attempt to understand the highly
40–80 Hz). Activation of these receptors induced penile erection, sexually aroused behaviors of males, with problematic and unclear
and genetic ablation of Krause corpuscles impaired intromission context from external sexual-related cues. In these cases, we reject
and ejaculation in mice (Qi et al., 2023). In men, the glans penis other explanations, such as forms of dominance, fetishism, or
stimulation induced contraction of the cavernous muscle, the sexual preference, and can assume that such behavior is related to
smooth muscle of the seminal vesicle, and the vassal ampullary. internal sexual cues.
These reflexes were inhibited by the glans penis, cavernous muscle, The critical question about the role of sensory information from
seminal vesicle, and vassal ampulla anesthesia (Shafik et al., 2007). internal genital organs on sexual arousal in mammals is currently
However, very little is known about internal sensory input from under verification. The first experiments were not promising, as
seminal vesicles or other parts of internal genitalia and their role removing seminal vesicles had no effect on copulation (Beach
in autonomic nervous system activity in mammals. In rats, can and Wilson, 1963; Larsson and Swedin, 1971). Beach and Wilson
be observed erections named touch-based (ex-copula) that can be (1963) suggest that the effect of internal genital receptors can be
evoked by constant pressure induced by a lead stick transverse to better visualized in animals with strictly seasonal sexual activity.
the penile root. Such pressure can be detected at the internal genital More recently, the connection between the filling state of seminal
tract as well (seminal vesicles). Ex-copula erections can be induced vesicles and copulation level in sexually naïve mice at the start
after a relatively long time–a few minutes (Sachs and Bitran, of the experiment has been found (Birkhäuser et al., 2012).
1990). It differed from erections generated during copulation by fMRI (functional magnetic resonance imaging) brain scans of
stimulation of the glans penis during intromission with rapid men also suggested that the CNS activity could be dependent

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to be involved in the reception of stretching, nociception,

and inflammation sensations (agonized by, e.g., capsaicin and
cinnamon). Despite SVs, TPRA1 receptors are present in the
prostate, corpus cavernosum, and glans penis (Figure 4). Thus,
these receptors probably participate in neuronal loops that cause
smooth muscle contraction and the release of neurotransmitters
important in the ejaculation reflex, i.e., NE from sympathetic
postganglionic fibers (Birowo et al., 2009); and 5-HT acting
probably as a paracrine mediator produces in neuroendocrine cells
of SV (Aumüller et al., 2012) and also speculatively of intestinal
origin (Jiménez-Trejo et al., 2021).
It is hypothesized that some form of dysregulation
in the sensory innervation of genitalia could be one
of the pathophysiological mechanisms of premature or
delayed ejaculation. SVs are innervated by sympathetic and
parasympathetic fibers which contain noradrenaline, acetylcholine,
FIGURE 4
Receptors, afferent and efferent innervation of the internal genital
serotonin, and neuropeptides VIP (vasoactive intestinal peptide),
organs and penis. TPRA1 receptors innervate internal genital organs PHI (peptide histidine isoleucine) NPY (neuropeptide Y),
as well as the penis. Krause corpuscles innervate the penis. The somatostatins, as well as CGPR and substance P (CGPR and
main afferent pathways conducting sensory signals from internal
genital organs travel along the pudendal nerve, hypogastric nerve,
substance P seems to be related to sensory afferent). The function
and pelvic nerves. The efferent pathways that innervate the internal of SVs depends also on androgens treatment (Pinho et al.,
genital organs and penis contain sympathetic fibers with NA 1997). Serotonin and augmented levels of this neurotransmitter
(noradrenergic) and NPY receptors, parasympathetic with Ach
(cholinergic) and VIP receptors, NANC with nitric oxide (NO),
caused by serotonin selective reuptake inhibitors (SSRI) modulate
serotoninergic with 5-HT receptors.; SVs, seminal vesicles; VDs, noradrenergic smooth muscle contraction (Birowo et al., 2010)
vasa deferentia; NTS, nucleus tractus solitarius; PB, parabrachial after hypogastric nerve stimulation (Kim and Paick, 2004) via 5-
nucleus; SPFp, parvocellular subparafascicular thalamic nucleus;
5-HTR, Serotonin receptor; AchR, Acetylcholine receptor; NAR,
HT1A, 1B, and 2C receptors (Waldinger et al., 1998). Regulation of
Noradrenergic receptor; TRPA1, Transient receptor potential cation serotonin levels is a crucial neurochemical mechanism controlling
channel subfamily A member 1; VIPR, Vasoactive intestinal peptide ejaculation, clearly depicted by pharmacological treatment
receptor; NPYR, Neuropeptide Y (tyrosine) receptor; NANC,
Non-adrenergic Non-cholinergic fibers; SNS, Sympathetic Nervous
of premature ejaculation using SSRI and 5-HT1A receptor
System; PSNS, Parasympathetic Nervous System. antagonists (Olivier et al., 2023).
