Indirect population dynamic benefits of altered life-history trade-offs in response to egg harvesting

Variations in demographic rates due to differential resource allocation between individuals are important considerations in the development of accurate population dynamic models. Systematic harvesting can alter age structure and/or reduce population density, conferring indirect positive benefits on the source population as a result of a consequent redistribution of resources between the remaining individuals. Independently of effects mediated through changes in density and competition, demographic rates can also be influenced by within-individual competition for resources. Harvesting dependent life stages can reduce an individual's current reproductive costs, allowing increased investment in its future fecundity and survival. Although such changes in demographic rates are well known, there has been little exploration of the potential impact on population dynamics. We use empirical data collected from a successfully reintroduced population of the Mauritius kestrel Falco punctatus to explore the population consequences of manipulating reproductive effort through harvesting. Consequent increases in an individual's future fecundity and survival allow source populations to withstand longer and more intensive harvesting regimes without being exposed to an increase in extinction risk, increasing maximum sustainable yields. These effects may also buffer populations against the impacts of stochastic events, but directional shifts in environmental conditions that increase reproductive costs may have detrimental population-level effects.

Life history trade-offs and community dynamics of small fishes in a seasonally pulsed wetland

We used a one-dimensional, spatially explicit model to simulate the community of small fishes in the freshwater wetlands of southern Florida, USA. The seasonality of rainfall in these wetlands causes annual fluctuations in the amount of flooded area. We modeled fish populations that differed from each other only in efficiency of resource utilization and dispersal ability. The simulations showed that these trade-offs, along with the spatial and temporal variability of the environment, allow coexistence of several species competing exploitatively for a common resource type. This mechanism, while sharing some characteristics with other mechanisms proposed for coexistence of competing species, is novel in detail. Simulated fish densities resembled patterns observed in Everglades empirical data. Cells with hydroperiods less than 6 months accumulated negligible fish biomass. One unique model result was that, when multiple species coexisted, it was possible for one of the coexisting species to have both lower local resource utilization efficiency and lower dispersal ability than one of the other species. This counterintuitive result is a consequence of stronger effects of other competitors on the superior species.

The endocrine basis for reproductive life-history trade-offs during the previtellogenic resting stage in the yellow fever mosquito, Aedes Aegypti

Juvenile hormone (JH) is the central hormonal regulator of life-history trade-offs in many insects. In Aedes aegypti, JH regulates reproductive development after emergence. Little is known about JH's physiological functions after reproductive development is complete or JH's role in mediating life-history trade-offs. By examining the effect of hormones, nutrition, and mating on ovarian physiology during the previtellogenic resting stage, critical roles were determined for these factors in mediating life-history trade-offs and reproductive output. The extent of follicular resorption during the previtellogenic resting stage is dependent on nutritional quality. Feeding females a low quality diet during the resting stage causes the rate of follicular resorption to increase and reproductive output to decrease. Conversely, feeding females a high quality diet causes resorption to remain low. The extent of resorption can be increased by separating the ovaries from a source of JH or decreased by exogenous application of methoprene. Active caspases were localized to resorbing follicles indicating that an apoptosis-like mechanism participates in follicular resorption. Accumulations of neutral lipids and the accumulation of mRNA's integral to endocytosis and oocyte development such as the vitellogenin receptor (AaVgR), lipophorin receptor (AaLpRov), heavy-chain clathrin (AaCHC), and ribosomal protein L32 (rpL32) were also examined under various nutritional and hormonal conditions. The abundance of mRNA's and neutral lipid content increased within the previtellogenic ovary as mosquitoes were offered increasing sucrose concentrations or were treated with methoprene. These same nutritional and hormonal manipulations altered the extent of resorption after a blood meal indicating that the fate of follicles and overall fecundity depends, in part, on nutritional and hormonal status during the previtellogenic resting stage. Mating female mosquitoes also altered follicle quality and resorption similarly to nutrition or hormonal application and demonstrates that male accessory gland substances such as JH III passed to the female during copulation have a strong effect on ovarian physiology during the previtellogenic resting stage and can influence reproductive output. Taken together these results demonstrate that the previtellogenic resting stage is not an inactive period but is instead a period marked by extensive life-history and fitness trade-offs in response to nutrition, hormones and mating stimuli.

