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PubMed2005

Historical perspective: Hormonal regulation of behaviors in amphibians.

This review focuses on research into the hormonal control of behaviors in amphibians that was conducted prior to the 21st century. Most advances in this field come from studies of a limited number of species and investigations into the hormonal mechanisms that regulate reproductive behaviors in male frogs and salamanders. From this earlier research, we highlight five main generalizations or conclusions. (1) Based on studies of vocalization behaviors in anurans, testicular androgens induce developmental changes in cartilage and muscles fibers in the larynx and thereby masculinize peripheral structures that influence the properties of advertisement calls by males. (2) Gonadal steroid hormones act to enhance reproductive behaviors in adult amphibians, but causal relationships are not as well established in amphibians as in birds and mammals. Research into the relationships between testicular androgens and male behaviors, mainly using castration/steroid treatment studies, generally supports the conclusion that androgens are necessary but not sufficient to enhance male behaviors. (3) Prolactin acts synergistically with androgens and induces reproductive development, sexual behaviors, and pheromone production. This interaction between prolactin and gonadal steroids helps to explain why androgens alone sometimes fail to stimulate amphibian behaviors. (4) Vasotocin also plays an important role and enhances specific types of behaviors in amphibians (frog calling, receptivity in female frogs, amplectic clasping in newts, and non-clasping courtship behaviors). Gonadal steroids typically act to maintain behavioral responses to vasotocin. Vasotocin modulates behavioral responses, at least in part, by acting within the brain on sensory pathways that detect sexual stimuli and on motor pathways that control behavioral responses. (5) Corticosterone acts as a potent and rapid suppressor of reproductive behaviors during periods of acute stress. These rapid stress-induced changes in behaviors use non-genomic mechanisms and membrane-associated corticosterone receptors.

Amphibians↗
PubMed2005

Intra-sexual variation in male reproduction in teleost fish: a comparative approach.

The occurrence of intra-sexual variation in reproduction is a widespread phenomenon in teleosts. One such form of variation consists in the occurrence of alternative male types: males that invest resources in mate attraction and males that exploit the investment of the former males, by trying to sneak fertilizations during spawning. These alternative reproductive tactics can be classified according to their plasticity during the life span of the individuals (i.e., fixed vs. sequential vs. reversible). Furthermore, the differences between morphs within a given species may involve a set of different traits, including reproductive behavior, the differentiation of male morphological traits, and the patterns of gonad tissue allocation and the differentiation of gonadal accessory glands. In this paper, we review the available data on four species exhibiting different types of intra-sexual plasticity in reproduction that have been studied in our lab. The data on the proximate mechanisms, androgens and forebrain arginine-vasotocin (AVT), underlying these alternative tactics suggest that between-morph differences in androgen levels, especially in 11-ketotestosterone, are especially present in species where the alternative male types have evolved morphological traits that are tactic-specific (i.e., sexual polymorphisms) and that differences in AVT appear to be related to between-morph differences in the expression of courtship behavior. Therefore, this comparative approach leads us to propose that the different endocrine systems are involved in the differentiation of different sets of traits that make up alternative phenotypes, and that the differentiation of alternative tactics is not controlled by a single endocrine system (e.g., androgens).

Adaptation, Physiological↗
PubMed2005

Testosterone increases display behaviors but does not stimulate growth of adult plumage in male golden-collared manakins (Manacus vitellinus).

In order to attract females, male golden-collared manakins gather in leks and perform a complex display consisting of acrobatics accompanied by loud "wingsnapping". During this display, males show off their yellow beard and yellow, black, and green plumage that is striking in comparison to the dull green plumage of young males and females. We investigated the role of testosterone (T) in activating the display of manakins and in stimulating the growth of the adult male plumage. T regulates song, copulation, and territorial aggression in temperate species. In tropical species, however, T levels can be relatively low year round, which has raised questions about the involvement of T in courtship display and male aggression in these species. In neither temperate nor tropical species has the role of hormones in the shift from juvenile to adult plumage been well studied. Therefore, we implanted green-plumaged birds and adult males with either a T pellet or an inert pellet (controls) and observed the display behaviors of these birds in the field and in captivity. In captive birds, we also plucked feathers from sexually dimorphic regions and observed color and regeneration rate of new feathers. We found that birds implanted with T increased several display behaviors compared to controls. All plucked feathers grew back the same color as prior to treatment; however, we observed some differences in feather growth rate between T-treated birds and controls.

