Portfolio theory and how parent birds manage investment risk

Investment theory is founded on the premise that higher returns are generally associated with greater risk, and that portfolio diversification reduces risk. Here I examine parental investment decisions in birds from this perspective, using data from a model system, a 16‐year study of breeding red‐winged blackbirds Agelaius phoeniceus. Like many altricial birds, blackbirds structure their brood into core (first‐hatched) and marginal (later‐hatched) elements that differ in risk profile. I measured risk in two ways: as the coefficient of variation in growth and survival of core and marginal offspring from a given brood structure; and using financial beta derived from the capital asset pricing model of modern portfolio theory. Financial beta correlates changes in asset value with changes in the value of a broader market, defined here as individual reproductive success vs. population reproductive success. Both measures of risk increased with larger core (but not marginal) brood size; and variation in growth and survival was significantly greater during ecologically adverse conditions. Core offspring showed low beta values relative to marginal progeny. The most common brood structures in the population exhibited the highest beta values for both core and marginal offspring: many parent blackbirds embraced rather than avoided risk. But they did so prudently with an investment strategy that resembled a financial instrument, the call option. A call option is a contingent claim on the future value of the asset, and is exercised only if asset value increases beyond a point fixed in advance. Otherwise the option lapses and the investor loses only the initial option price. Parents created high risk marginal progeny that were forfeited during ecological adversity (the option lapses) but raised otherwise (the option called); at the same time parents maintained a constant investment and return in low risk core progeny that varied little with changes in brood size or ecological conditions.     

Darwinian fitness

The term Darwinian fitness refers to the capacity of a variant type to invade and displace the resident population in competition for available resources. Classical models of this dynamical process claim that competitive outcome is a deterministic event which is regulated by the population growth rate, called the Malthusian parameter. Recent analytic studies of the dynamics of competition in terms of diffusion processes show that growth rate predicts invasion success only in populations of infinite size. In populations of finite size, competitive outcome is a stochastic process--contingent on resource constraints--which is determined by the rate at which a population returns to its steady state condition after a random perturbation in the individual birth and death rates. This return rate, a measure of robustness or population stability, is analytically characterized by the demographic parameter, evolutionary entropy, a measure of the uncertainty in the age of the mother of a randomly chosen newborn. This article appeals to computational and numerical methods to contrast the predictive power of the Malthusian and the entropic principles. The computational analysis rejects the Malthusian model and is consistent with of the entropic principle. These studies thus provide support for the general claim that entropy is the appropriate measure of Darwinian fitness and constitutes an evolutionary parameter with broad predictive and explanatory powers.     

Mortality plateaus and directionality theory

Recent large scale studies of senescence in animals and humans have revealed mortality rates that levelled off at advanced ages. These empirical findings are now known to be inconsistent with evolutionary theories of senescence based on the Malthusian parameter as a measure of fitness. This article analyses the incidence of mortality plateaus in terms of directionality theory, a new class of models based on evolutionary entropy as a measure of fitness. We show that the intensity of selection, in the context of directionality theory, is a convex function of age, and we invoke this property to predict that in populations evolving under bounded growth constraints, evolutionarily stable mortality patterns will be described by rates which abate with age at extreme ages. The explanatory power of directionality theory, in contrast with the limitations of the Malthusian model, accords with the claim that evolutionary entropy, rather than the Malthusian parameter, constitutes the operationally valid measure of Darwinian fitness.     

Financial Crisis: A New Measure for Risk of Pension Fund Portfolios

It has been argued that pension funds should have limitations on their asset allocation, based on the risk profile of the different financial instruments available on the financial markets. This issue proves to be highly relevant at times of market crisis, when a regulation establishing limits to risk taking for pension funds could prevent defaults. In this paper we present a framework for evaluating the risk level of a single financial instrument or a portfolio. By assuming that the log asset returns can be described by a multifractional Brownian motion, we evaluate the risk using the time dependent Hurst parameter H(t) which models volatility. To provide a measure of the risk, we model the Hurst parameter with a random variable with mixture of beta distribution. We prove the efficacy of the methodology by implementing it on different risk level financial instruments and portfolios.     

