The postnatal ontogeny of the sexually dimorphic vocal apparatus in goitred gazelles (Gazella subgutturosa)

This study quantitatively documents the progressive development of sexual dimorphism of the vocal organs along the ontogeny of the goitred gazelle (Gazella subgutturosa). The major, male‐specific secondary sexual features, of vocal anatomy in goitred gazelle are an enlarged larynx and a marked laryngeal descent. These features appear to have evolved by sexual selection and may serve as a model for similar events in male humans. Sexual dimorphism of larynx size and larynx position in adult goitred gazelles is more pronounced than in humans, whereas the vocal anatomy of neonate goitred gazelles does not differ between sexes. This study examines the vocal anatomy of 19 (11 male, 8 female) goitred gazelle specimens across three age‐classes, that is, neonates, subadults and mature adults. The postnatal ontogenetic development of the vocal organs up to their respective end states takes considerably longer in males than in females. Both sexes share the same features of vocal morphology but differences emerge in the course of ontogeny, ultimately resulting in the pronounced sexual dimorphism of the vocal apparatus in adults. The main differences comprise larynx size, vocal fold length, vocal tract length, and mobility of the larynx. The resilience of the thyrohyoid ligament and the pharynx, including the soft palate, and the length changes during contraction and relaxation of the extrinsic laryngeal muscles play a decisive role in the mobility of the larynx in both sexes but to substantially different degrees in adult females and males. Goitred gazelles are born with an undescended larynx and, therefore, larynx descent has to develop in the course of ontogeny. This might result from a trade‐off between natural selection and sexual selection requiring a temporal separation of different laryngeal functions at birth and shortly after from those later in life. J. Morphol. 277:826–844, 2016. © 2016 Wiley Periodicals, Inc.     

一种喉部疾病口语声诊断新方法

本文提出了声带的三质量块模型,并应用这模型模拟病噪产生的嘶哑语声,这些嘶声包括有声带闭合不全,声带小结,声带麻痹,喉炎,声带淀粉样变和声门癌等十六种典型情况。采用快速傅里叶变换,线性预测,倒谱技术和离荼余弦变换等分析各类喉病引起的嘶哑语声,实验结果表明声带模型分析法是喉病诊断的一种有效方法。     

Sexual Dimorphism of the Larynx of the Mongolian Gazelle (Procapra gutturosa Pallas, 1777) (Mammalia, Artiodactyla, Bovidae)

The larynges (except for the epiglottis) of two adult Mongolian gazelles, one male and one female, were dissected. This species is characterized by a pronounced sexual dimorphism of the larynx. Dimorphism with regard to the size of the entire larynx and of the thyroid cartilage is about 2:1 whereas the difference of mean body mass is about 1.3:1 between males and females. Unexpectedly, and in contrast to other bovids, the larynx of the male Mongolian gazelle has a paired lateral laryngeal ventricle. However, in contrast to horse, dog, pig and many primate species also possessing such a paired ventricle, its rostral opening in the Mongolian gazelle is situated lateral to the corniculate process of the arytenoid cartilage. The neck of the laryngeal ventricle is embraced by the bifurcated cuneiform process of the epiglottis. Despite the enlarged laryngeal cartilages, the vocal process of the male arytenoid cartilage is relatively shorter than that of the female. The male thyroarytenoid muscle is clearly separated into a rostral ventricular muscle and a caudal vocal muscle whereas the female's, as in other bovids, is almost uniform. The lateral sac of the two-chambered laryngeal ventricle in the male projects laterally between the ventricular and the vocal muscle. As in the domestic bovids and in many other artiodactyls the larynx of the male Mongolian gazelle is lacking any rostrally directed membraneous portion of the vocal fold. Instead, the thick and tough bow-like vocal fold projects caudally into the infraglottic cavity and is supported by a peculiar pan-like fibroelastic pad. This resilient element, situated medial to the bipartite thyroarytenoid muscle, might be a homologue of the vocal ligament, eventually including lateral portions of the elastic cone. A fibroelastic pad is absent in the female. The resilient floor of the laryngeal vestibulum, ventral to the fibroelastic pad, is rostrally and caudally subducted by tube-like spaces. Evolutionary enlargement of the male larynx, including the vocal folds, and of the caudal portions of the vocal tract may have shifted the fundamental and formant frequencies to a lower register. The paired lateral laryngeal ventricle might produce an amplitude increase of the vocalizations assisted by differential action of the bipartite thyroarytenoid muscle. In addition, the peculiar shape, size and tough consistency of the male vocal folds may, as in roaring felids, assist in producing high amplitude and low frequency vocalizations. Perhaps the biological role of the enlarged male larynx of Procapra gutturosa has evolved in relation to its mating system. In the rutting season, dominant males establish individual territories and maintain harems. During prolonged courtship prior to mating, these males perform an acoustic display uttering loud and guttural bellows. In addition, the bulging ventral neck region of males may serve as an optical attractant for the females. Thus, the evolution of the enlarged larynx of the male Mongolian gazelle may have been favoured by sexual selection.     