Therefore, the next question regards what pathways conduct
impulses from internal organs reaching the higher levels of the
autonomic nervous.
on the filling state of seminal vesicles (Weisstanner et al., 2022).
These results must be considered preliminary, with significant
limitations related to a lack of additional control groups. There Spinal reflexes
is a necessity to eliminate an alternative, central mechanism of In humans and animals with an intersected spinal cord,
sexual satiety to avoid misinterpretation. Such an effect could erections are still feasible to induce. Moreover, touch-based
be induced by shifting in the profile of neurotransmission, erections can be evoked significantly easier in mice and rats with
i.e., dopamine (Hull et al., 1999; Canseco-Alba et al., 2022), an intersected spinal cord when compared to control animals
serotonin (Matsumoto et al., 1997), noradrenaline (Rodríguez- (Meisel and Sachs, 1980; Sachs and Garinello, 1980). It indicates
Manzo and Fernández-Guasti, 1994), opioids (Rodríguez-Manzo that inputs coming from higher levels of CNS into the erection
and Fernández-Guasti, 1995), GABA (Rodríguez-Manzo and center in the spinal cord are predominantly inhibitory. However,
Canseco-Alba, 2017), endocannabinoids (Rodríguez-Manzo and after copulation to satiety, the inhibition of erection can’t be
Canseco-Alba, 2023), and glutamate (Rodríguez-Manzo, 2015). abolished by the thoracic spinal cord block. It suggests that such
This regulatory mechanism of sexual satiety emerged as copulation directly inhibits the local spinal intrinsic system (Sachs
a complex interconnected web with differentially expressed and Bitran, 1990). Together, these results show that in mammals,
neurochemical specificity of particular neuronal clusters that spinal reflexes alone are sufficient to induce erection. Analogical
participate in motivational, arousal, and sensory aspects. Thus, results were obtained during experiments with the ejaculatory
it could be challenging to definitely separate the mentioned mechanism, depicting the spinal intrinsic system for ejaculation.
interoceptive activation, especially in the context of reliable The autonomy of this system is underlined by the presence of
biomedical translation. pacemaker neurons, generating the efferent motoneuronal signal,
Surprisingly, there is very little experimental data on sensory mediating the contraction of muscles associated with the genital
innervation of seminal vesicles. Only one article showed that, in tract (see for review: Carro-Juárez and Rodríguez-Manzo, 2008).
