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Direct and indirect mate choice on leks.

Indirect mate choice is any behavior that restricts the individual's set of potential mates without discrimination of mate attributes directly, for example, by having preferences about where to mate. We analyzed a 14-year data set from great snipe (Gallinago media) leks for evidence of indirect mate choice based on relative and absolute position of lek territories. We found little or no effect of the centrality of territories on mating and no between-year consistency in the spatial distribution of matings within leks. Instead, the probability of matings occurring at a particular site increased if the current territory owner had mated the previous year. Furthermore, individual females returned in later seasons to mate with the same male as previously rather than at the same site. Previous work found that male interactions and dominance do not control matings and that females are very choosy about which territory they mate in. Here we show that this is because of the male occupying the territory rather than its position. We therefore conclude that direct female mate choice is the main behavioral process affecting variation in mating success among great snipe males, unlike in some lekking mammals where male competition and/or indirect mate choice appears more important.

Animals↗

Models of sexual selection on a quantitative genetic trait when preference is acquired by sexual imprinting.

The evolution of a quantitative genetic trait under stabilizing viability selection and sexual selection is modeled for a polygynous species in which female mating preferences are acquired by sexual imprinting on the parents and by exposure to the surviving population at large. Stabilizing viability selection acts equally on both sexes in the case of a sexually monomorphic trait and on males only in the case of a dimorphic trait. A genetically fixed sensory or perceptual bias defines the origin of the scale on which the trait is measured, and the possibility is incorporated that female preferences may deviate asymmetrically from the familiar-either toward or away from this origin. When viability selection is strong relative to sexual selection, the models predict that the mean trait value will evolve to the viability optimum. With intermediate ratios of the strength of viability to sexual selection, a stable equilibrium can occur on either side of this viability optimum, depending on the direction of asymmetry in female preferences. When viability selection is relatively weak and certain other conditions are also satisfied, runaway selection is predicted.

Animals↗

Mating preference functions of individual female barking treefrogs, Hyla gratiosa, for two properties of male advertisement calls.

A mating preference function describes the relationship between variation in a trait in potential mates and the strength of the preference for that trait. Few studies have measured mating preference functions either at a population level or for individuals. We used two-choice playback experiments to determine the mating preference functions of individual female barking treefrogs for two call characteristics: call-repetition rate and fundamental frequency. We tested each female four times with each pair of stimuli and with three to six pairs of stimuli. Individual females exhibited directional preferences for higher call rates and stabilizing preferences for intermediate fundamental frequency. These individual preference functions were similar to population-level preferences documented in previous studies. Within a stimulus type (call rate or fundamental frequency), nearly all females exhibited the same general preference function. However, females varied in the minimum difference between stimuli necessary to elicit a unanimous choice for the higher call rate, and they differed in both the intermediate fundamental frequency they preferred most and the minimum difference in fundamental frequency that elicited a unanimous choice for one of the two alternatives. The variation we observed among females was not repeatable; repeatabilities were in general low and statistically nonsignificant. The observed variation in the preferences of females, even if unrepeatable, should weaken selection on male traits relative to selection in the absence of such variation.

Animals↗

Evolution of a percutaneous fetoscopic access system for single-port tracheal occlusion.