Sexual dimorphism and bilateral asymmetry of syrinx and vocal tract in the European starling (Sturnus vulgaris)

Sexually dimorphic vocal behavior in zebra finches (Taeniopygia guttata) is associated with a 100% larger syrinx in males and other morphological adaptations of the sound source. The songbird syrinx consists of two independent sound sources, whose specialization for different spectral ranges may be reflected in morphological properties, but the morphology of labia and syringeal skeleton have not been investigated for lateralized specializations. Similarly, little is known whether the morphology of the songbird vocal tract reflects differences in vocal behavior. Here, we tested the hypothesis that different vocal behavior and specialization is reflected in the morphology. We investigated syringeal and upper vocal tract morphology of male and female European starlings (Sturnus vulgaris). Female starlings exhibit smaller vocal repertoires and sing at lower rates than males. In males, the left syrinx produces mostly low frequencies, while the right one is used for higher notes. Macroscopic and histological techniques were used to record nineteen measurements from the syrinx and the vocal tract which were tested for sexual differences in syrinx and vocal tract and for lateral asymmetry within the syrinx. Sexually dimorphic vocal behavior is reflected in the morphology of the starling syrinx. Males have a larger syrinx with the size difference attributable to increased muscle mass and three enlarged elements of the syringeal skeleton. The upper vocal tract, however, does not differ between males and females. Distinct lateralization was found in two elements of the syringeal skeleton of females, and the labia in the left syrinx are larger than those on the right in both sexes. The sexual dimorphism of the syringeal size is smaller in starlings (35%) than in zebra finches (100%), which is consistent with the different vocal behavior of females in both species. The morphological differences between the two sound sources are discussed in relation to their vocal performance. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Some aspects of dimorphism and human speech

The production of human speech involves thre supralaryngeal airway filtering the acoustic energy generated by either laryngeal or noise sources. Both the supralaryngeal vocal tract and the larynx are subject to sexual dimorphism in anatomically modernHomo sapiens, yielding acoustic signals that can convey a speaker’s sex. Formant frequencies can be lower because the adult male supralaryngeal vocal tract is often longer. The fundamental frequency of phonation also can be lower because of the enlargement of the thyroid cartilage of the larynx and concomitant increases in the length and mass of the vocal cords. Secondary, learned supralaryngeal vocal tract maneuvers can also lower or raise formant frequencies. These acoustic effects yield male-female «microphonetic» dialects that are ultimately related to the average dimorphism found in the adult population. The dimorphism anatomy of the supralaryngeal vocal tract and larynx of fossil hominoids can be inferred from their skulls.     

Sexual differentiation and hormonal regulation of the laryngeal synapse in Xenopus laevis