The role of oxidative stress as an underlying mechanism in life-history trade-offs

A key aspect underpinning life-history theory is the existence of trade-offs. Trade-offs occur because resources are limited, meaning that individuals cannot invest in all traits simultaneously, leading to costs for traits such as growth and reproduction. Such costs may be the reason for the sub-maximal growth rates that are often observed in nature, though the fitness consequences of these costs would depend on the effects on lifetime reproductive success. Recently, much attention has been given to the physiological mechanism that might underlie these life-history trade-offs, with oxidative stress (OS) playing a key role. OS is characterised by a build-up of oxidative damage to tissues (e.g. protein, lipids and DNA) from attack by reactive species (RS). RS, the majority of which are by-products of metabolism, are usually neutralised by antioxidants, however OS occurs when there is an imbalance between the two. There are two main theories linking OS with growth and reproduction. The first is that traits like growth and reproduction, being metabolically demanding, lead to an increase in RS production. The second involves the diversion of resources away from self-maintenance processes (e.g. the redox system) when individuals are faced with enhanced growth or reproductive expenditure. Previous research investigating trade-offs involving growth or reproduction and self-maintenance has been equivocal. One reason for this could be that associations among redox biomarkers can vary greatly so that the biomarker selected for analysis can influence the conclusion reached about an individual’s oxidative status. Therefore the first aim of my thesis was to explore the strength and pattern of integration of five biomarkers of OS (three antioxidants, one damage and one general oxidation measure) in wild blue tit (Cyanistes caeruleus) adults and nestlings (Chapter 2). In doing so, I established that all five biomarkers should be included in future analyses, thus using this collection of biomarkers I explored my next aims; whether enhanced growth (Chapters 3 and 4) or reproductive effort (Chapter 5) can lead to increased OS levels, if these traits are traded off against self-maintenance. I accomplished these aims using both a meta-analytic and experimental approach, the latter involving manipulation of brood size in wild blue tits in order to experimentally alter growth rate of nestlings and provisioning rate (a proxy for reproductive expenditure) of adults. I also investigated the potential for redox integration to be used as an index of body condition (Chapter 2), allowing predictions about future fitness consequences of changes to oxidative state to be made. A growth – self-maintenance trade off was supported by my meta-analytic results (Chapter 4) which found OS to be a constraint on growth. However, when faced with experimentally enhanced growth, animals were typically not able to adjust this trade-off so that oxidative damage resulted. This might support the idea that energetically expensive growth causes resources to be diverted away from the redox system; however, antioxidants did not show an overall reduction in response to growth in the meta-analysis suggesting that oxidative costs of growth may result from increased RS production due to the greater metabolism needed for enhanced growth. My experimental data (Chapter 3) showed a similar pattern, with raised protein damage levels (protein carbonyls; PCs) in the fastest growing blue tit chicks in a brood, compared with their slower growing sibs. These within-brood differences in OS levels likely resulted from within-brood hierarchies and might have masked any between-brood differences, which were not observed here. Despite evidence for a growth – self-maintenance trade off, my experimental results on blue tits found no support for the hypothesis that self-maintenance is also traded off against reproduction, another energetically demanding trait. There was no link between experimentally altered reproductive expenditure and OS, nor was there a direct correlation between reproductive effort and OS (Chapter 5). However, there are various factors that likely influence whether oxidative costs are observed, including environmental conditions and whether such costs are transient. This emphasises the need for longitudinal studies following the same individuals over multiple years and across a wide range of habitats that differ in quality. This would allow investigation into how key life events interact; it might be that raised OS levels from rapid early growth have the potential to constrain reproduction or that high parental OS levels constrain offspring growth. Any oxidative costs resulting from these life-history trade-offs have the potential to impact on future fitness. Redox integration of certain biomarkers might prove to be a useful tool in making predictions about fitness, as I found in Chapter 2, as well as establishing how the redox system responds, as a whole, to changes to growth and reproduction. Finally, if the tissues measured can tolerate a given level of OS, then the level of oxidative damage might be irrelevant and not impact on future fitness at all.