BACKGROUND/PURPOSE: Prenatal tracheal occlusion currently is being assessed as a treatment modality for congenital diaphragmatic hernia (CDH). The development of a totally percutaneous fetoscopic access system would help avoid the need for maternal laparotomy and reduce the morbidity rate of fetal surgical procedures for the mother. Laparoscopic radial expansion sheaths and Seldinger technique-based vascular catheters both have been advocated as means of achieving amniotic cavity access. The authors have investigated these 2 systems in an attempt to develop a reliable method for achieving safe percutaneous fetoscopic access and present the first successful attempt to deploy an intratracheal balloon using an entirely percutaneous approach through a single port in an ovine model. METHODS: A number of prototype systems were evaluated sequentially over a 3-year period in an ovine model: (1) the radially expanding InnerDyne step port system, (2) a new rigid cannula with a bulbous/sharp end preloaded onto the radially expanding InnerDyne port, (3) a conical removable addition to the rigid cannula in 2, (4) a modified bulbous/sharp ended cannula incorporating a circumferential protective insert, (5) a rigid split sheath with the radially expanding port placed through the lumen of the split sheath, (6) a flexible introducer and dilator with the split sheath (used in the Seldinger placement of central lines), and (7) a 2-needle approach using a superelastic shape-memory alloy Nickel-Titanium wire with the flexible dilator and sheath, incorporating a side perfusion port. For balloon tracheal occlusion, live anaesthetized time-mated pregnant ewes were used at 110 days' gestation. Tracheobronchoscopy was achieved using a 3-mm 0 degrees telescope, and the cutaneotracheal tract was secured by a 3.3-mm sheath incorporating a side-perfusion port. The rigid telescope was replaced by a flexible choledochoscope preloaded with a silicone balloon. The balloon was deployed 2 cm above the carina proximal to the right upper lobe bronchus. RESULTS: The many problems encountered in the evolution of the preferred system related mainly to separation and tenting of the chorioamniotic membranes in the ovine uterus and inconsistent access to the fetal parts of interest. Each resulted in significant modifications to our approach. Furthermore, the use of rigid access devices commonly caused fetal injury. Successful access to the intrauterine cavity and cannulation of the trachea was achieved consistently with minimal trauma, irrespective of fetal position by method 7. Multiple port placement allowed visualization of the entry of all components of the system confirming minimal chorioamniotic membrane separation and tenting. Single port tracheal occlusion was undertaken first on 6 cadavers before being performed successfully on 3 live anaesthetized ewes. Fetoscopic access and cannulation of the trachea was achieved consistently in all live animals irrespective of fetal position. CONCLUSIONS: This modified Seldinger technique using the unique properties of the memory-shaped alloy wire for initial uterine access offers a safe method for the percutaneous placement of fetoscopic ports in the ovine model for prenatal intervention. Successful placement of a tracheal balloon entirely through a single percutaneously placed port represents a further advance in prenatal therapy for CDH.

Animals↗

Dopamine activates masculine sexual behavior independent of the estrogen receptor alpha.

Estrogen receptor alpha (ERalpha) is believed to be a critical part of the regulatory processes involved in normal reproduction and sexual behavior. However, in this study we show the ERalpha is not required for display of masculine sexual behavior. Male and female, ERalpha knock-out (ERalphaKO) and wild-type mice were gonadectomized and implanted with testosterone. Sexual behavior and social preferences were tested after injection of the dopamine agonist, apomorphine (APO), or vehicle. All wild-type mice showed normal masculine behavior, including mounts and pelvic thrusts in females, and ejaculation in males. In agreement with past reports, ERalphaKO mice, given vehicle, failed to show mating behavior. Yet, ERalphaKO males given APO showed masculine copulatory behavior and chemoinvestigatory behavior directed at females. ERalphaKO females, treated with APO, mounted and thrusted when tested with receptive females. HPLC revealed that wild-type and ERalphaKO mice had equivalent catecholamine content in brain regions associated with masculine sexual behavior. These data show that the ERalpha is not essential during development or adulthood for the expression of masculine sexual behavior in mice. Moreover, dopamine can activate sexual behavior via a mechanism that either acts on an ER other than ERalpha or via an estrogen-independent pathway.

Animals↗

Voluntary beer drinking by pregnant rats: offspring sensitivity to ethanol and preference for beer.