humans, SVs contain multimodal sensory receptors type TRPA1 Men with transected spinal cords at the level of T10 still
[transient receptor potential cation channel, subfamily A, member can display some preserved genital reflexes (penile erection) after
(1) expressed on nerve fibers type C and epithelium]. They exposition to pornographic movies (see Sachs, 1995). In this case,
were found in fibers with colocalization with nNOS (neuronal the speculative pathway of transduction includes sensory fibers
nitric oxide synthase) and CGRP (calcitonin gene-related peptide) of the hypogastric nerve. The same level of spinal transection in
(Rahardjo et al., 2023). TRPA1 receptors are commonly known women does not eliminate the ability to achieve orgasm visible in

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fMRI brain activity during genital stimulation (Komisaruk et al., In vertebrates, sexual arousal consists of processes observable
2004), with a potential alternative transduction pathway through in fish, amphibians, and generally in groups of animals without
the parasympathetic vagus nerve (received supraspinal nucleus developed neocortex. These groups of animals have functional
solitary tract, NTS, and nucleus ambiguus). evolutionary older areas of the brain like the paleocortex, which
constitutes the olfactory system with the olfactory bulb, amygdala,
Supraspinal autonomic nervous system and piriform cortex, and also the archicortex, which forms the
loops—Brainstem level and forebrain hippocampus. These structures play an essential role in the
Information from internal organs, in this case from sexual detection of olfactory cues and regulation of sexual arousal and
organs, reaches the supraspinal level: lower brainstem with the sexual behavior, as well as hormonal status [migration and activity
most important structures such as nucleus solitary tract, nucleus of GnRH (LH-RH) neurons important in gonadotrophin release]
ambiguus, parabrachial nucleus, rostral ventrolateral medulla, (Schwanzel-Fukuda and Pfaff, 1989). In mammals, crucial for
nucleus raphe pallidus. The brainstem structures are connected regulating sexual behavior structures are the amygdala, bed nucleus
with and regulated by the hypothalamus, amygdala, insular cortex, of stria terminalis, and medial preoptic area, which create bilateral
and anterior cingulate cortex. The amygdala provides tagging of the reciprocal loops. This system displays anatomical and functional
biological significance of environmental stimuli, correspondingly sexual dimorphism with high expression of steroid receptors
with the detection of internal information and hormonal state of an (T, DHT, E, and P) (Hull et al., 2002). Medial amygdala and bed
organism. The insular cortex integrates multiple internal sensations nucleus of stria terminalis provide essential for the elevation of SA
and, by that, participates in the generation of visceroception and processing of sexual-related cues during the occurrence of stimuli
even more general interoceptive awareness. The dorsal posterior (Fiber et al., 1993; Liu et al., 1997; Kondo and Sachs, 2002).
insula functions as the primary interoceptive cortex by receiving Another source of sensory cues related to the elevation of sexual
and integrating inputs from visceral, pain, and thermal receptors arousal is tactile cues. The most important ones come from the
via the thalamus, parabrachial nucleus, and nucleus of the solitary genital region, which provokes ex-copula erection and erection
tract or nucleus tractus solitarius (NTS). Anterior insula integrates during copulation. Other tactile cues are also important, especially
interoceptive signals with emotional and cognitive processes, in sexually naïve rats. For instance, the reduction of tactile cues
thus participates in conscious body sensation. The insula is a from vibrissae in sexually naïve male rats significantly reduced the
primary viscerosensory as well as visceromotor area that controls number of males who started to copulate with receptive females
sympathetic and parasympathetic outputs. The insula exhibits (Biały and Beck, 1993). Other cues, like visual and auditory, are also
functional lateralization, with the right insula dominating in important and can be received on the level of the midbrain as well
sympathetic activation (Craig, 2003). Anterior cingulated cortex, as the amygdala and neocortex in mammals (Lenschow et al., 2022).
together with the insula, can work as a prediction error signal
(Kleckner et al., 2017) from internal organs with interoceptive
experience and via parabrachial nucleus and periaqueductal gray
(Lamotte et al., 2021) may convey interoceptive error attenuation.
Neuroimaging study in men and male
This predictive-signal analysis mechanism, suggests that this area rats during elevated sexual arousal
serves an important role in the perception of signal input (rather
than strictly reception of it) (Barrett and Simmons, 2015). Thus, Neuroimaging studies of brain activity are mainly based
dysregulation of this mechanism could hypothetically lead to on the measurement of changes in blood flow and neuronal
hyper/hypo-stimulation of lower regulatory loops, resulting in metabolism in various neuronal structures, using PET and fMRI
clinically visible sexual dysfunction. Furthermore, it is possible techniques. In rats with stress-induced depressive behavior, erectile
that if this mechanism doesn’t work properly it can contribute dysfunction correlates with alteration activity of such structures
to cenesthetic hallucinations, symptoms that sometimes occur as the amygdala, thalamus, hypothalamus, caudate putamen,
in patients with schizophrenia (or during non-schizophrenic cingulate cortex, insula, visual-sensory and motor cortices, and
psychotic episodes). cerebellum (Chen et al., 2018). Meta-analysis of men’s brain activity
during elevated levels of SA pointed out a high activity of regions
related to strictly autonomic sexual-related response, but also other
External cues related to sexual context regions corresponding to cognitive, emotional, and motivational
components (Kühn and Gallinat, 2011; Stoléru et al., 2012; Poeppl
Sachs (2007) distinguishes between arousal-related erections, et al., 2014). The data was mainly obtained from tests in which
which require sexual context, and erections occurring without men were exposed to erotic visual stimuli, mainly photos or videos.
