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The role of mating preferences in shaping interspecific divergence in mating signals in vertebrates.

Vertebrates represent one of the best-studied groups in terms of the role that mating preferences have played in the evolution of exaggerated secondary sexual characters and mating behaviours within species. Vertebrate species however, also exhibit enormous interspecific diversity in features of mating signals that has potentially led to reproductive isolation and speciation in many groups. The role that sexual selection has played in interspecific divergence in mating signals has been less fully explored. This review summarizes our current knowledge of how mating preferences within species have shaped interspecific divergence in mate recognition signals among the major vertebrate groups. Certain signal modalities appear to characterize mating signal diversification among different vertebrate taxa. Acoustic signals play an important role in mating decisions in anuran amphibians and birds. Here, different properties of the signal may convey information regarding individual, neighbor and species recognition. Mating preferences for particular features of the acoustic signal have led to interspecific divergence in calls and songs. Divergence in morphological traits such as colouration or ornamentation appears to be important in interspecific diversity in certain groups of fishes and birds. Pheromonal signals serve as the primary basis for species-specific mating cues in many salamander species, most mammals and even some fishes. The evolution of interspecific divergence in elaborate courtship displays may have played an important role in speciation of lizards, and particular groups of fishes, salamanders, birds and mammals. While much research has focused on the importance of mating preferences in shaping the evolution of these types of mating signals within species, the link between intraspecific preferences and interspecific divergence and speciation remains to be more fully tested. Future studies should focus on identifying how variation in mating preferences within a species shapes interspecific diversity in features of mating signals in order to better understand how sexual selection may have led to speciation in vertebrates.

Journal Article↗

Interactions and associations between age and sex classes in mouflon sheep (Ovis gmelini) during winter.

The fact that sexual segregation can occur in wild ruminants independently of sex-related differences in habitat use suggests that it results at least partly from social mechanisms. One of the simplest mechanisms which can be hypothesised is that individuals avoid the congener classes which frequently try to interact with them (be these interactions 'agonistic' or 'sexual', since courtship behaviour components typically induce withdrawal when performed outside the rut). The aim of the present study was to test, in mouflon sheep (Ovis gmelini), whether the age-sex classes which exchange frequent interactions on meeting occur rarely inside the same groups. Observations were made during winter, on a small population in Germany. Interactions were recorded at an attractive site where all age-sex classes met daily. Foot transects were conducted in parallel over the whole study area in order to determine group composition. As expected, adults of different sexes exchanged frequent interactions at the attractive site and were rarely found in the same groups under free-ranging conditions, while adult females, who interacted little with one another, were quite frequently associated. However, adult males simultaneously exhibited the highest rate of interaction at the attractive site and the highest frequency of association under free-ranging conditions. Unlike female-female and male-female pairs, in which interactions with a clear initiator and a clear recipient predominated, male-male pairs exchanged often spectacular but 'symmetrical' interactions (during which the two individuals behaved in the same way). The hypothesis that sexual segregation results from the avoidance of the congener classes which frequently engage interactions only holds for females. Adult rams interact frequently and seek companions with whom symmetrical interactions can be performed.

Journal Article↗

Variability of vibratory signals and mate choice selectivity in the southern green stink bug.

The southern green stink bug, Nezara viridula, has a complex mate recognition system that implicates chemical and acoustic signals. To localise a sexual partner acoustically, a male and female alternate between the male courtship song (MCrS) and the female calling song type 1 (FCS1). Although previous research has revealed that both signals show geographical variability, until now no studies have explored the form of this variability. We analysed the temporal and spectral characteristics of MCrS and FCS1 pulse trains of males and females from a French and a Guadeloupe population. Pulse train duration of the MCrS varied within and between populations. Likewise, spectral and temporal parameters of FCS1 varied within and between the two populations. Although females did not show any behavioural responses to pre-recorded MCrS, males responded to pre-recorded FCS1 by emitting a higher number of MCrS per minute. Furthermore, males modulated the repetition rate of their MCrS pulse trains to match those of the FCS1. All males responded to FCS1 from French and Guadeloupe females despite the temporal differences in these songs; however, they responded with a shorter latency and a higher rate of MCrS/FCS1 pulse trains to the songs of females from their own population. In choice experiments with two alternatives, responses to FCS1 were inhibited when males received a simultaneous female call from another Pentatomidae sympatric species, Acrosternum hilare. We conclude that, although males prefer FCS1 from their own population, they recognise FCS1 from French and Guadeloupe N. viridula females as species-specific female calls. Variability of vibratory signals might play a role in mate choice.