Female Long-Evans rats were allowed voluntary access to beer, food and water for three weeks prior to mating and throughout gestation; and were compared to controls. Offspring were tested for sensitivity to ethanol and preference for beer at 29 and 85 days of age. Offspring of beer drinkers had long-term alterations in sensitivity to ethanol as adults, although rates of ethanol metabolism were unaffected. The nature of responsivity to ethanol as adults was sex-dependent. Male offspring of beer drinkers were delayed in maintaining baseline body temperature at 29 days with recovery at 85 days; and were tolerant to the hypothermic effects of ethanol at 29 and 85 days. Female offspring had prolonged latency to respond to pain and temperature; and were tolerant to the effects of ethanol on motor coordination at 29 and 85 days. Although preference of the offspring for beer was not affected by maternal beer drinking, the pattern of fluid intake by offspring of beer drinkers differed from controls.

Alcohol Drinking↗

Evidence that sensory traps can evolve into honest signals.

Conventional models explaining extreme sexual ornaments propose that these reflect male genetic quality or are arbitrary results of genetic linkage between female preference and the ornament. The chase-away model emphasizes sexual conflict: male signals attract females because they exploit receiver biases. As males gain control of mating decisions, females may experience fitness costs through suboptimal mating rates or post-copulatory exploitation. Elaboration of male signals is expected if females increase their response threshold to resist such exploitation. If ornaments target otherwise adaptive biases such as feeding responses, selection on females might eventually separate sexual and non-sexual responses to the signal. Here we show that the terminal yellow band (TYB) of several Goodeinae species evokes both feeding and sexual responses; sexual responsiveness phylogenetically pre-dates the expression of the TYB in males and is comparable across taxa, yet feeding responsiveness decreases in species with more elaborated TYBs. Displaying a TYB is costly, and thus provides an example where a trait arose as a sensory trap but has evolved into an honest signal.

Animals↗

Drosophila song as a species-specific mating signal and the behavioural importance of Kyriacou & Hall cycles in D. melanogaster song.

'Species-specific' mating signals may not make a significant contribution to sexual isolation if differences between species are not matched by narrowly tuned differences in female preferences. The courtship song of the melanogaster species group of Drosophila has been studied by several researchers as a potential factor in sexual isolation, but there are few clear demonstrations that species differences are behaviourally significant. We played synthetic song with typical pulse interval parameters for D. melanogaster, D. simulans or D. sechellia to D. melanogaster or D. simulans females in the presence of mute males. Females mated most quickly when stimulated by song typical of their own species, confirming that song can influence sexual isolation between these species. This effect was greater in D. melanogaster than D. simulans, which correlates with reduced variability in mean interpulse interval within D. melanogaster. We also examined the role of 'Kyriacou & Hall cycles' (periodic cycling of mean interpulse intervals) in D. melanogaster. Contrary to some predictions, the presence of the Kyriacou & Hall cycle was more important in increasing mating speed than the increased variability of interpulse intervals that is inevitably present in song containing a cycle. The large stimulatory effect of cycles on mating speed is curious, given that the periodicity of cycles would probably be difficult for females to detect during courtship in field conditions. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Effects of maternal exposure to a reconstituted mixture of polychlorinated biphenyls on sex-dependent behaviors and steroid hormone concentrations in rats: dose-response relationship.

In a previous experiment, maternal exposure to a polychlorinated biphenyl (PCB) mixture reconstituted according to the congener pattern found in human breast milk resulted in decreased aromatase activity in the brain of newborn male rats, together with feminization of sweet preference behavior in adult male littermates. Both mixtures led to similar reductions of serum testosterone and testes weights. The purpose of the present study was (1) to examine the dose-response relationship for the reconstituted mixture and (2) to study if the rewarding properties of testosterone are affected at levels sufficient to alter sweet preference behavior. Female rats were fed diets with 0, 5, 20, or 40 mg PCBs/kg diet, resulting in an average daily intake of 0, 0.5, 2, or 4 mg/kg body wt. Exposure started 50 days prior to mating and was continued until birth of the offspring. A dose-dependent elevation of sweet preference was found in adult male offspring, indicating feminization of this sexually dimorphic behavior. Examination of conditioned place preference revealed a preference for the testosterone-paired side at the highest exposure condition. In weanling female offspring, dose-dependent reductions of serum testosterone and estradiol concentrations were detected. In addition, testosterone concentrations were reduced in a dose-dependent manner in adult male littermates long after termination of exposure. PCB concentrations in adipose tissue from offspring of the low dose group (0.5 mg/kg body wt) were approximately 10 times higher than values at the upper margin of current human exposure. Taken together, results indicate long-lasting and dose-dependent changes in sex-dependent behaviors and levels of sex steroid hormones in rats following developmental exposure to a PCB mixture that resembles the breast milk pattern.