In Xenopus laevis frogs, sex differences in adult laryngeal synapses contribute to sex differences in vocal behavior. This study explores the development of sex differences in types of neuromuscular synapses and the development and hormone regulation of sex differences in transmitter release. Synapses in the juvenile larynx have characteristics not found in adults: juvenile muscle fibers can produce subthreshold or suprathreshold potentials in response to the same strength of nerve stimulation and can also produce multiple spikes to a single nerve stimulus. Juvenile laryngeal muscle also contains the same synapse types (I, II, and III) as are found in adult laryngeal muscle. The distribution of laryngeal synapse types in juveniles is less sexually dimorphic than the distribution in adults. Analysis of quantal content indicates that laryngeal synapses characteristically release low amounts of transmitter prior to sexual differentiation. Quantal content values from male and female juveniles are similar to values for adult males and are lower than values for adult females. When juveniles are gonadectomized and treated with exogenous estrogen, quantal content values increase significantly, suggesting that this hormone may increase transmitter release at laryngeal synapses during development. Gonadectomy alone does not affect quantal content of laryngeal synapses in either sex. Androgen treatment decreases quantal content in juvenile females but not males; the effect is opposite to and smaller than that of estrogen. Thus, muscle fiber responses to nerve stimulation and transmitter release are not sexually dimorphic in juvenile larynges. Transmitter release is strengthened, or feminized, by the administration of estradiol, an ovarian steroid hormone. © 1995 John Wiley & Sons, Inc.     

Androgen directs sexual differentiation of laryngeal innervation in developing Xenopus laevis

In adult Xenopus laevis, innervation of the vocal organ is more robust in males than in females. This sex difference originates during tadpole development; at stage 56, when the gonads first differentiate, the number of axons entering the larynx is the same in the sexes, but by stage 62, innervation is greater in males. To determine if androgen secretion establishes sex differences in axon number, we treated tadpoles with antiandrogen or androgen beginning at stage 48 or 54 and counted laryngeal nerve axons at stage 62 using electron microscopy. When male tadpoles were treated with the antiandrogen hydroxyflutamide, axon numbers were reduced to female-typical values; axon numbers in females were unaffected by antiandrogen treatment. When female tadpoles were treated with the androgen DHT (dihydrotestosterone), axon numbers were increased to male-like values. These findings suggest that endogenous androgen secretion during late tadpole stages in males is required for the sexual differentiation of laryngeal innervation observed from stage 62 on. Because androgen treatment and laryngeal innervation affect myogenesis in postmetamorphic frogs, numbers of laryngeal dilator muscle fibers were determined for hormonally manipulated tadpoles. At stage 62, vehicle-treated males had more laryngeal axons than females; laryngeal muscle fiber numbers did not, however, differ in the sexes. Both male and female tadpoles, treated from stage 54 with DHT, had more muscle fibers at stage 62 than vehicle-treated controls. Thus, while endogenous androgen secretion during late tadpole stages is subthreshold for the establishment of masculinized muscle fiber numbers, laryngeal myogenesis is androgen sensitive at this time and can be increased by suprathreshold provision of exogenous DHT. A subgroup of tadpoles, DHT treated from stage 54 to 62, was allowed to survive, untreated, until postmetamorphic stage 2 (PM2: 5 months after metamorphosis is complete). Androgen treatment between tadpole stages 54 and 62 does not prevent the ontogenetic decrease in axon numbers characteristic of laryngeal development. In addition, the elevation in stage 62 axon numbers produced by DHT-treatment at late tadpole stages was not associated with elevated numbers of laryngeal muscle fibers at PM2. Juvenile males normally maintain elevated axon numbers (relative to final adult values) through PM2 and the presence of these additional axons may result from-rather than contribute directly to—laryngeal muscle fiber addition. 1994 John Wiley & Sons, Inc.     

Sexually distinct development of vocal pathways in Xenopus laevis

Deterministic rules, rather than experience, are thought to regulate the development of simple behaviors in vertebrates and invertebrates. We revisited this issue through examination of the sexually distinct vocalizations of African clawed frogs (Xenopus laevis), a reproductive behavior used by sexually mature males and females. We discovered that, as expected for simple behavior, female vocalizations develop through deterministic rules. The rare calls of juvenile females are indistinguishable from those of adult females. The vocal pathways of juvenile females, as measured by the contractile properties of the laryngeal muscles (the vocal muscles) and the laryngeal motoneuron somata (vocal motoneurons) size, are the developmental default and do not differentiate as they mature. Male Xenopus, in contrast, produce extensive vocalizations with rudimentary acoustic structure before reaching sexual maturity. Moreover, the functional properties of the vocal central pattern generator mature before muscle fibers and motoneuron size are fully masculinized. The results suggest that neuronal activity during development may be important in organizing the contractile properties of the muscle fibers in male, but not in female Xenopus. © 2010 Wiley Periodicals, Inc. Develop Neurobiol 70: 862–874, 2010     