Journal Article↗

Associative learning and memory in Drosophila: beyond olfactory conditioning.

The associative learning abilities of the fruit fly, Drosophila melanogaster, have been demonstrated in both classical and operant conditioning paradigms. Efforts to identify the neural pathways and cellular mechanisms of learning have focused largely on olfactory classical conditioning. Results derived from various genetic and molecular manipulations provide considerable evidence that this form of associative learning depends critically on neural activity and cAMP signaling in brain neuropil structures called mushroom bodies. Three other behavioral learning paradigms in Drosophila serve as the main subject of this review. These are (1) visual and motor learning of flies tethered in a flight simulator, (2) a form of spatial learning that is independent of visual and olfactory cues, and (3) experience-dependent changes in male courtship behavior. The present evidence suggests that at least some of these modes of learning are independent of mushroom bodies. Applying targeted genetic manipulations to these behavioral paradigms should allow for a more comprehensive understanding of neural mechanisms responsible for diverse forms of associative learning and memory.

Journal Article↗

Mild mutations in the pan neural gene prospero affect male-specific behaviour in Drosophila melanogaster.

The fruitfly Drosophila melanogaster is one of the most appropriate model organisms to study the genetics of behaviour. Here, we focus on prospero (pros), a key gene for the development of the nervous system which specifies multiple aspects from the early formation of the embryonic central nervous system to the formation of larval and adult sensory organs. We studied the effects on locomotion, courtship and mating behaviour of three mild pros mutations. These newly isolated pros mutations were induced after the incomplete excision of a transposable genomic element that, before excision, caused a lethal phenotype during larval development. Strikingly, these mutant strains, but not the strains with a clean excision, produced a high frequency of heterozygous flies, after more than 50 generations in the lab. We investigated the factors that could decrease the fitness of homozygotes relatively to heterozygous pros mutant flies. Flies of both genotypes had slightly different levels of fertility. More strikingly, homozygous mutant males had a lower sexual activity than heterozygous males and failed to mate in a competitive situation. No similar effect was detected in mutant females. These findings suggest that mild mutations in pros did not alter vital functions during development but drastically changed adult male behaviour and reproductive fitness.

Alleles↗

Genomic organisation and characterisation of the neural sex-determination gene fruitless (fru) in the Hawaiian species Drosophila heteroneura.

There are several mechanisms for the determination of sex. Sexual behaviour is part of the sex-determination cascade, and in Drosophila melanogaster male courtship is controlled in part by the fruitless gene. As part of a study of sexual behaviour in Hawaiian Drosophila, we have cloned the neural sex-determination gene fru from the Hawaiian picture-wing species Drosophila heteroneura. The fru gene has at least seven exons covering a region of 18kb and encodes three transcripts, fruA, fruB and fruC. Each transcript encodes a single ORF of 841, 678 and 691aa, respectively. The FRUA and FRUB proteins have a BTB protein-protein-binding domain and two zinc finger-like domains and are well conserved with the D. melanogaster proteins. The FRUC protein has a BTB domain but no zinc finger-like domains. The fru gene is expressed in 1-7 day old adult males as a 5.1kb transcript. This transcript is not seen in adult females, so the fru gene has a different pattern of sex-differential expression in the Hawaiian Drosophila compared with D. melanogaster.

Alternative Splicing↗

Diversity in frequency response properties of saccular afferents of the toadfish, Opsanus tau.