Inferring the Relative Resilience of Alternative States

Ecological systems may occur in alternative states that differ in ecological structures, functions and processes. Resilience is the measure of disturbance an ecological system can absorb before changing states. However, how the intrinsic structures and processes of systems that characterize their states affects their resilience remains unclear. We analyzed time series of phytoplankton communities at three sites in a floodplain in central Spain to assess the dominant frequencies or “temporal scales” in community dynamics and compared the patterns between a wet and a dry alternative state. The identified frequencies and cross-scale structures are expected to arise from positive feedbacks that are thought to reinforce processes in alternative states of ecological systems and regulate emergent phenomena such as resilience. Our analyses show a higher species richness and diversity but lower evenness in the dry state. Time series modeling revealed a decrease in the importance of short-term variability in the communities, suggesting that community dynamics slowed down in the dry relative to the wet state. The number of temporal scales at which community dynamics manifested, and the explanatory power of time series models, was lower in the dry state. The higher diversity, reduced number of temporal scales and the lower explanatory power of time series models suggest that species dynamics tended to be more stochastic in the dry state. From a resilience perspective our results highlight a paradox: increasing species richness may not necessarily enhance resilience. The loss of cross-scale structure (i.e. the lower number of temporal scales) in community dynamics across sites suggests that resilience erodes during drought. Phytoplankton communities in the dry state are therefore likely less resilient than in the wet state. Our case study demonstrates the potential of time series modeling to assess attributes that mediate resilience. The approach is useful for assessing resilience of alternative states across ecological and other complex systems.     

Relevance of evolutionary history for food web structure

Explaining the structure of ecosystems is one of the great challenges of ecology. Simple models for food web structure aim at disentangling the complexity of ecological interaction networks and detect the main forces that are responsible for their shape. Trophic interactions are influenced by species traits, which in turn are largely determined by evolutionary history. Closely related species are more likely to share similar traits, such as body size, feeding mode and habitat preference than distant ones. Here, we present a theoretical framework for analysing whether evolutionary history--represented by taxonomic classification--provides valuable information on food web structure. In doing so, we measure which taxonomic ranks better explain species interactions. Our analysis is based on partitioning of the species into taxonomic units. For each partition, we compute the likelihood that a probabilistic model for food web structure reproduces the data using this information. We find that taxonomic partitions produce significantly higher likelihoods than expected at random. Marginal likelihoods (Bayes factors) are used to perform model selection among taxonomic ranks. We show that food webs are best explained by the coarser taxonomic ranks (kingdom to class). Our methods provide a way to explicitly include evolutionary history in models for food web structure.     

Low HRV entropy is strongly associated with myocardial infarction.

Heart rate variability (HRV) is a marker of autonomous activity in the heart. An important application of HRV measures is the stratification of mortality risk after myocardial infarction. Our hypothesis is that the information entropy of HRV, a non-linear approach, is a suitable measure for this assessment. As a first step, to evaluate the effect of myocardial infarction on the entropy, we compared the entropy to standard HRV parameters. The entropy was estimated by compressing the tachogram with Bzip2. For univariate comparison, statistical tests were used. Multivariate analysis was carried out using automatically generated decision trees. The classification rate and the simplicity of the decision trees were the two evaluation criteria. The findings support our hypothesis. The meanNN-normalized entropy is reduced in patients with myocardial infarction with very high significance. One entropy parameter alone exceeds the discrimination strength of multivariate standards-based trees.     

Isolation by oceanographic distance explains genetic structure for Macrocystis pyrifera in the Santa Barbara Channel

Ocean currents are expected to be the predominant environmental factor influencing the dispersal of planktonic larvae or spores; yet, their characterization as predictors of marine connectivity has been hindered by a lack of understanding of how best to use oceanographic data. We used a high-resolution oceanographic model output and Lagrangian particle simulations to derive oceanographic distances (hereafter called transport times) between sites studied for Macrocystis pyrifera genetic differentiation. We build upon the classical isolation-by-distance regression model by asking how much additional variability in genetic differentiation is explained when adding transport time as predictor. We explored the extent to which gene flow is dependent upon seasonal changes in ocean circulation. Because oceanographic transport between two sites is inherently asymmetric, we also compare the explanatory power of models using the minimum or the mean transport times. Finally, we compare the direction of connectivity as estimated by the oceanographic model and genetic assignment tests. We show that the minimum transport time had higher explanatory power than the mean transport time, revealing the importance of considering asymmetry in ocean currents when modelling gene flow. Genetic assignment tests were much less effective in determining asymmetry in gene flow. Summer-derived transport times, in particular for the month of June, which had the strongest current speed, greatest asymmetry and highest spore production, resulted in the best-fit model explaining twice the variability in genetic differentiation relative to models that use geographic distance or habitat continuity. The best overall model also included habitat continuity and explained 65% of the variation in genetic differentiation among sites.     