Animals↗
PubMed2006

Elevated 11-ketotestosterone during paternal behavior in the Bluebanded goby (Lythrypnus dalli).

The relationship between androgens and paternal behavior is not straightforward, potentially because of the diversity of tasks a male must undertake to maximize reproductive success, notably alternating between courtship, aggression, and offspring care. In some species, these events are separated in time, but in others they are coincident. The endocrine profiles of species that simultaneously court, parent, and defend a nest, such as male bluebanded gobies (Lythrypnus dalli), are not well understood. We sampled a potent fish androgen, 11-ketotestosterone (KT), at different life history stages (experienced parenting males, experienced males not actively parenting, inexperienced males with their first clutch, and females), to examine this relationship. We found that experienced parenting L. dalli males have the highest KT levels of any group, while none of the other groups differed significantly. Males showed elevated KT levels when they had eggs compared to when they did not. Our data suggest that KT facilitates at least some aspects of parental care in L. dalli.

Animals↗
PubMed2006

Population divergence in plasticity of the AVT system and its association with aggressive behaviors in a Death Valley pupfish.

Behavioral differences can evolve rapidly in allopatry, but little is known about the neural bases of such changes. Allopatric populations of Amargosa pupfish (Cyprinodon nevadensis) vary in aggression and courtship behaviors in the wild. Two of these wild populations were recently found to differ in brain expression of arginine vasotocin (AVT)--a peptide hormone shown previously to modulate aggression in pupfish. These populations have been isolated for less than 4000 years, so it remained unclear whether the differences in behavior and neural AVT phenotype were evolved changes or plastic responses to ecologically dissimilar habitats. Here, I tested whether these population differences have a genetic basis by examining how aggressive behavior and neural AVT phenotype responded to ecologically relevant variation in salinity (0.4 ppt or 3 ppt) and temperature (stable or daily fluctuating). Pupfish from Big Spring were more aggressive than pupfish from the Amargosa River when bred and reared under common laboratory conditions. Morphometric analysis of preoptic AVT immunoreactivity showed that the populations differed in how the AVT system responded to salinity and temperature conditions, and revealed that this plasticity differed between parvocellular and magnocellular AVT neuron groups. Both populations also showed relationships between neural AVT phenotype and aggression in the rearing environment, although populations differed in how aggression related to variation in magnocellular AVT neuron size. Together, these results demonstrate that the pupfish populations have diverged in how physical and social conditions affect the AVT system, and provide evidence that the AVT system can evolve quickly to ecologically dissimilar environments.

Aggression↗
PubMed2006

Effects of artificial social stimuli on the reproductive schedule and hormone levels of yellow-eyed penguins (Megadyptes antipodes).

The effects of social stimuli on avian reproductive behaviors such as breeding schedules and courtship behaviors are well known due to numerous field studies. However, studies that have simultaneously examined the effects of social stimuli on reproductive behavior and the mediating endocrine mechanisms have been largely restricted to captive populations, which may not be representative of free-living populations. This study, conducted over two breeding seasons, aimed to simultaneously measure the effects of experimentally increasing auditory stimuli on the breeding schedule and endocrinology (levels of total androgen, estradiol, progesterone and prolactin) on free-living yellow-eyed penguins (Megadyptes antipodes). The yellow-eyed penguin is the least colonial of all penguins, nesting far apart from each other under dense vegetation, and, therefore, is presumed to experience much lower levels of social stimuli than other penguins. Egg laying was significantly more synchronous and tended to be earlier when birds were exposed to playbacks of the calls of conspecifics in 1 year of the study. We also found that levels of total androgen and estradiol of males in 1 year, and prolactin in another year, were proportionally higher among treated birds compared control birds that received no artificial auditory stimuli. These results show that even among supposedly solitary nesters, social stimuli could still play a role in influencing reproductive behavior and physiology. For the first time in free-living seabirds, we have demonstrated that behavioral responses to increased social stimuli are associated with hormonal changes.

Animals↗
PubMed2004

Phylogeny of the orchid bees (Hymenoptera: Apinae: Euglossini): DNA and morphology yield equivalent patterns.