Identification of trade-offs underlying the primary strategies of plants

Question: What are the key physiological and life-history trade-offs responsible for the evolution of different suites of plant traits (strategies) in different environments? Experimental methods: Common-garden experiments were performed on physiologically realistic model plants, evolved in contrasting environments, in computer simulations. This allowed the identification of the trade-offs that resulted in different suites of traits (strategies). The environments considered were: resource rich, low disturbance (competitive); resource poor, low disturbance (stressed); resource rich, high disturbance (disturbed); and stressed environments containing herbivores (grazed). Results: In disturbed environments, plants increased reproduction at the expense of ability to compete for light and nitrogen. In competitive environments, plants traded off reproductive output and leaf production for vertical growth. In stressed environments, plants traded off vertical growth and reproductive output for nitrogen acquisition, contradicting Grime's (2001) theory that slow-growing, competitively inferior strategies are selected in stressed environments. The contradiction is partly resolved by incorporating herbivores into the stressed environment, which selects for increased investment in defence, at the expense of competitive ability and reproduction. Conclusion: Our explicit modelling of trade-offs produces rigorous testable explanations of observed associations between suites of traits and environments.

Body reserves mediate trade-offs between life-history traits: new insights from small pelagic fish reproduction

Limited resources in the environment prevent individuals from simultaneously maximizing all life-history traits, resulting in trade-offs. In particular, the cost of reproduction is well known to negatively affect energy investment in growth and maintenance. Here, we investigated these trade-offs during contrasting periods of high versus low fish size and body condition (before/after 2008) in the Gulf of Lions. Female reproductive allocation and performance in anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) were examined based on morphometric historical data from the 1970s and from 2003 to 2015. Additionally, potential maternal effects on egg quantity and quality were examined in 2014/2015. After 2008, the gonadosomatic index increased for sardine and remained steady for anchovy, while a strong decline in mean length at first maturity indicated earlier maturation for both species. Regarding maternal effects, for both species egg quantity was positively linked to fish size but not to fish lipid reserves, while the egg quality was positively related to lipid reserves. Atresia prevalence and intensity were rather low regardless of fish condition and size. Finally, estimations of total annual numbers of eggs spawned indicated a sharp decrease for sardine since 2008 but a slight increase for anchovy during the last 5 years. This study revealed a biased allocation towards reproduction in small pelagic fish when confronted with a really low body condition. This highlights that fish can maintain high reproductive investment potentially at the cost of other traits which might explain the present disappearance of old and large individuals in the Gulf of Lions.

Density-dependent morphological plasticity and trade-offs among vegetative traits in Eichhornia crassipes (Pontederiaceae)

Density-dependent responses are an important component of the organism life-history, and the resource allocation theory is a central concept to the life-history theory. When resource allocation varies due to environmental changes, a plant may change its morphology or physiology to cope with the new conditions, a process known as phenotypic plasticity. Our study aimed to evaluate how plant density affects Eichhornia crassipes allocation patterns. A total of 214 individuals in high and low density were collected. The density effect was observed in all plant traits examined including biomass accumulation. All traits of E. crassipes demonstrated higher values in high density conditions, except for biomass of leaves. Density exhibited a high influence on vegetative traits of E. crassipes, but did not influence allocation pattern, since a trade-off among the vegetative traits was not found. The morphological plasticity and the absence of trade-offs were discussed as strategies to overcome neighbor plants in competition situations. In high density conditions, there were clear changes in the morphology of the plants which probably allows for their survival in a highly competitive environment.

Synergies and trade-offs between insect and pathogen resistance in maize leaves and roots

Determining links between plant defence strategies is important to understand plant evolution and to optimize crop breeding strategies. Although several examples of synergies and trade-offs between defence traits are known for plants that are under attack by multiple organisms, few studies have attempted to measure correlations of defensive strategies using specific single attackers. Such links are hard to detect in natural populations because they are inherently confounded by the evolutionary history of different ecotypes. We therefore used a range of 20 maize inbred lines with considerable differences in resistance traits to determine if correlations exist between leaf and root resistance against pathogens and insects. Aboveground resistance against insects was positively correlated with the plant's capacity to produce volatiles in response to insect attack. Resistance to herbivores and resistance to a pathogen, on the other hand, were negatively correlated. Our results also give first insights into the intraspecific variability of root volatiles release in maize and its positive correlation with leaf volatile production. We show that the breeding history of the different genotypes (dent versus flint) has influenced several defensive parameters. Taken together, our study demonstrates the importance of genetically determined synergies and trade-offs for plant resistance against insects and pathogens.