The frequency response of primary saccular afferents of toadfish (Opsanus tau) was studied in the time and frequency domains using the reverse correlation (revcor) method. Stimuli were noise bands with flat acceleration spectra delivered as whole-body motion. The recorded acceleration waveform was averaged over epochs preceding and following each spike. This average, termed the revcor, is an estimate of the response of an equivalent linear filter intervening between body motion and spike initiation. The spectrum of the revcor estimates the shape of the equivalent linear filter. Revcor responses were brief, damped oscillations indicative of relatively broadly tuned filters. Filter shapes were generally band-pass and differed in bandwidth, band edge slope, and characteristic frequency (74 Hz to 140 Hz). Filter shapes tend to be independent of stimulus level. Afferents can be placed into two groups with respect to characteristic frequency (74-88 Hz and 140 Hz). Some high-frequency afferents share a secondary peak at the characteristic frequency of low-frequency afferents, suggesting that an afferent may receive differently tuned peripheral inputs. For some afferents having similar filter shapes, revcor responses often differ only in polarity, probably reflecting inputs from hair cells oriented in opposite directions. The origin of frequency selectivity and its diversity among saccular afferents may arise from a combination of hair cell resonance and micromechanical processes. The resulting frequency analysis is the simplest yet observed among vertebrate animals. During courtship, male toadfish produce the 'boatwhistle' call, a periodic vocalization having several harmonics of a 130 Hz fundamental frequency. The saccule encodes the waveform of acoustic particle acceleration between < 50 and about 250 Hz. Thus, the fundamental frequency component of the boatwhistle is well encoded, but the successive higher harmonics are filtered out. The boatwhistle is thus encoded as a time-domain representation of its fundamental frequency or pulse repetition rate.

Acoustic Stimulation↗

Vasotocin and reproductive functions of the domestic chicken.

The neurohypophyseal hormone arginine vasotocin (AVT) combines both antidiuretic and reproductive activities. In the domestic chicken AVT produces assimetric effects on the reproductive functions of males and females. AVT synthesized in magnocellular diencephalic neurons is released into circulation in a highly coordinated manner contributing to the peripheral control of oviposition in hens. Conversely, parvocellular AVT cells located in the limbic system (bed nucleus of stria terminalis (BST)) are quite different in their properties and, possible, functions. In domestic chickens these cells express AVT in a sexually dimorphic manner and are found solely in males. This sexually dimorphic part of the AVT system is sensitive to gonadal steroids. Experimental data demonstrated that AVT modulates different aspects of reproductive behavior including courtship vocalization and copulation. Sexual differentiation of these limbic vasotocinergic cells show striking correlation with sexual differentiation of masculine behavior. Evidences coming from physiological, anatomical and ethological studies suggest strong implication of the vasotocinergic system in the control of reproductive functions.

Animals↗

Normal and abnormal sexual behavior.

This article includes a brief description of the reproductive patterns and specific behaviors comprising courtship and copulation in equids. The author then focuses on breeding behavior of the domestic stallion. Descriptions of normal breeding behavior under a variety of domestic management systems, comments on sexual behavior of donkeys and exotic equids, and the most common types and treatments of sexual behavior dysfunction are included.

Animals↗

For whom the bird sings: context-dependent gene expression.

Male zebra finches display two song behaviors: directed and undirected singing. The two differ little in the vocalizations produced but greatly in how song is delivered. "Directed" song is usually accompanied by a courtship dance and is addressed almost exclusively to females. "Undirected" song is not accompanied by the dance and is produced when the male is in the presence of other males, alone, or outside a nest occupied by its mate. Here, we show that the anterior forebrain vocal pathway contains medial and lateral "cortical-basal ganglia" subdivisions that have differential ZENK gene activation depending on whether the bird sings female-directed or undirected song. Differences also occur in the vocal output nucleus, RA. Thus, although these two vocal behaviors are very similar, their brain activation patterns are dramatically different.

Animals↗

Dissatisfaction encodes a tailless-like nuclear receptor expressed in a subset of CNS neurons controlling Drosophila sexual behavior.