Animals↗

How does offspring quality change with age in male Drosophila melanogaster?

We examined the relationship of genetic quality to age in male Drosophila melanogaster to test two contrasting hypotheses. The traditional hypothesis is that older males have proven their viability and therefore produce offspring of superior genetic quality. This hypothesis is often evoked as an explanation for female preference for older mates. In contrast, we have recently argued that older fathers may produce offspring of inferior genetic quality. Here, we present results from an experiment designed to measure the genetic quality of offspring produced by 2 day old, 2 week old and 5 week old male D. melanogaster. We found a statistically significant small reduction in larval viability and a similar but statistically non-significant reduction in son mating ability among the offspring of the 5 week old males. Daughter fecundity showed no apparent trend for a reduction nor an increase in performance with increasing age of the fathers. There was no evidence of a difference between the 2 day old and the 2 week old males for any of these three fitness components. These results are in somewhat better accordance with our alternative hypothesis, but the relatively weak and late occurring effects indicate that mate choice based on age may not be a viable strategy in this population.

Aging↗

The reinforcement of mating preferences on an island.

We develop a haploid model for the reinforcement of female mating preferences on an island that receives migrants from a continent. We find that preferences will evolve to favor island males under a broad range of conditions: when the average male display trait on the island and continent differ, when the preference acts on that difference, and when there is standing genetic variance for the preference. A difference between the mean display trait on the continent and on the island is sufficient to drive reinforcement of preferences. Additional postzygotic isolation, caused, for example, by either epistatic incompatibility or ecological selection against hybrids, will amplify reinforcement but is not necessary. Under some conditions, the degree of preference reinforcement is a simple function of quantities that can be estimated entirely from phenotypic data. We go on to study how postzygotic isolation caused by epistatic incompatibilities affects reinforcement of the preference. With only one pair of epistatic loci, reinforcement is enhanced by tighter linkage between the preference genes and the genes causing hybrid incompatibility. Reinforcement of the preference is also affected by the number of epistatically interacting genes involved in incompatibility, independent of the overall intensity of selection against hybrids.

Animals↗

The major histocompatibility complex and the chemosensory signalling of individuality in humans.

The chemosensory identity of mice and rats is determined partly by polymorphic genes of the major histocompatibility complex (MHC). In inbred strains of mice, as well as in seminatural populations, MHC-associated mating preferences selectively influence reproductive success, thus serving to promote heterozygocity in the MHC. In order to determine whether MHC-associated chemosignals are present in humans, two studies were conducted. In a first study, olfactory identification of MHC-associated chemosignals was conducted on 12 trained rats' responses to the urine odors of humans. In a second study, MHC-associated olfactory cues in humans were analyzed by means of gas chromatography. The results indicate that the urine odors of humans are associated with the MHC and demonstrate that the profile of volatile components in the urine odors shows some association with the MHC. Furthermore, results show that a profile of some specific components, as well as a few ubiquitous volatiles, constitutes MHC-associated odor signals in humans.

Animals↗

Estimating quantitative genetic parameters in haplodiploid organisms.

