Blockade of the pheromonal effects in rat by central deafferentation of the accessory olfactory system

Female rats reared without sex odours from male rats have a five day stral cycle. With exposure to male odour the estral cycle is shortened from five to four days. This pheromonal effect is blocked on deafferenting the vomeronasal system by electrolytically damaging both accessory olfactory bulbs.     

Emerging views on the distinct but related roles of the main and accessory olfactory systems in responsiveness to chemosensory signals in mice

In rodents, the nasal cavity contains two separate chemosensory epithelia, the main olfactory epithelium, located in the posterior dorsal aspect of the nasal cavity, and the vomeronasal/accessory olfactory epithelium, located in a capsule in the anterior aspect of the ventral floor of the nasal cavity. Both the main and accessory olfactory systems play a role in detection of biologically relevant odors. The accessory olfactory system has been implicated in response to pheromones, while the main olfactory system is thought to be a general molecular analyzer capable of detecting subtle differences in molecular structure of volatile odorants. However, the role of the two systems in detection of biologically relevant chemical signals appears to be partially overlapping. Thus, while it is clear that the accessory olfactory system is responsive to putative pheromones, the main olfactory system can also respond to some pheromones. Conversely, while the main olfactory system can mediate recognition of differences in genetic makeup by smell, the vomeronasal organ (VNO) also appears to participate in recognition of chemosensory differences between genetically distinct individuals. The most salient feature of our review of the literature is that there are no general rules that allow classification of the accessory olfactory system as a pheromone detector and the main olfactory system as a detector of general odorants. Instead, each behavior must be considered within a specific behavioral context to determine the role of these two chemosensory systems. In each case, one system or the other (or both) participates in a specific behavioral or hormonal response.     

Can audio–visual or visual stimuli from a prospective mate stimulate a reproductive neuroendocrine response in sheep?

Stimuli from a prospective mate increase the secretion of luteinising hormone (LH) in sheep. This 'male effect' in ewes and 'female effect' effect in rams is predominantly mediated by olfactory signals, though it is thought that non-olfactory signals play synergistic or substitutive roles. In this study, we tested whether exposure to visual or audio-visual stimuli from a prospective mate would stimulate an increase in LH secretion in ewes (Experiment 1) and rams (Experiment 2). In Experiment 1, groups of eight Merino ewes were exposed to one of three stimuli midway through a frequent blood-sampling regimen: full ram contact, still images of rams, a video of ewes and rams mating. Control ewes (n = 8) were completely isolated from rams. Exposure to still images of rams appeared to stimulate an increase in mean LH concentrations (P < 0.05) and tended to increase LH pulse frequency (P < 0.1), but the response was significantly smaller than that observed in ewes exposed to rams (P < 0.01). Audio-visual stimuli had no effect on any parameters of LH secretion (P > 0.1). In Experiment 2, Merino rams were allocated to either an Exposure (n = 7) or a Control (n = 7) group. Exposure rams underwent two exposure periods midway through a frequent blood-sampling regimen; exposure to still images of ewes and audio recorded during mating of ewes and rams (audio-visual exposure); exposure to oestrous ewes (ewe exposure). Control rams were sampled at the same frequency but remained isolated from ewe stimuli. Exposure of rams to the audio-visual stimuli did not affect any parameters of LH secretion (P > 0.1). In contrast, exposure to oestrous ewes increased LH pulse frequency (P < 0.05) and advanced the onset of the next LH pulse (P < 0.05). In conclusion, visual signals appear to be involved in eliciting the neuroendocrine response of ewes to rams and are of greater importance to this phenomenon in ewes (male effect) than rams (female effect). However, overall the visual and audio-visual signals used in this study were far less effective than stimulus animals, suggesting that these stimuli are less important than olfactory signals, or a combination of olfactory and audio-visual signals.     

Removal of the vomeronasal organ blocks the stress-induced hyperthermia response to alarm pheromone in male rats

Previously, we reported that male Wistar rats release alarm pheromone from their perianal region, which aggravates stress-induced hyperthermia (SIH) in pheromone-recipient rats. The subsequent discovery that this pheromone could be trapped in water enabled us to expose recipients to the pheromone in their home cages. Despite its apparent influence on autonomic and behavioral functions, we still had no clear evidence as to whether the alarm pheromone was perceived by the main olfactory system (MOS) or by the vomeronasal system. In this study, we investigated this question by exposing 3 types of recipients to alarm pheromone in their home cages: intact males (Intact), vomeronasal organ-excised males (VNX), and sham-operated males (Sham). The Intact and Sham recipients showed aggravated SIH in response to alarm pheromone, whereas the VNX recipients did not. In addition, the results of the habituation/dishabituation test and soybean agglutinin binding to the accessory olfactory bulb verified the complete ablation of the vomeronasal organ (VNO) with a functional MOS in the pheromone recipients. These results strongly suggest that male rats perceive alarm pheromone with the VNO.     