Orchid bees (Euglossini) are spectacular long-tongued Neotropical bees important in the pollination of Neotropical long-corolla flowers, particularly some orchids. Besides remarkably long tongues, males in particular exhibit other flower-related adaptations, including setal brushes on the foretarsi used for rasping the petals of orchids while collecting aromatic compounds. These compounds are stored in large swollen tibiae and are thought to play an important role in courtship behavior. Euglossini are also unusual in lacking sociality; they are the only tribe among the corbiculate bees that are not eusocial, and two of the genera are cleptoparasitic. Each genus exhibits distinct behavioral traits including nest architecture and host-parasite interactions, yet their evolution is unknown. Despite previous phylogenetic studies of on morphological characters, the relationships among the five euglossine genera remain under debate. We investigate euglossine generic relationships using DNA sequence data from four genes and new morphological characters. The morphological and molecular data yield congruent evolutionary patterns, and combining the data gives a fully resolved and well supported phylogeny of Euglossini.

Animals↗
PubMed2005

Phylogeny of Ensifera (Hexapoda: Orthoptera) using three ribosomal loci, with implications for the evolution of acoustic communication.

Representatives of the Orthopteran suborder Ensifera (crickets, katydids, and related insects) are well known for acoustic signals produced in the contexts of courtship and mate recognition. We present a phylogenetic estimate of Ensifera for a sample of 51 taxonomically diverse exemplars, using sequences from 18S, 28S, and 16S rRNA. The results support a monophyletic Ensifera, monophyly of most ensiferan families, and the superfamily Gryllacridoidea which would include Stenopelmatidae, Anostostomatidae, Gryllacrididae, and Lezina. Schizodactylidae was recovered as the sister lineage to Grylloidea, and both Rhaphidophoridae and Tettigoniidae were found to be more closely related to Grylloidea than has been suggested by prior studies. The ambidextrously stridulating haglid Cyphoderris was found to be basal (or sister) to a clade that contains both Grylloidea and Tettigoniidae. Tree comparison tests with the concatenated molecular data found our phylogeny to be significantly better at explaining our data than three recent phylogenetic hypotheses based on morphological characters. A high degree of conflict exists between the molecular and morphological data, possibly indicating that much homoplasy is present in Ensifera, particularly in acoustic structures. In contrast to prior evolutionary hypotheses based on most parsimonious ancestral state reconstructions, we propose that tegminal stridulation and tibial tympana are ancestral to Ensifera and were lost multiple times, especially within the Gryllidae.

Animals↗
PubMed2004

Acoustic communication in Okanagana rimosa (Say) (Homoptera: Cicadidae).

The cicada Okanagana rimosa (Say) has an acoustic communication system with three types of loud timbal sounds: (i) A calling song lasting several seconds to about 1 min which consists of a sequence of chirps at a repetition rate of 83 chirps per second. Each chirp of about 6 ms duration contains 4-5 pulses. The sound level of the calling song is 87-90 dB SPL at a distance of 15 cm. (ii) An amplitude modulated courtship song with increasing amplitude and repetition rate of chirps and pulses. (iii) A protest squawk with irregular chirp and pulse structure. The spectra of all three types are similar and show main energy peaks at 8-10 kHz. Only males sing, and calling song production is influenced by the songs of other males, resulting in an almost continuous sound in dense populations. In such populations, the calling songs overlap and the temporal structure of individual songs is obscured within the habitat. The calling song of the broadly sympatric, closely related species O. canadensis (Provander) is similar in frequency content, but distinct in the temporal pattern (24 chirps per second, 24 ms chirp duration, eight pulses per chirp) which is likely important for species separation in sympatric populations. The hearing threshold of the auditory nerve is similar for females and males of O. rimosa and most sensitive at 4-5 kHz. Experiments in the field show that female phonotaxis of O. rimosa depends on parameters of the calling song. Most females are attracted to calling song models with a 9 kHz carrier frequency (peak frequency of the calling song), but not to models with a 5 kHz carrier frequency (minimum hearing threshold). Phonotaxis depends on temporal parameters of the conspecific song, especially chirp repetition rate. Calling song production is influenced by environmental factors, and likelihood to sing increases with temperature and brightness of the sky. Correspondingly, females perform phonotaxis most often during sunny conditions with temperatures above 22 degrees C. Non-mated and mated females are attracted by the acoustic signals, and the percentage of mated females performing phonotaxis increases during the season.

