Polymorphisms in a desaturase 2 ortholog associate with cuticular hydrocarbon and male mating success variation in a natural population of Drosophila serrata

Elucidating the nature of genetic variation underlying both sexually selected traits and the fitness components of sexual selection is essential to understanding the broader consequences of sexual selection as an evolutionary process. To date, there have been relatively few attempts to connect the genetic variance in sexually selected traits with segregating DNA sequence polymorphisms. We set out to address this in a well-characterized sexual selection system - the cuticular hydrocarbons (CHCs) of Drosophila serrata - using an indirect association study design that allowed simultaneous estimation of the genetic variance in CHCs, sexual fitness and single nucleotide polymorphism (SNP) effects in an outbred population. We cloned and sequenced an ortholog of the D. melanogaster desaturase 2 gene, previously shown to affect CHC biosynthesis in D. melanogaster, and associated 36 SNPs with minor allele frequencies > 0.02 with variance in CHCs and sexual fitness. Three SNPs had significant multivariate associations with CHC phenotype (q-value < 0.05). At these loci, minor alleles had multivariate effects on CHCs that were weakly associated with the multivariate direction of sexual selection operating on these traits. Two of these SNPs had pleiotropic associations with male mating success, suggesting these variants may underlie responses to sexual selection due to this locus. There were 15 significant male mating success associations (q-value < 0.1), and interestingly, we detected a nonrandom pattern in the relationship between allele frequency and direction of effect on male mating success. The minor-frequency allele usually reduced male mating success, suggesting a positive association between male mating success and total fitness at this locus.

Load estimation with uncertainties from opportunistic sampling data: A semiparametric approach

We consider estimating the total load from frequent flow data but less frequent concentration data. There are numerous load estimation methods available, some of which are captured in various online tools. However, most estimators are subject to large biases statistically, and their associated uncertainties are often not reported. This makes interpretation difficult and the estimation of trends or determination of optimal sampling regimes impossible to assess. In this paper, we first propose two indices for measuring the extent of sampling bias, and then provide steps for obtaining reliable load estimates that minimizes the biases and makes use of informative predictive variables. The key step to this approach is in the development of an appropriate predictive model for concentration. This is achieved using a generalized rating-curve approach with additional predictors that capture unique features in the flow data, such as the concept of the first flush, the location of the event on the hydrograph (e.g. rise or fall) and the discounted flow. The latter may be thought of as a measure of constituent exhaustion occurring during flood events. Forming this additional information can significantly improve the predictability of concentration, and ultimately the precision with which the pollutant load is estimated. We also provide a measure of the standard error of the load estimate which incorporates model, spatial and/or temporal errors. This method also has the capacity to incorporate measurement error incurred through the sampling of flow. We illustrate this approach for two rivers delivering to the Great Barrier Reef, Queensland, Australia. One is a data set from the Burdekin River, and consists of the total suspended sediment (TSS) and nitrogen oxide (NO(x)) and gauged flow for 1997. The other dataset is from the Tully River, for the period of July 2000 to June 2008. For NO(x) Burdekin, the new estimates are very similar to the ratio estimates even when there is no relationship between the concentration and the flow. However, for the Tully dataset, by incorporating the additional predictive variables namely the discounted flow and flow phases (rising or recessing), we substantially improved the model fit, and thus the certainty with which the load is estimated.

Human mating in the informal market for sperm donation: Preferences and decision making

Choosing a mate is one of the largest (economic) decisions humans make. This thesis investigates this large scale decision and how the process is changing with the advent of the internet and the growing market for online informal sperm donation. This research identifies individual factors that influence female mating preferences. It explores the roles of behavioural traits and physical appearance, preferences for homogamy and hypergamy, and personality, and how these impact the decision to choose a donor. Overall, this thesis makes contributions to both the literature on human behaviour, and that on decision-making in extreme and highly important situations.

Effects of methyl eugenol feeding on mating compatibility of Asian population of Bactrocera dorsalis (Diptera: Tephritidae) with African population and with B. carambolae

Males of some species included in the Bactrocera dorsalis complex are strongly attracted to methyl eugenol (ME) (1,2-dimethoxy-4-(2-propenyl) benzene), a natural compound occurring in a variety of plant species. ME feeding of males of the B. dorsalis complex is known to enhance their mating competitiveness. Within B. dorsalis, recent studies show that Asian and African populations of B. dorsalis are sexually compatible, while populations of B. dorsalis and Bactrocera carambolae are relatively incompatible. The objectives of this study were to examine whether ME feeding by males affects mating compatibility between Asian and African populations of B. dorsalis and ME feeding reduces male mating incompatibility between B. dorsalis (Asian population) and B. carambolae. The data confirmed that Asian and African populations of B. dorsalis are sexually compatible for mating and showed that ME feeding only increased the number of matings. Though ME feeding also increased the number of matings of B. dorsalis (Asian population) and B. carambolae males but the sexual incompatibility between both species was not reduced by treatment with ME. These results conform to the efforts resolving the biological species limits among B. dorsalis complex and have implications for fruit fly control programs in fields and horticultural trade.