Role of the vomeronasal organ in neonatal offspring recognition in sheep

Twenty-five pregnant Dorsett ewes were randomly divided into three groups to test if ewes use their vomeronasal organs for offspring recognition during nursing. One group of eight ewes (procaine) were made anosmic by irrigation of the nasal olfactory apparatus with a zinc sulphate procaine solution. The second group of nine ewes (cauterized) had their vomeronasal organs rendered nonfunctional by cauterization of the nasoincisive duct. The third group of eight ewes were the controls. Parturition was synchronized in all ewes with betamethasone on Day 145 of gestation. Maternal responsiveness was tested two separate times with 1- to 2-day-old alien lambs. Each alien lamb trial was conducted 24 h apart. Cauterized ewes allowed alien lambs to suckle and they were unable to distinguish alien lambs from their own lambs, whereas the ewes in both groups with functional vomeronasal organs (procaine and control) violently rejected any alien lamb's attempt to suckle. Thus, female sheep use their vomeronasal organs for neonatal offspring recognition.     

Mammalian social odours: attraction and individual recognition

Mammalian social systems rely on signals passed between individuals conveying information including sex, reproductive status, individual identity, ownership, competitive ability and health status. Many of these signals take the form of complex mixtures of molecules sensed by chemosensory systems and have important influences on a variety of behaviours that are vital for reproductive success, such as parent-offspring attachment, mate choice and territorial marking. This article aims to review the nature of these chemosensory cues and the neural pathways mediating their physiological and behavioural effects. Despite the complexities of mammalian societies, there are instances where single molecules can act as classical pheromones attracting interest and approach behaviour. Chemosignals with relatively high volatility can be used to signal at a distance and are sensed by the main olfactory system. Most mammals also possess a vomeronasal system, which is specialized to detect relatively non-volatile chemosensory cues following direct contact. Single attractant molecules are sensed by highly specific receptors using a labelled line pathway. These act alongside more complex mixtures of signals that are required to signal individual identity. There are multiple sources of such individuality chemosignals, based on the highly polymorphic genes of the major histocompatibility complex (MHC) or lipocalins such as the mouse major urinary proteins. The individual profile of volatile components that make up an individual odour signature can be sensed by the main olfactory system, as the pattern of activity across an array of broadly tuned receptor types. In addition, the vomeronasal system can respond highly selectively to non-volatile peptide ligands associated with the MHC, acting at the V2r class of vomeronasal receptor. The ability to recognize individuals or their genetic relatedness plays an important role in mammalian social behaviour. Thus robust systems for olfactory learning and recognition of chemosensory individuality have evolved, often associated with major life events, such as mating, parturition or neonatal development. These forms of learning share common features, such as increased noradrenaline evoked by somatosensory stimulation, which results in neural changes at the level of the olfactory bulb. In the main olfactory bulb, these changes are likely to refine the pattern of activity in response to the learned odour, enhancing its discrimination from those of similar odours. In the accessory olfactory bulb, memory formation is hypothesized to involve a selective inhibition, which disrupts the transmission of the learned chemosignal from the mating male. Information from the main olfactory and vomeronasal systems is integrated at the level of the corticomedial amygdala, which forms the most important pathway by which social odours mediate their behavioural and physiological effects. Recent evidence suggests that this region may also play an important role in the learning and recognition of social chemosignals.     