Modelling the production of complex calls in the túngara frog (Physalaemus pustulosus)

Relatively few studies have used experimental manipulations to investigate the mechanics of vocal production in frogs and toads, even though many frogs produce complex signals with multiple components and/or nonlinearities. Modelling approaches can add to empirical studies by illuminating how various components of the vocal system interact to produce communication signals. In this study, we use bond graphs, a lumped-element modelling technique, to explore how the combination of active modulation of vocal production and the passive dynamics of a unique laryngeal structure result in the complex calls produced by males in an anuran model system. The túngara frog (Physalaemus (= Engystomops) pustulosus) produces advertisement calls with a ‘whine’ and a facultative ‘chuck’. Whines are amplitude and frequency modulated. The chuck is characterized by its spectral complexity and appears to contain a period doubling bifurcation resulting in subharmonics. In our model, we focus on how a fibrous mass attached to the vocal cords results in subharmonics in the chuck. Our models suggest that active (neural) modulation of the fibrous mass is not necessary for the transition between the spectral characteristics of the whine and chuck. Rather, it is possible that the vibratory mode of the fibrous mass and thus the vocal cords changes passively as a result of changes in airflow through the system.     

Allometry, bilateral asymmetry and sexual differences in the vocal tract of common eiders Somateria mollissima and king eiders S. spectabilis

Intraspecific sexual differences, high variation, and positive allometry of sexually-selected external display structures are common. Many sexually-selected anatomical specializations occur in the avian vocal tract but intraspecific variation and allometry have been investigated little. The tracheal bulla bulla syringealis occurs in males of most duck species. We quantified variation and size-scaling of the bulla, plus sexual differences in size of trachea, bronchi, and vocal muscles, for 62 common eiders Somateria mollissima and 51 king eiders S. spectabilis. Trends were similar in both species. Bullar ossification and definitive size occurred early in life: bullar size did not differ between first-year and older males. Bullar size did not vary more than size of other body parts (CVs of 3.4–7.0% for bullar length and breadth). Bullar size scaled to body size with negative allometry or isometry. Vocal muscles were 10–50% thicker in males than females, a much greater sexual difference than in body size (CVs of 3–6% on linear body-size variables). Vocal muscles were larger on the left side in both sexes and bilateral asymmetry was slightly more pronounced in males. Low variation and a trend towards negative allometry suggest that bullar size is under stabilizing selection; if bullar size affects vocal attributes of voice, then the latter cannot be condition-dependent. We recommend comparative research on vocal communication, vocal individuality and vocal-tract anatomy and function in eiders and other ducks.     

Sexual Dimorphism of the Zebra Finch Syrinx Indicates Adaptation for High Fundamental Frequencies in Males

Background

In many songbirds the larger vocal repertoire of males is associated with sexual dimorphism of the vocal control centers and muscles of the vocal organ, the syrinx. However, it is largely unknown how these differences are translated into different acoustic behavior.

Methodology/Principal Findings

Here we show that the sound generating structures of the syrinx, the labia and the associated cartilaginous framework, also display sexual dimorphism. One of the bronchial half rings that position and tense the labia is larger in males, and the size and shape of the labia differ between males and females. The functional consequences of these differences were explored by denervating syringeal muscles. After denervation, both sexes produced equally low fundamental frequencies, but the driving pressure generally increased and was higher in males. Denervation strongly affected the relationship between driving pressure and fundamental frequency.

Conclusions/Significance

The syringeal modifications in the male syrinx, in concert with dimorphisms in neural control and muscle mass, are most likely the foundation for the potential to generate an enhanced frequency range. Sexually dimorphic vocal behavior therefore arises from finely tuned modifications at every level of the motor cascade. This sexual dimorphism in frequency control illustrates a significant evolutionary step towards increased vocal complexity in birds.     