Effects of laboratory colonization on Bactrocera dorsalis (Diptera, Tephritidae) mating behaviour: ‘what a difference a year makes

Laboratory-reared insects are widely known to have significantly reduced genetic diversity in comparison to wild populations; however, subtle behavioural changes between laboratory-adapted and wild or ‘wildish’ (i.e., within one or very few generations of field collected material) populations are less well understood. Quantifying alterations in behaviour, particularly sexual, in laboratory-adapted insects is important for mass-reared insects for use in pest management strategies, especially those that have a sterile insect technique component. We report subtle changes in sexual behaviour between ‘wildish’ Bactrocera dorsalis flies (F1 and F2) from central and southern Thailand and the same colonies 12 months later when at six generations from wild. Mating compatibility tests were undertaken under standardised semi-natural conditions, with number of homo/heterotypic couples and mating location in field cages analysed via compatibility indices. Central and southern populations of B. dorsalis displayed positive assortative mating in the 2010 trials but mated randomly in the 2011 trials. ‘Wildish’ southern Thailand males mated significantly earlier than central Thailand males in 2010; this difference was considerably reduced in 2011, yet homotypic couples from southern Thailand still formed significantly earlier than all other couple combinations. There was no significant difference in couple location in 2010; however, couple location significantly differed among pair types in 2011 with those involving southern Thailand females occurring significantly more often on the tree relative to those with central Thailand females. Relative participation also changed with time, with more southern Thailand females forming couples relative to central Thailand females in 2010; this difference was considerably decreased by 2011. These results reveal how subtle changes in sexual behaviour, as driven by laboratory rearing conditions, may significantly influence mating behaviour between laboratory-adapted and recently colonised tephritid fruit flies over a relatively short period of time.

The mating system of the true fruit fly Bactrocera tryoniand its sister-species,Bactrocera neohumeralis

The frugivorous ‘true’ fruit fly, Bactrocera tryoni (Queensland fruit fly), is presumed to have a non-resourced-based lek mating system. This is largely untested, and contrary data exists to suggest Bactrocera tryoni may have a resource-based mating system focused on fruiting host plants. We tested the mating system of Bactrocera tryoni, and its close sibling Bactrocera neohumeralis, in large field cages using laboratory reared flies. We used observational experiments that allowed us to determine if: (i) mating pairs were aggregated or non-aggregated; (ii) mating system was resource or non-resource based; (iii) flies utilised possible landmarks (tall trees over short) as mate-rendezvous sites; and (iv) males called females from male-dominated leks. We recorded nearly 250 Bactrocera tryoni mating pairs across all experiments, revealing that: (i) mating pairs were aggregated; (ii) mating nearly always occurred in tall trees over short; (iii) mating was non-resource based; and (iv) that males and females arrived at the mate-rendezvous site together with no evidence that males preceded females. Bactrocera neohumeralis copulations were much more infrequent (only 30 mating pairs in total), but for those pairs there was a similar preference for tall trees and no evidence of a resource-based mating system. Some aspects of Bactrocera tryoni mating behaviour align with theoretical expectations of a lekking system, but others do not. Until evidence for unequivocal female choice can be provided (as predicted under a true lek), the mating system of Bactrocera tryoni is best described as a non-resource based, aggregation system for which we also have evidence that land-marking may be involved. This article is protected by copyright. All rights reserved

Effects of 17α-ethinyl estradiol on the mating system of the sand goby (Pomatoschistus minutus)