The dissatisfaction (dsf) gene is necessary for appropriate sexual behavior and sex-specific neural development in both sexes. dsf males are bisexual and mate poorly, while mutant females resist male courtship and fail to lay eggs. Males and females have sex-specific neural abnormalities. We have cloned dsf and rescued both behavioral and neural phenotypes. dsf encodes a nuclear receptor closely related to the vertebrate Tailless proteins and is expressed in both sexes in an extremely limited set of neurons in regions of the brain potentially involved in sexual behavior. Expression of a female transformer cDNA under the control of a dsf enhancer in males leads to dsf-like bisexual behavior.

Amino Acid Sequence↗

Gender roles and interactions in drinking and drug use.

In gender-focused discussions of alcohol and other drug use and problems, the emphasis has usually been on the individual male or female or on the genders as aggregates of individuals. But most drinking and much drug use have strong social and interactional elements, where gender roles and often gendered interactions come into play. Drawing on the existing literature, opportunities for research on gender roles and interactions in drinking and drug use and problems are discussed under the following headings: courtship and affectional preference; sexuality; marriage and partnership; parenthood; friendship and peer relations; work roles; informal social control (spouse, relatives, friends); and domination, violence, and abuse.

Alcoholism↗

Multimodal sensory integration in weakly electric fish: a behavioral account.

The ability to integrate multisensory information is a fundamental characteristic of the brain serving to enhance the detection and identification of external stimuli. Weakly electric fish employ multiple senses in their interactions with one another and with their inanimate environment (electric, visual, acoustic, mechanical, chemical, thermal, and hydrostatic pressure) and also generate signals using some of the same stimulus energies (electric, acoustic, visual, mechanical). A brief overview provides background on the sensory and motor channels available to the fish followed by an examination of how weakly electric fish 'benefit' from integrating various stimulus modalities that assist in prey detection, schooling, foraging, courtship, and object location. Depending on environmental conditions, multiple sensory inputs can act synergistically and improve the task at hand, can be redundant or contradictory, and can substitute for one another. Over time, in repeated encounters with familiar surrounds, loss of one modality can be compensated for through learning. Studies of neuronal substrates and an understanding of the computational algorithms that underlie multisensory integration ought to expose the physiological corollaries to widely published concepts such as internal representation, sensory expectation, sensory generalization, and sensory transfer.

Animals↗

Neuroethology and life history adaptations of the elasmobranch electric sense.

The electric sense of elasmobranch fishes (sharks and rays) is an important sensory modality known to mediate the detection of bioelectric stimuli. Although the best known function for the use of the elasmobranch electric sense is prey detection, relatively few studies have investigated other possible biological functions. Here, we review recent studies that demonstrate the elasmobranch electrosensory system functions in a wide number of behavioral contexts including social, reproductive and anti-predator behaviors. Recent work on non-electrogenic stingrays demonstrates that the electric sense is used during reproduction and courtship for conspecific detection and localization. Electrogenic skates may use their electrosensory encoding capabilities and electric organ discharges for communication during social and reproductive interactions. The electric sense may also be used to detect and avoid predators during early life history stages in many elasmobranch species. Embryonic clearnose skates demonstrate a ventilatory freeze response when a weak low-frequency electric field is imposed upon the egg capsule. Peak frequency sensitivity of the peripheral electrosensory system in embryonic skates matches the low frequencies of phasic electric stimuli produced by natural fish egg-predators. Neurophysiology experiments reveal that electrosensory tuning changes across the life history of a species and also seasonally due to steroid hormone changes during the reproductive season. We argue that the ontogenetic and seasonal variation in electrosensory tuning represent an adaptive electrosensory plasticity that may be common to many elasmobranchs to enhance an individual's fitness throughout its life history.

Adaptation, Biological↗

Electric signaling behavior and the mechanisms of electric organ discharge production in mormyrid fish.