Neuroeffectors for vocalization in Xenopus laevis: hormonal regulation of sexual dimorphism

South African clawed frogs use sex-specific vocalizations during courtship. In the male, vocalizations are under the control of gonadal androgen. Though females have moderate levels of circulating androgen, they do not give male-typical mate calls. Both muscles of the vocal organ and neurons of the central nervous system (CNS) vocal pathway are sexually dimorphic and androgen-sensitive. Recent studies suggest that the failure of androgen to masculinize adult females results from a male-specific, androgen-regulated developmental program. At metamorphosis the larynx is sexually monomorphic and feminine in morphology, muscle fiber number and androgen receptor content. During the next six months, under the influence of increasing androgen titers and high receptor levels, myoblasts proliferate in the male and muscle fibers increase at an average rate of 100/day. Females have much lower hormone levels, receptor values decline and they display no net addition of fibers. At metamorphosis, both males and females have approximately 4000 muscle fibers. By adulthood, males have eight times the female fiber number. In the CNS, adult laryngeal motor neurons are more numerous with larger somata and dendritic trees in males than in females. Certain connections of neurons in the vocal pathway are also less robust in females. Unlike the periphery, motor neuron number does not appear to be established by androgen-induced proliferation. Our current hypothesis is that androgen acts at the level of laryngeal muscle to produce more muscle fibers and thus provide more target for motor neurons in the male. This process could regulate cell number by ontogenetic cell death. In the CNS, androgen-target neurons become capable of accumulating hormone shortly before metamorphosis. Androgen receptor in laryngeal motor neurons may permit the dendritic growth characteristic of males by increasing sensitivity to afferent stimuli. Such a process could account for the observed differences in CNS vocal "circuitry" in X. laevis and thus behavioral differences between the sexes.     

Different patterns of cytokeratin expression in the normal epithelia of the upper respiratory tract

The distribution and type of cytokeratins present in the normal human epithelia of the nasopharynx, oropharynx, tongue, palatine tonsil, epiglottis, vocal cord, and laryngeal ventricle were studied using immunohistochemical techniques and by gel electrophoresis of cytoskeletal proteins microdissected from frozen tissues. Noncornifying stratified epithelia covering the oropharynx, tongue, surface of the palatine tonsil, pharyngeal surface of the epiglottis, and vocal cord were all found to contain cytokeratins nos. 4, 5, 6, 13, 14, and 15, together with minor amounts of cytokeratin no. 19, i.e., a pattern similar to that previously reported for esophageal epithelium. The immunohistochemical reaction with KA4, an antibody specific for cytokeratins nos. 14, 15, 16, and 19, revealed reactivity confined to the basal epithelial cells of the tongue, oropharynx, pharyngeal epiglottis, and two out of five samples of vocal cords. This same antibody reacted with the entire thickness of three out of the five true vocal cords which were shown by gel electrophoresis to also contain cytokeratins nos. 16 and 17. Gel electrophoresis revealed that the pseudostratified columnar epithelium covering the laryngeal ventricle was more complex, in that it contained cytokeratins nos. 5, 13, 14, 15, and 17, which are typical of stratified epithelia, as well as cytokeratins nos. 7, 8, 18, and 19, which are characteristic of simple epithelia. This pattern is similar to that found in bronchial epithelium. The laryngeal surface of the epiglottis exhibited cytokeratins nos. 4, 5, 7, 8, 13, 14, 15, 17, 18, and 19, i.e., a pattern combining features of both esophageal- and bronchial-type epithelia. The reaction of these epithelia containing columnar cells with antibody RGE-53, which is specific for cytokeratin no. 18, revealed a staining reaction confined to the superficial columnar cells, whereas KA1 stained only the basal cells of these epithelia. The results of our study make it possible to distinguish two types of noncornifying stratified squamous epithelium, namely the 'esophageal type' which covers the tongue, oropharynx, and pharyngeal surface of the epiglottis, and another type which overlies the vocal cords and the transitional zone between the pharyngeal and laryngeal surfaces of the epiglottis. Furthermore, there appear to be variants of pseudostratified columnar epithelium, i.e., the usual bronchial type lining the laryngeal ventricle, and a type with a thicker subcolumnar cell compartment that is found on the laryngeal surface of the epiglottis. The patterns of expression of cytokeratins in the respiratory tract are compared with those of other epithelia.     