Do synergies improve accuracy? A study of speed-accuracy trade-offs during finger force production

We explored possible effects of negative covariation among finger forces in multifinger accurate force production tasks on the classical Fitts's speed-accuracy trade-off. Healthy subjects performed cyclic force changes between pairs of targets ""as quickly and accurately as possible."" Tasks with two force amplitudes and six ratics of force amplitude to target size were performed by each of the four fingers of the right hand and four finger combinations. There was a close to linear relation between movement time and the log-transformed ratio of target amplitude to target size across all finger combinations. There was a close to linear relation between standard deviation of force amplitude and movement time. There were no differences between the performance of either of the two ""radial"" fingers (index and middle) and the multifinger tasks. The ""ulnar"" fingers (little and ring) showed higher indices of variability and longer movement times as compared with both ""radial"" fingers and multifinger combinations. We conclude that potential effects of the negative covariation and also of the task-sharing across a set of fingers are counterbalanced by an increase in individual finger force variability in multifinger tasks as compared with single-finger tasks. The results speak in favor of a feed-forward model of multifinger synergies. They corroborate a hypothesis that multifinger synergies are created not to improve overall accuracy, but to allow the system larger flexibility, for example to deal with unexpected perturbations and concomitant tasks.

Life-history consequences of divergent selection on egg size in Drosophila melanogaster

Life histories are generally assumed to evolve via antagonistic pleiotropy (negative genetic correlations) among traits, and trade-offs between life-history traits are typically studied using either phenotypic manipulations or selection experiments. We investigated the trade-off between egg size and fecundity in Drosophila melanogaster by examining both the phenotypic and genetic relationships between these traits after artificial selection for large and small eggs, relative to female body size. Egg size responded strongly to selection in both directions, increasing in the large-egg selected lines and decreasing in the small-egg selected lines. Phenotypic correlations between egg size and fecundity in the large-egg selected lines were negative, but no relationship between these traits occurred in either the control or small-egg selected lines. There was no negative genetic correlation between egg size and fecundity. Total reproductive allocation decreased in the small-egg selected lines but did not increase in the large-egg lines. Our results have three implications. First, our selection procedure may have forced females selected for large eggs into a physiological trade-off not reflected in a negative genetic correlation between these traits. Second, the lack of a negative genetic correlation between egg size and number suggests that the phenotypic trade-off frequently observed between egg size and number in other organisms may not evolve over the short term via a direct genetic trade-off whereby increases in egg size are automatically accompanied by decreased fecundity. Finally, total reproductive allocation may not evolve independently of egg size as commonly assumed.

All-Optical Label Stacking: Easing the Trade-offs Between Routing and Architecture Cost in All-Optical Packet Switching

All-optical label swapping (AOLS) forms a key technology towards the implementation of all-optical packet switching nodes (AOPS) for the future optical Internet. The capital expenditures of the deployment of AOLS increases with the size of the label spaces (i.e. the number of used labels), since a special optical device is needed for each recognized label on every node. Label space sizes are affected by the way in which demands are routed. For instance, while shortest-path routing leads to the usage of fewer labels but high link utilization, minimum interference routing leads to the opposite. This paper studies all-optical label stacking (AOLStack), which is an extension of the AOLS architecture. AOLStack aims at reducing label spaces while easing the compromise with link utilization. In this paper, an integer lineal program is proposed with the objective of analyzing the softening of the aforementioned trade-off due to AOLStack. Furthermore, a heuristic aiming at finding good solutions in polynomial-time is proposed as well. Simulation results show that AOLStack either a) reduces the label spaces with a low increase in the link utilization or, similarly, b) uses better the residual bandwidth to decrease the number of labels even more

Direct and indirect root defences of milkweed (Asclepias syriaca): trophic cascades, trade-offs and novel methods for studying subterranean herbivory