sexual context (e.g., noctuary erections). He assumed that sexual The right insula (>60% of analyzed articles) and the left anterior
arousal is not only the activation of the autonomic nervous cingulate cortex (>60% of analyzed articles) are the structures that
system–parasympathetic with sympathetic nervous system but increase their activity during elevated SA. The insula is a structure
also neuronal networks that detect sexually important cues to involved in visceral sensory processing (Craig, 2003), and the right
elevate sexual arousal. This point of view is complementary to insula receives sensory information from the penis and is activated
that presented herein. However, in most experimental schedules, upon genital stimulation (Holstege and Huynh, 2011). The level of
during exposition to sexual stimuli, similar structures that detect insula activation correlates with penile erection rigidity (Moulier
and integrate stimuli control partly both motivational processing et al., 2006; Wang et al., 2017). Additionally, to the insula, the
and arousing response. Focusing only on autonomic activation activity of the anterior cingulate cortex (particularly Brock areas
allows us to avoid oversimplification and obtain data precisely 24 and 32) correlates with penile erection, and stimulation of this
related to SA. area can trigger an erection (see Holstege and Huynh, 2011). On

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Bogacki-Rychlik et al. 10.3389/fnbeh.2023.1330460

the other hand, elevated thalamic activity is observed in >50% of

articles, and it may reflect the elevated level of general arousal. Very
often, elevated levels of activity have been observed in structures
which activity is linked to the analysis of visual stimuli, such as
the occipital, parietal, and temporal cortex (Stoléru et al., 2012). It’s
worth mentioning that there were no differences in the activity of
cortical regions between heterosexual and homosexual men.
Some differences between men and women, as well
as heterosexual and homosexual men, can be seen in the
hypothalamus response to olfactory stimuli. In homosexual
men and heterosexual women, the ventromedial nucleus of the
hypothalamus and the preoptic area respond to olfactory cues after
exposure to an androgen-like compound. In turn, similar activity
of the dorsomedial and paraventricular nuclei of the hypothalamus FIGURE 5

was detected in heterosexual and homosexual men after exposure Scheme of the chamber for the NCE test. Based on Sachs et al.
(1994) and Bogacki-Rychlik et al. (2022).
to estrogen-like compounds (Savic et al., 2001, 2005).
Ejaculation leads to strong activation of the right insula,
dorsal temporal lobe, and globus pallidus. In addition, some
structures, such as the amygdala and the parietal, prefrontal, medial
hand, during PEI, such NCEs are not influenced by any frustrating
orbitofrontal, and temporal cortices, mainly on the left side, can
context related to the separation of individuals.
decrease their activity (Holstege and Huynh, 2011).
Non-contact erections mostly depend on odor cues from a
receptive female (Sachs, 1997), detected by the main olfactory
system (Kondo et al., 1999). Lesion of olfactory bulbs significantly
Animal model for research psychogenic reduces NCEs (Edwards and Davis, 1997). The medial amygdala
sexual arousal in males (NCE-test) seems to be one of the key structures in the regulation of NCEs.
Lesion of the medial amygdala eliminates NCEs (Kondo et al.,
Sachs et al. (1994) proposed a new model–a non-contact 1997; Kondo and Sachs, 2002). Moreover, the number of non-
erections test (NCEs) to investigate sexual arousal in male rats. This contact erections correlates with medial amygdala volume (Cooke
type of erection is proposed to be closely related to psychogenic et al., 2000). A significant effect also can be observed after the bed
erections in men (Sachs, 1995). In this model, sexual arousal can nucleus of the stria terminalis (BNST) lesion (Liu et al., 1997).
be measured by the latency to the first erection and the number Interestingly, a lesion of the medial preoptic area (MPOA) can have
of non-contact erections during the particular trial, usually 20– no significant effect on NCEs (Liu et al., 1997). It suggests that
30 min. In the NCEs test, the sexual context is well-established and
the main output from the amygdala and BNST important to evoke
relatively easy to modify, which enables an investigation of sexually
NCEs is not directed to MPOA. However, MPOA can be involved
arousing factors. Animals in the chamber (Figure 5) are separated,
in NCEs as elevated levels of dopamine in MPOA can enhance
without the possibility of receiving tactile stimulation, but the male
NCEs (Adachi et al., 2003), and this structure also regulates the
is exposed in a controlled manner to the visual, auditory, and odor
tonus of the sympathetic and parasympathetic nervous system
cues, depending on the barrier type and direction of airflow. This
(Hull and Dominguez, 2007).