Folding kinetics of two-state proteins: effect of circularization,permutation, and crosslinks

Protein folding kinetics has recently been probed by clever experiments using circular permutants and other topological mutations. A circular permutant is created from a wild-type protein by covalently linking together the chain ends and cleaving elsewhere in the chain. An interesting puzzle is why circular permutation causes no apparent change in the folding mechanism of CI2, but dramatic changes in the folding mechanisms of S6 and of an SH3 domain, as determined by Phi-value experiments. Here, we use a computational model to predict the folding routes of topological variants, based on a measure (effective contact order) of the chain entropy loss at each folding step. The predictions are consistent with the experiments, leading to insights into the folding routes and into the meaning of Phi-values in general. We find that Phi-values do not always describe time sequences of folding events, or positions along a single reaction coordinate; rather, Phi reflects only the degree of rate control. For example, the circular permutant P(40-41) of CI2 is predicted to reverse the time sequence of the formation of beta(1)beta(4) relative to beta(2)beta(3), without changing the diffuse Phi-value distribution, while the circular permutant P(13-14) of S6 switches the rate-limiting step from the formation of beta(1)beta(4) to beta(1)beta(3), changing the Phi-value distribution from diffuse to strongly polarized. As a test of the model, we propose mutations that should reverse these outcomes.     

Computing parametric beta diversity with unequal plot weights: a solution based on resampling methods

Jost (Ecology, 88:2427–2439, 2007) recently showed that the Shannon diversity is the only standard diversity measure that can be partitioned into meaningful independent alpha and beta components when plot weights are unequal. This conclusion is very disappointing if one wants to calculate the beta diversity of unequal weighted plots using a parametric measure with varying sensitivities to the occurrence of rare and abundant species. To overcome this impasse, at least partially, in this paper, I propose a parametric measure of beta diversity that is based on the combination of Shannon’s entropy with Hurlbert’s ‘expected species diversity’. Unlike most parametric measures of diversity, the proposed index has a clear probabilistic interpretation, allowing at the same time a multiplicative partition of diversity into independent alpha and beta components for unequally weighted plots.     

林冠结构是局域尺度木本植物功能性状beta多样性形成的重要驱动力

功能性状beta多样性反映了群落间功能性状组成的差异, 解析其形成机制是群落生态学研究的核心内容之一。本研究以云南西双版纳热带季节雨林20 ha动态监测样地为研究对象, 测定木本植物11个重要的功能性状, 采用多度加权的平均最近邻体性状距离度量不同取样尺度的功能性状beta多样性, 基于距离矩阵的多元回归方法解析林冠结构差异、环境异质性、空间距离在功能性状beta多样性格局形成中的相对作用。结果表明, 对于所有木本植物个体(DBH ≥ 1 cm)而言, 同时考虑林冠结构、环境和空间距离的模型为解释功能性状beta多样性格局的最优模型; 在3个不同取样尺度上, 林冠结构差异和环境距离都对功能性状beta多样性具有较大的解释力, 且随着取样尺度的增大而上升, 空间距离的作用基本可以忽略。本研究证实了林冠结构是局域尺度木本植物功能性状beta多样性格局形成的重要驱动力, 这一发现更新了环境异质性和空间距离是驱动功能性状beta多样性格局形成的主要因素的传统认知, 为将来研究功能性状beta多样性形成机制提供新的视角, 并证实了取样尺度在解析木本植物功能性状beta多样性格局形成机制中的重要性。     

Parental care tactics of three-spined sticklebacks living in a harsh environment

The three-spined stickleback (Gasterosteus aculeatus) is characterizedby paternal care of fertilized eggs. We carried out field experimentsto evaluate the response of males to simulated risks to theirbroods and themselves in relation to brood size and age, timeof breeding season, age of parent, nest-site characteristics(depth, distance from shore, distance to nearest nest, degreeof nest cover), and water temperature. For males with eggs intheir nests, we chased the male away from the nest to simulatethe arrival of a bird predator and measured the time until return,a measure of risk taking. There was no relationship betweenreturn time and the number or the age of eggs. There was a smallbut significant relationship between the distance of the nestfrom the shore and return time. Males with nests farther fromthe shore tended to return sooner. No other environmental variableexplained a significant proportion of the variation in returntime. Return time was unrelated to either time of breeding seasonor male age. In another experiment, a model of a rival maleconspecific was placed near the guardian's nest, simulatinga potential threat to the eggs. Again, we found no relationshipbetween aggression to the model and egg number or egg age. Maleswith nests in deeper water were less aggressive than those withnests in shallower water, but no other environmental variableexplained a significant proportion of the variation in aggression.Male age was unrelated to the amount of aggression, but malesbecame more aggressive as the season progressed. Our resultscontrast with previous studies of parental behavior in thisspecies. Possible reasons for these differences are discussed.     

Are there latitudinal gradients in species turnover?