The rodent accessory olfactory system

The accessory olfactory system contributes to the perception of chemical stimuli in the environment. This review summarizes the structure of the accessory olfactory system, the stimuli that activate it, and the responses elicited in the receptor cells and in the brain. The accessory olfactory system consists of a sensory organ, the vomeronasal organ, and its central projection areas: the accessory olfactory bulb, which is connected to the amygdala and hypothalamus, and also to the cortex. In the vomeronasal organ, several receptors—in contrast to the main olfactory receptors—are sensitive to volatile or nonvolatile molecules. In a similar manner to the main olfactory epithelium, the vomeronasal organ is sensitive to common odorants and pheromones. Each accessory olfactory bulb receives input from the ipsilateral vomeronasal organ, but its activity is modulated by centrifugal projections arising from other brain areas. The processing of vomeronasal stimuli in the amygdala involves contributions from the main olfactory system, and results in long-lasting responses that may be related to the activation of the hypothalamic–hypophyseal axis over a prolonged timeframe. Different brain areas receive inputs from both the main and the accessory olfactory systems, possibly merging the stimulation of the two sensory organs to originate a more complex and integrated chemosensory perception.     

The "male effect" in sheep and goats: a review of the respective roles of the two olfactory systems

In sheep and goats, exposure of seasonally anestrous females to sexually active males results in activation of luteinizing hormone (LH) secretion and synchronized ovulation. This phenomenon is named "the male effect" and seems to constitute a major factor in the control of reproductive events. This effect depends mostly on olfactory cues and is largely mimicked by exposure to male fleece only. In sheep, preventing the vomeronasal organ (VNO) from functioning does not affect the female responses to male odor suggesting that, unlike in rodents, the accessory olfactory system does not play the major role in the perception of this pheromonal cue. Female responses also seem to depend on previous experience, an effect that is not common for pheromones and renders this model of special interest. The aim of the present report is to summarize our current knowledge concerning the "male effect" and in particular to clarify the respective roles of the two olfactory systems in the processes involved in this effect.     

Experiential and endocrine dependence of gonadotropin responses in male mice to conspecific urine

Previous research has shown that a urinary pheromone of female mice acts via the vomeronasal organ of the accessory olfactory system to elicit rapid release of luteinizing hormone (LH) in conspecific males. Several experiments were conducted to examine the importance of sexual experience for gonadotropin responses in male mice to female urine, male urine, saline, or mixtures of these stimuli. Both sexually naive and sexually experienced male mice had significantly higher plasma LH levels after presentations of female urine than after presentations of male urine. However, sexual experience appeared to increase the reliability of the short-latency gonadotropin response to female urine relative to a sexually neutral component of urine such as sodium chloride, and male urine appeared to suppress spontaneous LH secretion episodes in both naive and sexually experienced males. Subsequent experiments with sexually experienced subjects demonstrated that male mouse urine is a powerful suppressant of LH release in other males. Specifically, female mouse urine mixed with male urine failed to elicit LH responses in male subjects, whereas female urine mixed with saline was highly effective. Urine obtained from castrated male donors was as potent as urine from intact males in suppressing the gonadotropin response to female urine. The suppressive activity in male mouse urine thus does not appear to be critically dependent on gonadal hormones. The existence of a potent stimulatory pheromone in female urine and a potent suppressive pheromone in male urine makes male mice an excellent model system for studying the neural regulation of LH secretion.     

GnRH-induced gonadotrophin secretion in ovariectomized Booroola ewes with hypothalamic-pituitary disconnection

To test whether the F gene-specific differences in the plasma concentrations of FSH and LH are due to differences in the pituitary responsiveness to exogenous GnRH, ovariectomized Booroola ewes with hypothalamic-pituitary disconnection (HPD-ovx) were treated with GnRH (250 ng i.v.) once every 2 h for up to 5 weeks. In Exp. 1, jugular venous blood was collected once weekly from 13 FF and 14 ++ HPD-ovx ewes for 6 weeks before GnRH treatment and every 2nd, 3rd or 6th day for 5 weeks during treatment. In Exp. 2, jugular venous blood was collected from another 8 FF and 7 ++ HPD-ovx ewes at 5- or 10-min intervals over 4 GnRH pulses (250 ng i.v. once every 2 h) on 3 separate occasions after the animals had been subjected to the GnRH pulse regimen for approximately 7 days beforehand. Also in Exp. 2, the animals were extensively sampled around a larger (10 micrograms) i.v. injection of GnRH and the pituitary FSH and LH contents assessed after the animals had been re-exposed to the once every 2 h GnRH (250 ng i.v.) pulse regimen for several days following the larger GnRH bolus. In Exp. 3 the distributions of mean plasma concentrations of FSH and LH in individual GnRH-treated HPD-ovx ewes were compared with those in ovariectomized and ovary-intact FF and ++ ewes. During the 6 weeks before GnRH treatment (Exp. 1), the plasma concentrations of FSH (approximately 1 ng/ml) and LH (less than or equal to 0.8 ng/ml) were not different between the genotypes. After GnRH treatment both the mean FSH and LH concentrations increased significantly (P less than 0.01) above basal values after 2 days with F gene-specific differences being noted for FSH but not LH (FSH; FF greater than ++; P less than 0.05). Thereafter, the mean FSH but not LH concentrations increased at a faster rate in FF than in ++ ewes with the overall mean FSH concentrations between the genotypes being significantly different (P less than 0.05). In Exp. 2 considerable between-animal variation in the pulsatile pattern of FSH but not LH concentrations was seen in ewes of both genotypes during GnRH treatment. The overall mean FSH concentrations were higher in FF than in ++ ewes (P less than 0.05) and the mean FSH response to each GnRH pulse was significantly higher in FF than in ++ ewes (P less than 0.05).(ABSTRACT TRUNCATED AT 400 WORDS)     