Comparative analyses of age- and sex-specific patterns of vocal behaviour in four species of Old World monkeys

Sound recordings and behavioural data were collected from four primate species of two genera (Macaca, Presbytis). Comparative analyses of structural and behavioural aspects of vocal communication revealed a high degree of intrageneric similarity but striking intergeneric differences. In the two macaque species (Macaca silenus, Macaca radiata), males and females shared the major part of the repertoire. In contrast, in the two langurs (Presbytis johnii, Presbytis entellus), many calls were exclusive to adult males. Striking differences between both species groups occurred with respect to age-specific patterns of vocal behaviour. The diversity of vocal behaviour was assessed from the number of different calls used and the proportion of each call in relation to total vocal output for a given age/sex class. In Macaca, diversity decreases with the age of the vocalizer, whereas in Presbytis the age of the vocalizer and the diversity of vocal behaviour are positively correlated. A comparison of the data of the two genera does not suggest any causal relationship between group composition (e.g. multi-male vs. one-male group) and communication system. Within each genus, interspecific differences in vocal behaviour can be explained by differences in social behaviour (e.g. group cohesion, intergroup relation, mating behaviour) and functional disparities. Possible factors responsible for the pronounced intergeneric differences in vocal behaviour between Macaca and Presbytis are discussed.     

Syringeal anatomy and allometry in murres (Alcidae: Uria)

Species and sexual differences in vocalizations and the vocal tract are widespread. We studied the vocal tract of Common (Uria aalge) and Thick-billed (U. lomvia) Murres. We predicted anatomical or allometric differences in adults between species and sexes due to vocal differences related to social behavior (Common Murres nest at higher densities; males are the more aggressive sex, etc.). The vocal tract was anatomically simple and similar between species and sexes. The trachea was mainly cartilage, but the posteriormost 4–6 tracheal rings were calcified and fused as the tympanum, as part of the syrinx. The syrinx included the (unfused) first bronchial semirings, which were enlarged and calcified beneath the insertion of M. tracheolateralis. Weak bilateral asymmetry (right side larger) occurred in widths of M. tracheolateralis and M. sternotrachealis. The trachea was ∼10% longer in Common Murres; the tympanum and M. sternotrachealis width were relatively larger in Thick-billed Murres. Vocal-tract morphology and size did not differ between the sexes in either species. In allometric analyses on adults, isometry characterized (1) tympanum size in relation to head + bill length, and to humerus length (respectively), in Thick-billed Murres, and (2) M. sternotrachealis width in relation to tarsometatarsal length in both species. Comparative field studies on species and sexes are needed to clarify the functional significance of our findings in relation to vocalization.     

The sexually dimorphic larynx of Xenopus laevis: development and androgen regulation

The aims of this study were to characterize sexual dimorphism in the larynx of adult Xenopus laevis and to determine how sex differences arise during postmetamorphic development. The larger male larynx is a result of greater cell numbers in both cartilage and muscle. The dilator laryngis muscle of the male larynx has 6-7 times more muscle fibers than that of the female. At metamorphosis, the larynx is sexually monomorphic and feminine in phenotype. The DNA content of the male larynx doubles during the first 6 months following metamorphosis; there is no net DNA increase in the female larynx during this time. Both sexes experience a marked increase in laryngeal DNA content and mass between 6 months and adulthood. The number of muscle fibers in the male larynx increases at an average rate of 150 fibers a day during the first 10 months of postmetamorphic development. There is no net change in fiber numbers in the female larynx from metamorphosis to adulthood. Administration of the antiandrogen Flutamide to metamorphic frogs prevents the net addition of laryngeal muscle fibers in males. Thus, we propose that addition of postmetamorphic laryngeal muscle fibers in males is dependent upon the presence of circulating androgens. Exogenous testosterone administration results in an increase in laryngeal mass, DNA content, and cellular proliferation in juvenile frogs. Using [3H]thymidine injections to probe ongoing, as well as testosterone-induced, cell proliferation, we conclude that cellular proliferation is regulated differently in males and females during development. Thus androgen-induced proliferation is one cellular mechanism responsible for the sexual dimorphism observed in adults.     