Journal Article↗
PubMed2002

Pheromonal activation of vomeronasal neurons in plethodontid salamanders.

Pheromones from the mental glands of male plethodontid salamanders increase sexual receptivity in conspecific females. The pheromone enters the vomeronasal organ during courtship to produce this effect. Vomeronasal neurons from female Plethodon shermani were examined following exposure to male pheromone or saline placed on the nares. Agmatine was used in conjunction with the pheromone to enable immunocytochemical visualization of chemosensory neurons that were activated by the pheromone. Olfactory neurons exposed to pheromone or saline, and vomeronasal neurons exposed to saline did not demonstrate significant labeling. A population of vomeronasal neurons was intensely labeled following exposure to the pheromone. This study suggests that a specific population of vomeronasal neurons in a female plethodontid salamander is responsible for transmitting pheromonal information to the brain to produce modifications in behavior.

Animals↗
PubMed1999

Testosterone increases the number of substance P-like immunoreactive neurons in a specific sub-division of the lateral hypothalamus of the weakly electric, brown ghost knifefish, Apteronotus leptorhynchus.

During spawning, male and female brown ghost knife fish modulate their electric organ discharge to produce discrete courtship signals known as chirps. However, non-spawning fish show clear sex differences in chirp responsiveness to electrosensory stimuli; males consistently chirp, whereas females do not. This behavioral dimorphism is paralleled by sex differences in substance P-like immunoreactivity (SPl-ir) in the prepacemaker nucleus (PPn) which regulates chirping. The PPn is densely innervated by SPl-ir fibers in males, but not in females. However, we have shown that both female chirping behavior and the expression of SPl-ir in the PPn are enhanced following adult testosterone treatment [J.G. Dulka, L. Maler, W. Ellis, Androgen-induced changes in electrocommunicatory behavior are correlated with changes in substance P-like immunoreactivity in the brain of the electric fish, Apteronotus leptorhynchus, J. Neurosci. 15 (1995) 1879-1890]. Thus, testosterone-induced changes in SPl-ir input to the PPn may modulate female chirping during spawning. Recent evidence suggests that SPl-ir projections to the PPn may originate from SPl-ir neurons in the lateral hypothalamus (Hl). If so, one might expect to see a greater number of SPl-ir neurons in the Hl of testosterone-implanted females compared to Blank-implanted controls. In making this comparison, we found that both groups of females had similar numbers of SPl-ir neurons in the anterior Hl, but that testosterone-implanted females had significantly (p<0.01) more SPl-ir neurons in a distinct part of the posterior Hl. This sub-population of testosterone-sensitive, SPl-ir neurons may contribute to the regulation of chirping, since an increase in their number is positively correlated with the appearance of SPl-ir fibers in the PPn [J.G. Dulka, L. Maler, W. Ellis, Androgen-induced changes in electrocommunicatory behavior are correlated with changes in substance P-like immunoreactivity in the brain of the electric fish, Apteronotus leptorhynchus, J. Neurosci. 15 (1995) 1879-1890].

Animals↗
PubMed2003

Plasma steroid and nutrient levels during the active season in wild Testudo horsfieldi.

Plasma concentrations of sex steroids (testosterone and progesterone), proteins (total protein and albumin), lipids (phospholipids, cholesterol, and triglycerides), and minerals (calcium and phosphorus) were measured in wild Testudo horsfieldi in Uzbekistan, during the short, 3-month activity period (March-May, 1998). Testosterone concentration in males was highest (52 ng/ml) when they had just emerged from brumation (hibernation) in mid-March, which was also the beginning of the mating period, and fell in April. In females, progesterone peaked in mid-April (at 10 ng/ml), just before ovulation of the first clutches at the end of April and beginning of May. Testosterone levels in females and progesterone levels in males were low (<3 ng/ml) throughout the activity period. In general, the plasma concentrations of proteins, lipids, and phosphorus increased slowly in males, but more rapidly in females, during the activity season. These increases were particularly strong in females in the second half of April, coinciding with the peaks in female hormone levels. The changes in plasma hormones and nutrients reflected the timing of the different behaviours. The four first weeks of above-ground activity (mid-March to mid-April) by males, when they had high testosterone levels, were primarily allocated to fighting other males, courtship, and mating, while females spent much of that time feeding. Thereafter, both sexes concentrated on feeding. Females were probably preparing to ovulate in late-April, when their progesterone levels were highest and when plasma nutrient levels increased considerably.