Role of the olfactory systems and importance of learning in the ewes' response to rams or their odors

In sheep, exposure of seasonally anestrous females to the male or its fleece results in activation of luteinizing hormone (LH) secretion and synchronized ovulation. The study of the neural pathways involved in this phenomenon, commonly named "male effect", show that the main olfactory system plays a critical role in the detection and the integration of the male odor. The accessory olfactory system participates in the perception of the ram odor but does not seem necessary for the endocrine response. According to the hypothesis that the neuroanatomical differences between the two olfactory systems could be associated with different functional roles, we investigated the importance of sexual experience and learning processes in the male effect. Our results showed that female responses depend on previous sexual experience. We also demonstrated that the LH response to male odor could result from an associative learning process. The aim of the present report was to summarize our current knowledge concerning the "male effect" and in particular to clarify the role of sexual experience and learning in the processes involved in this effect.     

哺乳动物主要嗅觉系统和犁鼻系统信息识别的编码模式

哺乳动物具有两套嗅觉系统, 即主要嗅觉系统和犁鼻系统。前者对环境中的大多数挥发性化学物质进行识别, 后者对同种个体释放的信息素进行识别。本文从嗅觉感受器、嗅球、嗅球以上脑区三个水平综述了这两种嗅觉系统对化学信息识别的编码模式。犁鼻器用较窄的调谐识别信息素成分, 不同于嗅上皮用分类性合并受体的方式识别气味; 副嗅球以接受相同受体输入的肾丝球所在区域为单位整合信息, 而主嗅球通过对肾丝球模块的特异性合并编码信息; 在犁鼻系统, 信息素的信号更多地作用于下丘脑区域, 引起特定的行为和神经内分泌反应。而在主要嗅觉系统, 嗅皮层可能采用时间模式编码神经元群, 对气味的最终感受与脑的不同区域有关。犁鼻系统较主要嗅觉系统的编码简单, 可能与其执行的功能较少有关。     

哺乳类两大嗅觉系统功能的研究进展

近年来对于嗅觉系统的研究已成为动物学研究的热点之一,本文通过对以往关于哺乳类两大嗅觉系统功能的研究进行总结和回顾,对目前犁鼻器系统(VOE-AOB)和主嗅觉系统(MOE-MOB)功能结论上的争议做了初步探讨。通过概括和总结,发现目前对两大嗅觉系统功能还存在争议,其原因可能有以下几点:以前的研究方法上可能有不完善之处;不同的研究采用的物种不同,结论上的争议也许与不同物种间存在种间差异有关;动物的社会经验对研究结论可能也有一定影响。希望通过本文能进一步促进今后此方面的研究。     

Importance of learning in the response of ewes to male odor

Exposure of anestrous ewes to a ram or its odor results in the activation of the luteinizing hormone (LH) secretion leading to reinstatement of cyclicity in most females. Sexual experience and learning have been suggested as important factors to explain the variability of the female responses. In experiment 1, we compared the behavioral and endocrine responses of four groups of anestrous females that differed in age (young or adult) and previous exposure to males [naive (no exposure) or experienced (courtship behavior for young and numerous mating for adults)]. Age did not seem to affect the LH response to males or their odor. In contrast, sexual experience was a critical factor: the proportion of females exhibiting an LH response to male odor was significantly higher in experienced than in naive ewes. Sexual experience affected the response to male odor, but did not have an effect on responses to the male himself. A second experiment investigated whether the LH response to male odor could result from an associative learning process. Accordingly, we tested the effectiveness of a conditioned stimulus (lavender odor) previously associated with the male, in inducing the endocrine response. The results indicate that the odor of lavender activated LH secretion only in ewes that have been previously exposed to scented males. This demonstrates that ewes are able to learn the association between a neutral odor and their sexual partner.     