Haplodiploid organisms (diploid females and haploid males) are genetically distinct in that they have asymmetrical genetic segregation, gene dosage compensation in the haploid male, maternal effects, and a general propensity to inbreed. Taking these characteristics into account, we demonstrate how conventional experimental designs and statistics derived to measure quantitative genetic parameters for diploid organisms can be applied to haplodiploid ones. First, the C matrix of the joint-scaling test is modified to test the additive-dominance model and then the calculations for fractions in the C matrices (for males and females) are shown for an infinite number of generations with both random and brother-sister mating. Second, analytical adjustments are outlined for the derivation of covariance between relatives using populations at either Hardy-Weinberg equilibrium or those that prefer to inbreed. Finally, four of the conventional designs (sib-analysis, offspring-parental regression, North Carolina III (NC III), and diallel cross) are modified and then compared according to their ability to deal with the experimental conditions encountered in haplodiploid systems. Although all designs can be used with caution, our analysis suggests that the NC III design is the most broadly applicable because it consistently meets the theoretical assumptions. The parameters we derive here for inbred populations are recommended for analysing the typically skewed genotypic distributions found in many natural populations of haplodiploid organisms. The theoretical and applied aspects of our work as well as the difference between our work and that for X-linked genes are discussed.

Animals↗

Genetic benefits of mate choice separated from differential maternal investment in red junglefowl (Gallus gallus).

Females may choose more attractive mates to obtain better viability or attractiveness genes for their offspring. A number of studies have demonstrated a positive relationship between paternal attractiveness and offspring quality. However, this pattern could be due to inheritance of paternal genes and/or it could be due to increased maternal investment in the offspring of more attractive males. To isolate female responses to male appearance from paternal genetic effects, I housed female red junglefowl (Gallus gallus) with vasectomized (sterile) males and artificially inseminated them. Male junglefowl with larger combs are more attractive to females. Females laid more eggs when housed with a large-combed, as opposed to a small-combed, vasectomized mate. Neither egg volume nor offspring body condition was associated with comb size of the mother's vasectomized mate. Paternal genetics appeared important. Body condition and comb size were greater for the sons of large-combed sperm donor males. This is consistent with the hypothesis that genetic benefits to offspring maintain female preference for the most ornate males. It is possible that greater body condition and comb size in sons of large-combed sires was not caused by genetic differences, but instead was due to compounds in the ejaculate of large-combed sperm donors inducing greater reproductive investment from females. However, females artificially inseminated by large-combed males did not produce more or larger eggs than females artificially inseminated by small-combed males, and thus there is no other evidence consistent with ejaculate-induced differential investment. Furthermore, only in older chicks was body condition significantly related to sire comb size, suggesting genetic rather than differential investment mechanisms.

Analysis of Variance↗

Runaway sexual selection with paternal transmission of the male trait and gene-culture determination of the female preference.

Sexual selection is modeled with a male viability-reducing trait and a female mating preference for that trait both of which are culturally transmitted. Both the male trait and the female preference are transmitted only between same-sex individuals, so that non-random association between the trait and the preference, which would give rise to a Fisherian runaway process, cannot arise. Inclusion of an autosomal gene that confers a female predisposition to acquire a certain preference is shown to allow the coevolution of the male trait and the female preference by a Fisherian process. This holds true even when the female preference has a slight viability cost, provided the male cultural transmission is not perfect. It is also suggested that a Fisherian process can be more easily initiated in these models than in the conventional genetic models. Furthermore, a Fisherian process may cause cultural transmission of female preference to evolve. Additionally, polymorphism can be maintained at the predisposition locus if heterozygous females have a stronger predisposition to acquire the preference than homozygotes. Our models may be applicable to the case when the male trait is a Y-linked genetic or environmentally determined trait.

Animals↗

Flower choices by raspberry beetles: cues for feeding and oviposition.