In aquatic environments, endocrine disrupting chemicals (EDCs) that interfere with the endocrinology of males and females form a threat to the maintenance of populations. EDCs are a diverse group of natural and manmade chemicals that already at very low concentrations (at nanogram levels) can have severe effects on reproduction by individuals, e.g. complete sex reversal, feminisation of males, impaired reproduction even resulting in near extinction of populations. With regard to fish, despite the extensive literature on physiological effects of EDCs, very little is known about potential population-level effects. In this thesis, I examined how 17α-ethinyl estradiol (EE2), a synthetic estrogen used in oral contraceptive pills, affects the reproductive behaviour of a marine fish, the sand goby (Pomatoschistus minutus). The aims were fourfold. First, I investigated how exposure to EE2 affects courtship and parental care of sand goby males. Secondly, I looked at effects on the mating system and sexual selection. In the third study, I observed the effects of exposure in a social context where exposed males had to compete with non-exposed males for resources and mates. Finally, I studied the effects of exposure on male-male competition and male aggressive behaviour. This work revealed that EE2 exposure impairs the ability of males to acquire and defend a nest, as well as diminishes the attractiveness of males to females by decreasing their courtship and aggressive behaviour. These effects are harmful for a male whose reproductive success is determined by the ability to compete for limited resources and to attract mates. Furthermore, this thesis showed that selection on male size was relaxed after EE2 exposure and male size had a smaller effect on mating success. These effects can be of a profound nature as they interfere with sexual selection, and may in the long run lead to the loss of traits maintained through sexual selection. The thesis shows that an exposure to environmentally relevant levels of EE2 clearly reduces the chances of individuals to reproduce successfully. Furthermore, it strongly suggests that several types of biomarkers should be used to detect and assess the effects of EDC exposure because severe behavioural effects can sometimes be seen before effects are detectable at the molecular or morphometric level. Behavioural assays should be considered an important complementary tool for the standard ecotoxicological assays because observed behavioural changes have direct and negative effects on fitness, while the connection between changes in molecular expression and fitness may be less obvious.

The mating system of the true fruit fly Bactrocera tryoni and its sister-species, Bactrocera neohumeralis

The frugivorous 'true' fruit fly, Bactrocera tryoni (Queensland fruit fly), is presumed to have a non-resourced-based lek mating system. This is largely untested, and contrary data exists to suggest Bactrocera tryoni may have a resource-based mating system focused on fruiting host plants. We tested the mating system of Bactrocera tryoni, and its close sibling Bactrocera neohumeralis, in large field cages using laboratory reared flies. We used observational experiments that allowed us to determine if: - (i) mating pairs were aggregated or non-aggregated; - (ii) mating system was resource or non-resource based; - (iii) flies utilised possible landmarks (tall trees over short) as mate-rendezvous sites, and; - (iv) males called females from male-dominated leks. We recorded nearly 250 Bactrocera tryoni mating pairs across all experiments, revealing that: - (i) mating pairs were aggregated; - (ii) mating nearly always occurred in tall trees over short; - (iii) mating was non-resource based, and; - (iv) that males and females arrived at the mate-rendezvous site together with no evidence that males preceded females. Bactrocera neohumeralis copulations were much more infrequent (only 30 mating pairs in total), but for those pairs there was a similar preference for tall trees and no evidence of a resource-based mating system. Some aspects of Bactrocera tryoni mating behaviour align with theoretical expectations of a lekking system, but others do not. Until evidence for unequivocal female choice can be provided (as predicted under a true lek), the mating system of Bactrocera tryoni is best described as a non-resource based, aggregation system for which we also have evidence that land-marking may be involved. This article is protected by copyright. All rights reserved.

Do SMEs cluster around innovation activities? Discovering active, incremental and opportunistic innovators

Do SMEs cluster around different types of innovation activities? Are there patterns of SME innovation activities? To investigate we develop a taxonomy of innovation activities in SMEs using a qualitative study, followed by a survey. First, based upon our qualitative research and literature review we develop a comprehensive list of innovation activities SMEs typically engage in. We then conduct a factor analysis to determine if these activities can be combined into factors. We identify three innovation activity factors: R&D activities, incremental innovation activities and cost innovation activities. We use these factors to identify three clusters of firms engaging in similar innovation activities: active innovators, incremental innovators and opportunistic innovators. The clusters are enriched by validating that they also exhibit significant internal similarities and external differences in their innovation skills, demographics, industry segments and family business ownership. This research contributes to innovation and SME theory and practice by identifying SME clusters based upon their innovation activities.

Assortative mating and the adaptive landscape

The ability of a population to shift from one adaptive peak to another was examined for a two-locus model with different degrees of assortative mating, selection, and linkage. As expected, if the proportion of the population that mates assortatively increases, so does its ability to shift to a new peak. Assortative mating affects this process by allowing the mean fitness of a population to increase monotonically as it passes through intermediate gene frequencies on the way to a new, higher, homozygotic peak. Similarly, if the height of the new peak increases or selection against intermediates becomes less severe, the population becomes more likely to shift to a new peak. Close linkage also helps the shift to a new adaptive peak and acts similarly to assortative mating, but it is not necessary for such a shift as was previously thought. When a population shifts to a new peak, the number of generations required is significantly less than that needed to return to the original peak when that happens. The short period of time required may be an explanation for rapid changes in the geological record. Under extremely high degrees of assortative mating, the shift takes longer, presumably because of the difficulty of breaking up less favored allelic combinations.