Sexual experience does not compensate for the disruptive effects of zinc sulfate--lesioning of the main olfactory epithelium on sexual behavior in male mice

Recent studies point to an important role for the main olfactory epithelium (MOE) in regulating sexual behavior in male mice. We asked whether sexual experience could compensate for the disruptive effects of lesioning the MOE on sexual behavior in male mice. Male mice, which were either sexually naive or experienced, received an intranasal irrigation of either a zinc sulfate solution to destroy the MOE or saline. Sexual behavior in mating tests with an estrous female was completely abolished in zinc sulfate-treated male mice regardless of whether subjects were sexually experienced or not before the treatment. Furthermore, zinc sulfate treatment clearly disrupted olfactory investigation of both volatile and nonvolatile odors. Destruction of the MOE by zinc sulfate treatment was confirmed by a significant reduction in the expression of Fos protein in the main olfactory bulb following exposure to estrous female urine. By contrast, vomeronasal function did not seem to be affected by zinc sulfate treatment: nasal application of estrous female urine induced similar levels of Fos protein in the mitral and granule cells of the accessory olfactory bulb (AOB) of zinc sulfate- and saline-treated males. Likewise, the expression of soybean agglutinin, which stains the axons of vomeronasal organ neurons projecting to the glomerular layer of the AOB, was similar in zinc sulfate- and saline-treated male mice. These results show that the main olfactory system is essential for the expression of sexual behavior in male mice and that sexual experience does not overcome the disruptive effects of MOE lesioning on this behavior.     

Gonadotrophin release in ovariectomized ewes fed different amounts of coumestrol

Three experiments were conducted to study changes in pulsatile secretion of LH and FSH during the breeding season or anoestrus in ovariectomized Ile-de-France ewes fed different amounts of the phyto-oestrogen coumestrol. In Exp. 1, conducted during the breeding season, ewes (3-4 per group) were fed lucerne supplying 4, 18 or 30 mg coumestrol per ewe per day for 15 days. Experiments 2 and 3 were conducted during seasonal anoestrus. In Exp. 2, ewes (4 per group) were fed lucerne supplying coumestrol concentrations ranging from 4 to 38 mg/ewe/day for 15 days. In Exp. 3, ewes (10 per group) were fed lucerne supplying 14 or 125 mg coumestrol/ewe/day for 15 days. During the breeding season, an increased concentration of coumestrol in the diet significantly decreased the amplitude of LH pulses. There were no effects on LH pulse frequency or on FSH concentrations. During seasonal anoestrus, there were no significant effects on LH pulse frequency, or amplitude and no significant effect on FSH concentration. These results show that high concentrations of coumestrol in lucerne diets would not explain seasonal variation in LH pulse frequency in ovariectomized ewes. However, lucerne diets with increased coumestrol concentrations can influence LH release during the breeding season.     

Olfactory receptors and signalling elements in the Grueneberg ganglion

The Grueneberg ganglion (GG) is a cluster of neurones present in the vestibule of the anterior nasal cavity. Although its function is still elusive, recent studies have shown that cells of the GG transcribe the gene encoding the olfactory marker protein (OMP) and project their axons to glomeruli of the olfactory bulb, suggesting that they may have a chemosensory function. Chemosensory responsiveness of olfactory neurones in the main olfactory epithelium (MOE) and the vomeronasal organ (VNO) is based on the expression of either odorant receptors or vomeronasal putative pheromone receptors. To scrutinize its presumptive olfactory nature, the GG was assessed for receptor expression by extensive RT-PCR analyses, leading to the identification of a distinct vomeronasal receptor which was expressed in the majority of OMP-positive GG neurones. Along with this receptor, these cells expressed the G proteins Go and Gi, both of which are also present in sensory neurones of the vomeronasal organ. Odorant receptors were expressed by very few cells during prenatal and perinatal stages; a similar number of cells expressed adenylyl cyclase type III and G(olf/s), characteristic signalling elements of the main olfactory system. The findings of the study support the notion that the GG is in fact a subunit of the complex olfactory system, comprising cells with either a VNO-like or a MOE-like phenotype. Moreover, expression of a vomeronasal receptor indicates that the GG might serve to detect pheromones.     