Mormyrid fish communicate and navigate using electric organ discharges (EODs). The EOD is highly stereotyped and provides information on sender identity, including species, sex, reproductive condition, and possibly relative status and individual identity. By contrast, the sequence of pulse intervals (SPI) is variable and plays more of a role in signaling behavioral states. Various types of SPI displays may be produced, including tonic patterns such as 'random' and 'regularized', and phasic patterns such as 'bursts' and cessations'. Certain displays have been linked to specific behaviors such as aggression, submission, courtship and active exploration. In addition, interacting pairs of fish may produce stereotyped displays involving the relative timing of their EODs. The EOD waveform is controlled by the morphological and physiological properties of cells in the electric organ termed electrocytes. Differences in the innervation, morphology, size and membrane characteristics of electrocytes have been directly linked to species and sex differences in the EOD. The generation of each EOD is initiated in the medullary command nucleus (CN), which thereby determines the timing of EOD output. CN does not have any properties of a pacemaker, but rather appears to integrate descending inputs that affect the probability of EOD production. The precommand nucleus (PCN) provides a major source of excitatory input to CN and is itself inhibited by corollary discharge feedback following the production of each EOD. Changes in the activity of PCN and its inhibitory feedback neurons modify EOD output, and therefore drive the generation of SPI patterns. Current studies are addressing the mechanisms underlying the generation of these patterns and preliminary results suggest that different types of signals may be controlled by distinct components of the electromotor system. This is similar to findings in other electrogenic teleosts, suggesting that it may be a general feature in the motor control of signaling behavior.

Action Potentials↗

Separated dental laminae are present in the upper jaw of Mesoamerican lungless salamanders (Amphibia, Plethodontidae).

Plethodontid salamander species from Northern America exhibit a single, unseparated dental lamina in the upper jaw, which consistently provides teeth for the unpaired premaxillary and the two maxillary bones. During the distinct mating season, adult males of these species bear long, monocuspid, unbladed teeth in the entire upper jaw, which are used during courtship. However, plethodontid salamander species from Southern Central America studied so far show either three separated dental laminae or a single dental lamina with three distinguishable tooth-producing parts connected by non-producing parts. These species reproduce aseasonally and sexually mature males permanently bear long, monocuspid, unbladed teeth on the premaxillary only. As the formation of these long, monocuspid teeth depends on the presence of androgen-receptors in the tooth-forming tissues, the morphological separation of the dental lamina in plethodontid salamanders from Southern Central America is very likely to be a prerequisite for a region-specific expression of androgen-receptors in the tooth-forming tissues controlling the differentiated shape formation of teeth in these species. Seven species of plethodontid salamanders from Costa Rica and Panama were examined and, in all, a tendency to exhibit three separate dental lamina in the upper jaw was found, although the morphological separation is definitive only in Bolitoglossa colonnea, B. lignicolor, B. pesrubra, B. schizodactyla and B. striatula. B. dofleini and B. marmorea show a continuous dental lamina with non-producing parts at the transitional sites between the parts associated with the premaxillary and the maxillary bones.

Animals↗

Behavioral functions of the insect mushroom bodies.

New methods of intervention in Drosophila and other insect species reveal that the mushroom bodies are involved in a diverse set of behavioral functions. The intrinsic Kenyon cells (those neurons with projections within the mushroom bodies) house part of the short-term memory trace for odors and are required for courtship conditioning memory. A pair of extrinsic mushroom body neurons (neurons with projections both inside and outside the mushroom bodies) provides a neuropeptide important for 1-hour olfactory memory. In addition, the mushroom bodies are necessary for context generalization in visual learning and for regulating the transition from walking to rest.

Animals↗

A kinder, gentler genetic analysis of behavior: dissection gives way to modulation.

Many of the mutations and genetic variants that affect behavior in Drosophila have proved to be mild lesions of genes that are capable of more severe phenotypes. Examples of such variants affecting ion channels, transcription factors and protein kinases in studies of courtship and learning have anticipated recent findings on the naturally occurring variants in circadian rhythms and foraging.

Animals↗