However, other authors do not exclude the possibility of USVs production by female mice because during interaction it is not easy to distinguish the animal that produces vocalizations. Using devocalizing males, no USVs were detected during male-female interactions and this supports the idea that is male mice that emitted vocalizations (Sugimoto et al., 2011). This type of calls is primarily attributed to males to attract females and has an important role in social/sexual behaviors (Egnor and Seagraves, 2016). The most characterized adult mice USVs are those emitted during male-female interactions and/or in presence of female odor cues/urine. Also adult mice produce USVs in different situations such as courtship, mating and social interaction. The second type of USVs concerns vocalizations of juvenile mice during social interactions and it is correlated with social bonding and motivational level of mice (Panskepp et al., 2007 Peleh et al., 2019). D’Amato and colleagues reported an increased number of isolation calls in pups born from mothers with a lower maternal responsiveness such as BALB/c females in comparison with C57BL/6 mothers (D’Amato et al., 2005). In addition, it is also interesting the idea that emotional state and responsiveness of mother influence the emission of pups USVs. Moreover, experiments using mice with alleles linked to social bonding and separation distress, like oxytocin or mu-opioid receptor, demonstrated that isolation-induced USVs are emitted in response to affective variations (Winslow et al., 2000 Moles et al., 2004). Indeed, different types of calls are produced in response to particular conditions such as the presence of odor of an unfamiliar, potentially infanticidal, adult male (Branchi et al., 1998). In addition, alterations in the pups USVs features reveal modifications in emotional states of pups and therefore in arousal states of mother (D’Amato et al., 2005 Lahvis et al., 2011). Pups USVs represent an early communicative behavior of the mother-pup dyad indeed USVs trigger maternal care and facilitate communication between mother and offspring (D’Amato et al., 2005 Hernandez-Miranda et al., 2017). For the first time, Zippelius and Schleidt described USVs emitted by pups during separation from the mother and the littermates and they referred to USVs as “whistles of loneliness” able to elicit mother retrieval (Zippelius and Schleidt, 1956). Three types of USVs have been largely studied in laboratory mice: - isolation-induced USVs in pups, - interaction-induced USVs in juvenile mice and - interaction-induced USVs in adult mice ( Figure 1). Mice predominantly communicate using the ultrasonic vocalizations (USVs) with a frequency from 30 to 110 kHz (Holy and Guo, 2005). Mouse communication occurs both in the audible and ultrasonic range of sound frequencies (Ehret and Bernecker, 1986 Zippelius and Schleidt, 1956). Indeed, several NDDs murine models exist and an intense study of USVs is currently in progress, with the aim of both performing an early diagnosis and to find a pharmacological/behavioral intervention to improve patients’ quality of life. A particular focus will be devoted to mouse USVs in the context of NDDs. This review covers the topics of USVs characteristics in mice, contexts for USVs emission and factors that modulate their expression. They have long been studied in the ethological context, but recently they emerged as a precious instrument to study pathologies characterized by deficits in communication, in particular neurodevelopmental disorders (NDDs), such as autism spectrum disorders. The understanding of different calls meaning is still missing, and it will represent an essential step forward in the field of USVs. Indeed, different ultrasonic call categories exist and have already been defined. USVs can be acquired by means of specific tools and later analyzed on the base of both quantitative and qualitative parameters. USVs have been greatly characterized in specific life phases and contexts, such as mother isolation-induced USVs for pups or female-induced USVs for male mice during courtship. Mice use ultrasonic vocalizations (USVs) to communicate each other and to convey their emotional state.
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