A new strategy for superior reproductive performance of ewes bred out-of-season utilizing progestagen supplement prior to withdrawal of intravaginal pessaries

Two experiments were conducted to examine the effect of progestagen supplement 24h prior to intravaginal pessary withdrawal on reproductive performance of seasonal anestrous ewes. Ewes in each experiment were allocated to treatment and control and all were induced to estrus using either intravaginal MAP (Exp. 1; n=24) or CIDR-G (Exp. 2; n=28) pessaries for 12 days. Half of the ewes in each experiment were supplemented 24h before withdrawal of pessaries with either 10mg oral MAP tablets (Exp. 1) or 25mg i.m. progesterone (P(4)) administration (Exp. 2; P(4)-supplement-treated group). Fertile rams were allowed with the ewes at sponge removal (Day 0, 0h) and estrus was monitored at 6-h intervals for 3 days. Blood samples were collected for measurements of P(4) (Exp. 1 and Exp. 2) and LH (Exp. 2). In both experiments, the percent of ewes in estrus was greater (P<0.05) and intervals to estrus were longer (P<0.05) in progestagen-supplement-treated than control ewes. In Exp. 2, the occurrence and magnitude of LH surges were greater (P<0.01) and intervals to onset of LH surge were longer (P<0.01) in P(4)-supplement-treated than control ewes. In Exp. 2, P(4) supplement elevated P(4) levels from 1.8+/-0.1ng/mL on Day -1 to 4.2+/-0.3 on Day 0 (P<0.001). Following pessaries removal, P(4) concentrations fell to basal values on Day 1 in both groups and remained low until Day 5. Then, P(4) concentrations increased and remained elevated through Day 19 in all (100%) progestagen-supplement-treated in Exp. 1 (12/12) and Exp. 2 (14/14) and in only 5/12 (41.7%) and 6/14 (42.9%) control ewes, respectively. These ewes were confirmed pregnant by ultrasonography and lambed on Day 149.2+/-0.2 following Day 0. In conclusion, progestagen supplement 24h prior to removal of pessary can be used successfully to improve reproductive performance of ewes bred out-of-season.     

A map of pheromone receptor activation in the mammalian brain

In mammals, the detection of pheromones is mediated by the vomeronasal system. We have employed gene targeting to visualize the pattern of projections of axons from vomeronasal sensory neurons in the accessory olfactory bulb. Neurons expressing a specific receptor project to multiple glomeruli that reside within spatially restricted domains. The formation of this sensory map in the accessory olfactory bulb and the survival of vomeronasal organ sensory neurons require the expression of pheromone receptors. In addition, we observe individual glomeruli in the accessory olfactory bulb that receive input from more than one type of sensory neuron. These observations indicate that the organization of the vomeronasal sensory afferents is dramatically different from that of the main olfactory system, and these differences have important implications for the logic of olfactory coding in the vomeronasal organ.     

Discrimination of conspecific sex and reproductive condition using chemical cues in axolotls (<Emphasis Type="Italic">Ambystoma mexicanum</Emphasis>)

Chemosensory cues play an important role in the daily lives of salamanders, mediating foraging, conspecific recognition, and territorial advertising. We investigated the behavioral effects of conspecific whole-body odorants in axolotls, Ambystoma mexicanum, a salamander species that is fully aquatic. We found that males increased general activity when exposed to female odorants, but that activity levels in females were not affected by conspecific odorants. Although males showed no difference in courtship displays across testing conditions, females performed courtship displays only in response to male odorants. We also found that electro-olfactogram responses from the olfactory and vomeronasal epithelia were larger in response to whole-body odorants from the opposite sex than from the same sex. In males, odorants from gravid and recently spawned females evoked different electro-olfactogram responses at some locations in the olfactory and vomeronasal epithelia; in general, however, few consistent differences between the olfactory and vomeronasal epithelia were observed. Finally, post hoc analyses indicate that experience with opposite-sex conspecifics affects some behavioral and electrophysiological responses. Overall, our data indicate that chemical cues from conspecifics affect general activity and courtship behavior in axolotls, and that both the olfactory and vomeronasal systems may be involved in discriminating the sex and reproductive condition of conspecifics.Abbreviations EOG electro-olfactogram - VNO vomeronasal organ