chamber can be modified by various actions. A transparent barrier
Paraventricular nucleus (PVN) is a structure that has a direct
enables the exposition of visual cues from females. If we would like
oxytocinergic output on sacral spinal cord interneurons and can
to reduce the significance of visual cues, there is a possibility to use
induce penile erection, which can be spontaneous without visible
black walls instead of transparent ones (Sachs, 1997).
sexual context as well as NCE in a sexual context (Argiolas and
To reduce the influence of auditory ultrasonic vocalizations,
an additional barrier above the chamber eliminates the detection Melis, 1995; Melis et al., 1999; Succu et al., 2007). Another structure
of vocalizations emitted by another animal (Bogacki-Rychlik et al., involved in NCEs is the nucleus accumbens (NAcc). A lesion in
2022). the NAcc reduces NCEs (Kippin et al., 2004), and an elevated
If we would like to reduce odor cues from the female, we can level of dopamine in NAcc increases the incidence of NCEs (Sanna
use sucking air from the female’s side (Sachs, 1997). et al., 2009, 2015). On the other hand, LHA (lateral hypothalamus)
Detection of a non-contact erection during trial required direct exhibits inhibitory effects on NCEs. Lesion located at the anterior
observation of penile erection. However, if it is not possible, two part of LHA facilitates NCEs. The same lesion inhibits ejaculation
additional co-occurring elements of male behavior could be used during copulation (Kippin et al., 2004). Lesion of the nucleus
as indicators of such response, i.e., tiptoe with genital grooming paragigantocellularis located in the medulla has no effect on NCEs,
(similar to movement and posture observed after intromission) although it has a significant influence on reflexive erection and
or genital grooming with hip movement. Similar non-contact ejaculation (Liu and Sachs, 1999).
erections can be observed during the post-ejaculatory period, Analysis of neuronal activity measured by c-Fos
usually shortly before the end of 22-kHz vocalizations (Sachs and immunostaining showed differences related to NCEs at the
Bialy, 2000; Bialy and Sachs, 2002). Such post-ejaculatory erections NAcc core and shell, medial amygdala (anterior and posterior
can be a more convenient method to analyze NCEs, but with part), BNST, MPOA (Kelliher et al., 1999), as well as at the
less possibility to modify the sensory environment. On the other orbitofrontal cortex, piriform cortex, PVN, anterior hypothalamus

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(AHA), ventromedial hypothalamus (VMH), basolateral amygdala erection on the spinal and supraspinal level determines the level
(BLA), and central nucleus of amygdala (CeA) (Coria-Avila et al., of sexual arousability. Based on these assumptions, we analyzed
2018; Ramírez-Rodríguez et al., 2021). We should keep in mind autonomic reflexes and a network of interconnectedness of ANS
that elevated C-Fos immunostaining as well as c-fos expression on different neuronal levels. We also visualized these relations by
do not simply reflect neuronal activity but activity with induction presenting data from brain imaging of humans and rats. Next, we
of gene expression and neuronal plasticity (Bialy et al., 1992; described one of the animal models–the NCEs test, to investigate
Kaczmarek, 1992, 1993, 2018; Bialy and Kaczmarek, 1996) so all SA, closely related to psychogenic erections evoked by sexual-
active structures can be not visible using this technique. related cues. This model is based on top-down regulation of the
As we mentioned, NCEs depend on androgens–testosterone autonomic nervous system related to SA.