Raspberry beetles, Byturus tomentosus, use raspberry as the host plant for feeding, mating and oviposition; the larvae are a major pest on the fruit. This study seeks reasons for their choices between flowers of different cultivars. Beetles showed consistent diurnal movements between different cultivars of raspberry, which were linked to differential behavioural patterns; they fed predominantly on Glen Clova canes in the morning and moved to Glen Prosen later in the day where oviposition behaviour was commoner and substantially more effective. Hence two closely similar cultivars of the same plant species received very different levels of flower damage (from adult feeding) and of fruit damage (from larval infestation). These patterns were linked to subtle differences in cues from the host plant; Glen Clova produces unusually copious nectar which may limit oviposition opportunities because of osmotic threats to eggs and larvae, whereas Glen Prosen has drier flowers for egg laying and apparently emits a short-lived volatile cue that principally attracts female beetles. Selective breeding of the host plant has therefore had unexpected side-effects on interactions with a major pest species, mediated by the behaviour of the insect, and perhaps accentuating natural behavioural preferences that serve to reduce competition between adult and larval stadia. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Duet-splitting and the evolution of gibbon songs.

Unlike the great apes and most other primates, all species of gibbons are known to produce elaborate, species-specific and sex-specific patterns of vocalisation usually referred to as "songs". In most, but not all, species, mated pairs may characteristically combine their songs in a relatively rigid pattern to produce coordinated duet songs. Previous studies disagree on whether duetting or the absence of duetting represented the primitive condition in gibbons. The present study compares singing behaviour in all gibbon species. Various vocal characteristics were subjected to a phylogenetic analysis using previously published phylogenetic trees of the gibbon radiation as a framework. Variables included the degree of sex-specificity of the vocal repertoire, the occurrence of solo songs, and the preference for a specific time of day for song-production. The results suggest the following scenario for the evolution of gibbon songs: (1) The last common ancestor of recent gibbons produced duet songs. (2) Gibbon duets probably evolved from a song which was common to both sexes and which only later became separated into male-specific and female-specific parts (song-splitting theory). (3) A process tentatively called "duet-splitting" is suggested to have led secondarily from a duetting species to a non-duetting species, in that the contributions of the pair-partners split into temporally segregated solo songs. This appears to be the first time that a non-duetting animal can be shown to be derived from a duetting form. (4) The return to exclusive solo singing may be related to the isolated island distribution of the non-duetting species.

Acoustics↗

Selective breeding for variations in patterns of mystacial vibrissae of mice. Bilaterally symmetrical strains derived from ICR stock.

The establishment of certain patterns of mystacial vibrissae in mice has been the aim of an extensive breeding program carried on in this laboratory since 1977. In a companion paper we have reported on variations in this pattern in an outbred population of ICR mice. Starting with 21 ICR animals we bred, mostly by brother-sister mating, for 13 bilaterally symmetric patterns of mystacial vibrissae characterized by the presence (or absence) of supernumerary whiskers (SWs). The strains are classified as follows: I, a mouse strain with the standard pattern; II, eight strains bred for the occurrence of SWs at a given site or sites; and III, four mouse strains bred for a maximal number of SWs in different regions of the whiskerpad. Commonly, SWs occur in regions that coincide with the zones of mergence between the three facial processes except for two class II strains in which we bred for SWs in the "straddler" row of vibrissae, and for one class III strain, in which we cultivated the tendency (that appeared late in our program) to have SWs at the crest of a facial process. For classes I and II we analyzed the results for about 18 generations in terms of "improvement," meaning an increase in the percentages of animals with the desired phenotype together with a decreased frequency of undesired SWs. For class III, success in breeding meant the increase of the mean number of the desired SWs. All results led to the same conclusion: there is a genetic basis for the occurrence of SWs. The side preference of a particular SW is not strain dependent. It disappears in those class I and II strains in which almost 100% of animals obtained the desired phenotype. The increase in number of SWs in one zone of mergence does not depend on the presence of SWs in the other. Where tested, we almost always found a representation of an SW in a topologically equivalent location within the "barrelfield" area of the somatosensory cerebral cortex. Except for some diseases early in the breeding program, and some side effects of inbreeding that were eliminated, the population was without obvious defects. Where tested, there was no correlation between the occurrence of SWs and sex. The observed variations in pattern of mystacial vibrissae and their genetic background led us to propose a morphogenetic model for the formation of the pattern of mystacial vibrissae.

Animals↗