Adult plasticity in hormone-sensitive motoneuron morphology: methodological/behavioral confounds

Changes in androgen levels can alter the structure of motoneurons in the spinal nucleus of the bulbocavernosus (SNB), a motor nucleus that innervates perineal muscles involved in copulatory behavior. While sexual activity can alter androgen levels in normal males, it has no effect on SNB motoneuron soma size or dendritic morphology (Beversdorf, Kurz, and Sengelaub, 1990). However, Breedlove (1997) reported reductions in the size of SNB somata, nuclei, and target muscles of copulating versus noncopulating castrated rats maintained on subphysiological testosterone. To reconcile the results obtained using intact versus implant paradigms, we tested the hypothesis that the implant/behavior paradigm could produce differences in hormone levels, potentially confounding sexual behavior effects on the morphology of this androgen-sensitive neuromuscular system. Young adult male rats were castrated and immediately given 5-mm Silastic implants containing crystalline testosterone. One week later, blood samples were drawn and the males were housed with receptive females (copulators) or nonreceptive females (noncopulators) or housed alone (singles). After 27 days, blood samples were drawn again, and SNB target muscles and spinal cords removed. No differences in target muscle weight or SNB somata and nuclei size were observed between copulators, noncopulators, or singles; as expected, all measures were significantly reduced relative to intact males. Radioimmunoassay showed that testosterone declined differentially over the course of the behavioral manipulation across groups, being greatest in copulators and least pronounced in single males. These data indicate that differences in sexual or housing experience can alter testosterone titers under these implant conditions, potentially confounding hormone-sensitive measures of morphology.     

Sex Differences and Age Gradations in Vocalizations of Japanese and Lion-tailed Monkeys (Macaca fuscata and Macaca silenus)

SYNOPSIS. Vocal repertoires of male and female non-human primatescan differ by presenceor absence of sounds, acoustic form, andfunction or usage. Examples of sex-specificity in vocalizationsare taken from field studies of two macaque species that differin social organizationand habitat. They are used to illustratesex-specific acoustic patterns and vocalizations thatare employedin socially different fashion by males and females. Some vocalsex differences develop from sounds shared in common by younganimals of both sexes while others arise late in life by anacoustic pattern suddenly emerging only in males. Differencesmay be due to sexual selection, social suppression, morphologicalfeatures, or because vocal sounds are linked to circumstancesexperienced by only male or female adults. The relation betweensocial usage of some vocalizations by adults and youngstersaffords insights into their signal value and into the underlyingstate of the vocalizing animal.     

Apneic laryngeal chemoreflex and naloxone

The apneic laryngeal chemoreflex (QRL), elicited by water on the vocal cords and by mechanical stimulation (MRL) has been compared in dog. Both stimuli cause apnea, bradycardia, hypotension and constriction of the glottis. In QRL apnea predominates white in MRL bradycardia is more intense. All the components of the response decrease with naloxone at a 400 micrograms X kg-1 doses, suggesting that the reflex inhibition depends on endogenous opioids located in the respiratory centre.     

Respiratory activities of intralaryngeal branches of the recurrent laryngeal nerve

To distinguish experimentally between motor nerve activity destined for vocal cord abductor muscles and that bound for muscles that adduct the cords, we recorded efferent activities of intralaryngeal branches of the recurrent laryngeal nerve (RLN) in decerebrate, vagotomized, paralyzed, ventilated cats. Activities of the whole RLN and phrenic nerve were also recorded. Nerve activities were assessed at several steady-state end-tidal O2 and CO2 concentrations. The nerve to the thyroarytenoid (TA) muscle, a vocal cord adductor, was only slightly active under base-line (normocapnic, hyperoxic) conditions but in most cats developed strong activity during expiration in hypocapnia or hypoxia. In severe hypocapnia, phasic expiratory TA activity persisted even during phrenic apnea, indicating continuing activity of the respiratory rhythm generator. The nerve to the posterior cricoarytenoid (PCA) muscle, the vocal cord abductor, was always active in inspiration but often showed expiratory activity as well. This expiratory activity was usually enhanced by hypercapnia and often inhibited by hypoxia. The results are consistent with previous electromyographic findings and emphasize the importance of distinguishing abductor from adductor activity in studies of laryngeal control.