(T) and dihydrotestosterone (DHT), but not estrogens (E2) therapy For the sake of clarity and to avoid the informational excessive,
after castration (Manzo et al., 1999; Cooke et al., 2000). Androgens we didn’t discuss the mechanism of ejaculation and its relationship
(T and DHT) but not estradiol implanted into sexually dimorphic to the measurement of SA. Ejaculation has to be considered as
postero-dorsal medial amygdala maintain NCEs after castration, the most prominent indicator of sexual arousal accumulation.
and latency to first NCE positively correlates with the proximity However, neuronal loops regulating this mechanism especially
of androgen implantation to the postero-dorsal region of medial specific to ejaculation lumbar spinothalamic (LSt) cells (Truitt and
amygdala (Bialy and Sachs, 2002). On the other hand, the blockade Coolen, 2002) are different from erection on discussed levels of a
of the androgen receptors but not the inhibition of aromatase of neuronal system (Coolen et al., 2004; Veening et al., 2014; Veening
testosterone in the postero-dorsal medial amygdala inhibits NCEs and Coolen, 2014). It’s visible by the fact that erection can be
(Bialy et al., 2011). The hormonal profile and neuronal network of maintained without the triggering of ejaculation, and oppositely,
NCEs differ from the erection observed during copulation. Erection ejaculation can be induced without the occurrence of erection
during initiation of copulation (measured by intromission latency) (Clement and Giuliano, 2016). This problem should be discussed
depends on estradiol but not androgen therapy (Meisel et al., more in consecutive review.
1984; Cooke et al., 2003). Furthermore, estradiol and the activity From a clinical perspective of urological practice, important
of aromatase (which transforms testosterone to estradiol) in MeA questions concerned with the anatomy of innervation of genitals
and medial preoptic area are crucial for erection during copulation and the physiological construction of reflex arches. To maximize
(Dominguez et al., 2001; Putnam et al., 2003; Hull and Dominguez, the therapeutic effect while minimizing the side effects, the
2007). Results from NCEs tests and copulation show that different optimal assessment of the resection range is crucial. Therefore,
networks are involved in erection depending on the context. the knowledge of afferent sensory input and efferent output could
Non-contact erection test tests are useful to investigate arousal answer a question of, e.g., point of resection of seminal vesicles
in the context of sex preference: same-sex, opposite-sex, or juvenile- or type of prostatectomy. Paradoxically, innervations of internal
object (pedophilic-like) preference. In some circumstances (e.g., genital organs and reflexes related to sexual arousal on the level
prenatal inhibition of aromatase of testosterone by letrozole), of spinal cord are relatively poorly understood and require future
enhanced same-sex preference can be visible in non-contact tests investigation to complete a picture of down-top and top-down
according to the fact that males display significantly more erections regulation of sexual arousal.
when exposed to other male compared to receptive female.
Interestingly, such same-sex preferences in NCEs test do not reflect
preference in copulation, in which they prefer to mate with a Author contributions
receptive female (Olvera-Hernández et al., 2015).
In adult males, same-sex preference, or preference to young WB-R: Writing – original draft, Writing – review and editing.
males, can be evoked by conditioning to male marked by a KG: Writing – review and editing. MB: Writing – original draft,
neutral odor (almond as a CS) associated with D2 dopamine Writing – review and editing.
receptor activation. Such conditioning evoked NCEs preferentially
when male was exposed to almond-scented male (Coria-Avila
et al., 2018) or young juvenile male compared to receptive
female. Also, in these experiments, preferences were visible in the
Funding
NCEs test, but not in the copulation test with receptive female
The author(s) declare financial support was received for the
(Ramírez-Rodríguez et al., 2021).
research, authorship, and/or publication of this article. This study
was supported by the Medical University of Warsaw and the Young
Researcher Grant: 1MA/5/M/MB/N/20/20 and carried out with the
Summary and implication for future use of the CePT infrastructure financed by the European Union—
research the European Regional Development Fund within the Operational
Program “Innovative Economy” for 2007–2013.
In this review, we clarified the term male sexual arousal and
its relation to general arousal and sexual motivation. We did this
to reconceptualize some issues that emerged from behavioral and Conflict of interest
physiological results obtained on different models and conceptual
paradigms. In this review, we made two core assumptions: (1) The authors declare that the research was conducted in the
penile erection is the most important, pivotal for other parameters, absence of any commercial or financial relationships that could be
indicator of SA; (2) the activity of parts of ANS that regulate the construed as a potential conflict of interest.
Frontiers in Behavioral Neuroscience 10 [Link]
Bogacki-Rychlik et al. 10.3389/fnbeh.2023.1330460

Publisher’s note organizations, or those of the publisher, the editors and the
reviewers. Any product that may be evaluated in this article, or
All claims expressed in this article are solely those of the claim that may be made by its manufacturer, is not guaranteed or
authors and do not necessarily represent those of their affiliated endorsed by the publisher.

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