OPSM - Opportunistic Power Save mode for Infrastructure IEEE 802.11 WLAN

We focus on the energy spent in radio communication by the stations (STAs) in an IEEE 802.11 infrastructure WLAN. All the STAs are engaged in web browsing, which is characterized by a short file downloads over TCP, with short duration of inactivity or think time in between two file downloads. Under this traffic, Static PSM (SPSM) performs better than CAM, since the STAs in SPSM can switch to low power state (sleep) during think times while in CAM they have to be in the active state all the time. In spite of this gain, performance of SPSM degrades due to congestion, as the number of STAs associated with the access point (AP) increases. To address this problem, we propose an algorithm, which we call opportunistic PSM (OPSM). We show through simulations that OPSM performs better than SPSM under the aforementioned TCP traffic. The performance gain achieved by OPSM over SPSM increases as the mean file size requested by the STAs or the number of STAs associated with the AP increases. We implemented OPSM in NS-2.33, and to compare the performance of OPSM and SPSM, we evaluate the number of file downloads that can be completed with a given battery capacity and the average time taken to download a file.

Providing quality of service via opportunistic splitting using stochastic approximation

We consider the problem of wireless channel allocation to multiple users. A slot is given to a user with a highest metric (e.g., channel gain) in that slot. The scheduler may not know the channel states of all the users at the beginning of each slot. In this scenario opportunistic splitting is an attractive solution. However this algorithm requires that the metrics of different users form independent, identically distributed (iid) sequences with same distribution and that their distribution and number be known to the scheduler. This limits the usefulness of opportunistic splitting. In this paper we develop a parametric version of this algorithm. The optimal parameters of the algorithm are learnt online through a stochastic approximation scheme. Our algorithm does not require the metrics of different users to have the same distribution. The statistics of these metrics and the number of users can be unknown and also vary with time. Each metric sequence can be Markov. We prove the convergence of the algorithm and show its utility by scheduling the channel to maximize its throughput while satisfying some fairness and/or quality of service constraints.

Opportunistic splitting for scheduling via stochastic approximation

We consider the problem of scheduling a wireless channel among multiple users. A slot is given to a user with a highest metric (e.g., channel gain) in that slot. The scheduler may not know the channel states of all the users at the beginning of each slot. In this scenario opportunistic splitting is an attractive solution. However this algorithm requires that the metrics of different users form independent, identically distributed (iid) sequences with same distribution and that their distribution and number be known to the scheduler. This limits the usefulness of opportunistic splitting. In this paper we develop a parametric version of this algorithm. The optimal parameters of the algorithm are learnt online through a stochastic approximation scheme. Our algorithm does not require the metrics of different users to have the same distribution. The statistics of these metrics and the number of users can be unknown and also vary with time. We prove the convergence of the algorithm and show its utility by scheduling the channel to maximize its throughput while satisfying some fairness and/or quality of service constraints.

Opportunistic scheduling of wireless links

We consider the problem of scheduling of a wireless channel (server) to several queues. Each queue has its own link (transmission) rate. The link rate of a queue can vary randomly from slot to slot. The queue lengths and channel states of all users are known at the beginning of each slot. We show the existence of an optimal policy that minimizes the long term (discounted) average sum of queue lengths. The optimal policy, in general needs to be computed numerically. Then we identify a greedy (one step optimal) policy, MAX-TRANS which is easy to implement and does not require the channel and traffic statistics. The cost of this policy is close to optimal and better than other well-known policies (when stable) although it is not throughput optimal for asymmetric systems. We (approximately) identify its stability region and obtain approximations for its mean queue lengths and mean delays. We also modify this policy to make it throughput optimal while retaining good performance.

Opportunistic Scheduling in Cellular Systems in the Presence of Noncooperative Mobiles

A central scheduling problem in wireless communications is that of allocating resources to one of many mobile stations that have a common radio channel. Much attention has been given to the design of efficient and fair scheduling schemes that are centrally controlled by a base station (BS) whose decisions depend on the channel conditions reported by each mobile. The BS is the only entity taking decisions in this framework. The decisions are based on the reports of mobiles on their radio channel conditions. In this paper, we study the scheduling problem from a game-theoretic perspective in which some of the mobiles may be noncooperative or strategic, and may not necessarily report their true channel conditions. We model this situation as a signaling game and study its equilibria. We demonstrate that the only Perfect Bayesian Equilibria (PBE) of the signaling game are of the babbling type: the noncooperative mobiles send signals independent of their channel states, the BS simply ignores them, and allocates channels based only on the prior information on the channel statistics. We then propose various approaches to enforce truthful signaling of the radio channel conditions: a pricing approach, an approach based on some knowledge of the mobiles' policies, and an approach that replaces this knowledge by a stochastic approximations approach that combines estimation and control. We further identify other equilibria that involve non-truthful signaling.