Animals↗
PubMed2003

Is a wild mammal kept and reared in captivity still a wild animal?

This study compared domestic guinea pigs (Cavia aperea f. porcellus; DGP) and two different populations of the wild cavy (Cavia aperea), its ancestor, to examine whether rearing of wild mammals in captivity affects their behavior and physiological stress responses. One population of wild cavies consisted of wild-trapped animals and their first laboratory-reared offspring (WGP-1). The animals of the other population were reared in captivity for about 30 generations (WGP-30). The spontaneous behavior of each of six groups of WGP-1 and WGP-30 and nine groups of DGP, each consisting of one adult male and two adult females, was analyzed quantitatively. Blood samples of the males were taken to determine cortisol, epinephrine, and norepinephrine concentrations. In addition, the exploratory behavior of 60-day-old male WGP-1, WGP-30, and DGP was investigated in an exploration apparatus. The domesticated animals displayed significantly less aggression, but significantly more sociopositive and male courtship behavior than their wild ancestors. In addition, DGP were much less attentive to their physical environment. Surprisingly, no behavioral difference was found between WGP-1 and WGP-30. Basal cortisol concentrations did not differ between wild and domestic guinea pigs. Catecholamine concentrations, however, as well as the challenge values of cortisol, were distinctly reduced in the DGP. WGP-1 and WGP-30 did not differ with respect to their endocrine stress responses. In the exploration apparatus both forms of wild cavies were much more explorative than the domestic animals. These data suggest that the long-term breeding and rearing of wild guinea pigs in captivity do not result in significant changes in behavior and hormonal stress responses. It appears to take much longer periods of time and artificial selection by humans to bring about characters of domestication in wild animals.

Aggression↗
PubMed2003

Rapid corticosterone-induced impairment of amplectic clasping occurs in the spinal cord of roughskin newts (taricha granulosa).

Courtship clasping, a reproductive behavior in male roughskin newts (Taricha granulosa), is rapidly blocked by an action of corticosterone (CORT) at a specific neuronal membrane receptor. The CORT-induced impairment of clasping in behaving newts appears to be mediated partly by an elimination of clasping-related activity in medullary reticulospinal neurons. Previous studies of rapid CORT actions in Taricha have focused on the brain, so existence of CORT action in the spinal cord or peripheral nervous system has not been assessed. The present study used newts with a high cervical spinal transection to examine potential spinal or peripheral CORT effects on clasping by the hindlimbs in response to pressure on the cloaca. Spinal transection causes clasps elicited by cloacal stimulation to be very sustained beyond the termination of the eliciting stimulus. In spinally transected newts, CORT caused a dose-dependent depression in the duration as well as quality of the clasp that appeared within 10 min of injection. CORT selectively impaired the usual sustained maintenance of a clasp after termination of cloacal stimulation, but not clasp elicitation during stimulation. These effects were not produced by dexamethasone, a synthetic glucocorticoid that binds poorly to the CORT membrane receptor. The CORT effect on clasp maintenance but not clasp elicitation implies selective action on an intraspinal generator for clasping but not on sensory or efferent neuromuscular aspects of the response. These results indicate the presence in the newt spinal cord of the CORT membrane receptor that exerts functional effects distinctly different from those on the brainstem.

Animals↗
PubMed2003

Endocrine correlates of sexual behavior in the Mohor gazelle (Gazella dama mhorr).