P>1. Entomopathogenic nematodes can function as indirect defence for plants that are attacked by root herbivores. By releasing volatile organic compounds (VOCs), plants signal the presence of host insects and thereby attract nematodes.2. Nonetheless, how roots deploy indirect defences, how indirect defences relate to direct defences, and the ecological consequences of root defence allocation for herbivores and plant biomass are essentially unknown.3. We investigate a natural below-ground tritrophic system, involving common milkweed, a specialist root-boring beetle and entomopathogenic nematodes, and asked whether there is a negative genetic correlation between direct defences (root cardenolides) and indirect defences (emission of volatiles in the roots and nematode attraction), and between constitutive and inducible defences.4. Volatiles of roots were analysed using two distinct sampling methods. First, we collected emissions from living Asclepias syriaca roots by dynamic headspace sampling. This method showed that attacked A. syriaca plants emit five times higher levels of volatiles than control plants. Secondly, we used a solid phase micro-extraction (SPME) method to sample the full pool of volatiles in roots for genetic correlations of volatile biosynthesis.5. Field experiments showed that entomopathogenic nematodes prevent the loss of biomass to root herbivory. Additionally, suppression of root herbivores was mediated directly by cardenolides and indirectly by the attraction of nematodes. Genetic families of plants with high cardenolides benefited less from nematodes compared to low-cardenolide families, suggesting that direct and indirect defences may be redundant. Although constitutive and induced root defences traded off within each strategy (for both direct and indirect defence, cardenolides and VOCs, respectively), we found no trade-off between the two strategies.6. Synthesis. Constitutive expression and inducibility of defences may trade off because of resource limitation or because they are redundant. Direct and indirect defences do not trade off, likely because they may not share a limiting resource and because independently they may promote defence across the patchiness of herbivore attack and nematode presence in the field. Indeed, some redundancy in strategies may be necessary to increase effective defence, but for each strategy, an economy of deployment reduces overall costs.

Speed-cohesion trade-offs in collective decision making in ants and the concept of precision in animal behaviour

In decision making, speed-accuracy trade-offs are well known and often inevitable because accuracy depends on being well informed and gathering information takes time. However, trade-offs between speed and cohesion, that is the degree to which a group remains together as a single entity, as a result of their decision making, have been comparatively neglected. We combine theory and experimentation to show that in decision-making systems, speed-cohesion trade-offs are a natural complement to speed-accuracy trade-offs and are therefore of general importance. We then analyse the decision performance of 32 rock ant, Temnothorax albipennis, colonies in experiments in which accuracy of collective decision making was held constant, but time urgency varied. These experiments reveal for the first time an adaptive speed-cohesion trade-off in collective decision making and how this is achieved. In accord with different time constraints, colonies can decide quickly, at the cost of social unity, or they can decide slowly with much greater cohesion. We discuss the similarity between cohesion and the term precision as used in statistics and engineering. This emphasizes the generality of speed versus cohesion/precision trade-offs in decision making and decision implementation in other fields within animal behaviour such as sexually selected motor displays and even certain aspects of birdsong. We also suggest that speed versus precision trade-offs may occur when individuals within a group need to synchronize their activity, and in collective navigation, cooperative hunting and in certain escape behaviours.

All-Optical Label Stacking: Easing the Trade-offs Between Routing and Architecture Cost in All-Optical Packet Switching

All-optical label swapping (AOLS) forms a key technology towards the implementation of all-optical packet switching nodes (AOPS) for the future optical Internet. The capital expenditures of the deployment of AOLS increases with the size of the label spaces (i.e. the number of used labels), since a special optical device is needed for each recognized label on every node. Label space sizes are affected by the way in which demands are routed. For instance, while shortest-path routing leads to the usage of fewer labels but high link utilization, minimum interference routing leads to the opposite. This paper studies all-optical label stacking (AOLStack), which is an extension of the AOLS architecture. AOLStack aims at reducing label spaces while easing the compromise with link utilization. In this paper, an integer lineal program is proposed with the objective of analyzing the softening of the aforementioned trade-off due to AOLStack. Furthermore, a heuristic aiming at finding good solutions in polynomial-time is proposed as well. Simulation results show that AOLStack either a) reduces the label spaces with a low increase in the link utilization or, similarly, b) uses better the residual bandwidth to decrease the number of labels even more