In this study, we quantitatively examined male sexual behavior in relation to fecal estrogen and progesterone concentrations in female Mohor gazelles. We investigated the hypothesis that, during natural mating, males detect cues relating to the potential for successful conception and pregnancy. Time series analysis revealed that males could detect the approach of estrus 2-3 days before female fecal estrogens and estrogen/progestagen (E/P) ratio reached their peak values. Males also paid closer attention to those females excreting higher fecal estrogen concentrations. Mounting and copulation frequencies were positively correlated with both peri-ovulatory fecal estrogen concentrations, and the frequency of pre-copulatory courtship behaviors. These data suggested that males invest their reproductive effort selectively by mating the most fertile females, assuming that estrogen is a valid index of fertility. This assumption was investigated by examining sequential phases of the reproductive cycle for evidence that oocytes and follicles produced in a more estrogenic environment would lead to the formation of the most competent corpora lutea, thereby maximizing the chance of sustaining pregnancy. Associations between sexual behavior and hormone excretion support the hypothesis that males may use this mechanism to assess female fertility.

Animals↗
PubMed2000

Brain control of mating behavior in the male cricket Gryllus bimaculatus DeGeer: brain neurons responsible for inhibition of copulation actions.

We localized brain neurons responsible for the inhibition of copulation actions in the male cricket, Gryllus bimaculatus. Males with one connective-cut between the brain and the terminal abdominal ganglion (TAG) were able to perform normal reproductive behaviors such as courtship, copulation and spermatophore protrusion. However, copulation response tests using a female dummy showed that they failed to exhibit a transient inhibitory state after application of noxious stimulation in the mating stage and to exhibit a tonic inhibitory state in the post-copulatory sexual refractory stage. A partial cut of a single connective (hemilateral connective-cut) combined with the copulation response test and an axonal backfilling demonstrated that approximately 40 brain neurons (per one connective), whose somata were located in the posterior region of the protocerebrum and mainly contralaterally to their descending axons running through the dorso-medial part of the connective, were candidates for the inhibition of copulation actions, at least in the mating stage. In addition, we report here for the first time, that copulation actions can be elicited shortly after copulation even in intact males. This unusual behavior was frequently observed in fledglings after the final molt. The results are discussed from the perspective of the operation of the brain inhibition system on the central pattern generator (CPG) for copulation.

Journal Article↗
PubMed1991

Sexual isolation in Drosophila. I. Theoretical models for multiple-choice experiments.

Several simple models are developed to calculate expected mating frequencies in ethological isolation experiments. They take into account the effect that the peculiar sexual behavior of Drosophila species can have in multiple-choice experiments. These models depend on only three basic parameters: male competitive ability (C), female receptivity (R) and the coefficient of females acceptance (A). Two types of model can be distinguished: (1) models with discrete preferences, in which A is a measure of the percentage of females accepting a particular kind of male and (2) models with continuous preferences, in which A represents the probability of acceptance for each courtship. It is demonstrated that the information rendered by just one experiment, although effective for determining whether sexual isolation exists, it is insufficient to estimate its degree or to demonstrate that it is asymmetrical. Further developments of the models under more complex conditions as well as their implications for reinforcement and founder effect theories are discussed.

Animals↗
PubMed2003

Pregnant female guinea pigs adapt easily to a new social environment irrespective of their rearing conditions.

For male guinea pigs, the social rearing conditions are crucial for the course of social interactions and endocrine stress responses in later life. While colony-reared males adjust easily to new social situations, high levels of aggression and extreme endocrine responses are found in males that were raised in pairs [Psychoneuroendocrinology 23 (1998) 891]. This study investigated whether the same finding also applies to females. Therefore, 8 females reared in mixed-sexed colonies of 8-10 males and 12-13 females (designated as CF) and 8 females reared in mixed-sexed pairs (designated as PF) were transferred to an unknown colony for 3 days. Control females remained in their respective housing conditions. Serum cortisol concentrations (CORT) were determined before, during and after the transfer. The behavior of the CF and PF was recorded on the first, second and third day of transfer. All females adjusted easily to the new social situation, but frequencies of some behavioral patterns were significantly different between CF and PF. In the new social situation, PF displayed social orientation as well as defensive aggressive and avoidance behavior more frequently than CF. On the day of transfer, resident males displayed courtship behavior more frequently towards PF than CF. CORT did not differ between PF and CF either before, during or after the transfer. Furthermore, the transfer to the new social situation did not result in significantly increased CORT 4 h later either in CF or PF. We conclude that-in contrast to male guinea pigs-females are able to adapt to unfamiliar conspecifics independently of their social rearing conditions.

Adaptation, Psychological↗