Aim To examine the effect on the observed relationship between spatial turnover and latitude of both the measure of beta diversity used and the method of analysis. Location The empirical analyses presented herein are for the New World. Methods We take the spatial distributions of the owls of the New World as an exemplar data set to investigate the patterns of beta diversity across latitudes revealed by different analytical methods. To illustrate the strengths and weaknesses of alternative measures of beta diversity and different analytical approaches, we also use a simple random distribution model, focusing in particular on the influence of richness gradients and landmass geometry. Results Our simple spatial model of turnover demonstrates that different combinations of analytical approach and measure of beta diversity can give rise to strikingly different relationships between turnover and latitude. The analyses of the bird data for the owls of the New World demonstrate that this observation extends to real data. Conclusions For the particular assemblage considered, we present strong evidence that species richness declines at higher latitudes, and there is also some evidence that species turnover is greater nearer the equator, despite conceptual and practical difficulties involved in analysing spatial patterns of species turnover. We suggest some ways of overcoming these difficulties.     

Quantifying Predation Risk for Refuging Animals: A Case Study with Golden Marmots

Although a variety of behaviors expose animals to some risk of predation, there is no accepted way to compare their relative risk. For animals that retreat to refugia when alarmed by predators, the proportion of time devoted to each out-of-refuge behavior multiplied by the total time required to return to a refuge can be used to compare a behavior's relative predation risk. Total time to return to a refuge is a function of both response time - the time required to respond to an increased risk of predation — and travel time — the time required to flee to a refuge once alarmed. Quantifying these components can illustrate how animals minimize exposure to predators. Golden marmots (Marmota caudata aurea) were a refuging prey species used to examine the utility of this measure and to understand how marmots minimized their risk of exposure to predation. Golden marmots devoted different amounts of time to looking, foraging, self-grooming, and playing. To estimate the behavior-specific time required to return to refugia, the location of different activities was noted and a behavior-specific travel time was calculated. Alarm calls were played back to marmots engaged in different behaviors to determine, in a standardized manner, if there were behavior-specific response times. Marmots appeared to minimize their predation risk by performing most behaviors close to refugia. Results suggest that foraging was the riskiest behavior, largely because marmots foraged far from refugia and spent about 30% of their time foraging. While sample sizes were small, results also suggested that play, a rare adult behavior, exposed animals to predation because of a relatively long response time.     

Phylogenetic,spatial and environmental components of extinction risk in carnivores

Aim Extinction risk is non‐randomly distributed across phylogeny and space and is influenced by environmental conditions. We quantified the relative contribution of these factors to extinction risk to unveil the underlying macroecological processes and derive predictive models. Location Global. Methods Based on the IUCN global assessments, we divided 192 carnivore species into two dichotomous classes representing different levels of extinction risk. We used spatial proximity, phylogenetic relationship and environmental variables together with phylogenetic eigenvector regression and spatial eigenvector filters to model and predict threat status. Results Our full models explained between 57% and 96% of the variance in extinction risk. Phylogeny and spatial proximity roughly explained between 21% and 70% of the total variation in all analyses, while the explanatory power of environmental conditions was relatively weaker (up to 15%). Phylogeny and spatial proximity contributed equally to the explained variance in the lower threat level, while spatial proximity was the most important factor in the models of the higher threat level. Prediction of threat status achieved 97% correct assignments. Main conclusions Our approach differs fundamentally from current studies of extinction risk because it does not necessarily rely on life‐history information. We clearly show that instead of treating phylogenetic inertia and spatial signal as statistical nuisances, space and phylogeny should be viewed as very useful in explaining a wide range of phenomena in comparative studies.     

Trade-offs between storage and survival affect diapause timing in capital breeders

Many organisms spend the unfavourable part of the year, such as the winter season, in diapause or dormancy and reproduce in spring shortly after emergence. Reserves are acquired prior to diapause to cover metabolic costs and in some species also reproduction (capital breeding) directly after diapause. Storage is then a component of future reproduction, and capital breeders consequently pay a pre-breeding cost of reproduction as they risk dying while obtaining and carrying the reserves. How large should the reserves be, and to what extent should optimal storage, and thereby timing of diapause, depend on predation risk and reproductive strategy? We present a general and simplistic life history model of an arthropod (e.g. crustaceans or insects) that is exposed to background mortality risk when it accumulates reserves before diapause. The model optimizes diapause timing and resultant reserves for income, mixed and capital breeders, and predicts how mortality risk affects the degree of capital breeding. For income breeders, timing of diapause is insensitive to the risk while obtaining reserves as they, regardless of risk, acquire the minimum amount needed to survive the winter. For capital breeders, the higher the risk the earlier the diapause and less is consequently stored. Mixed breeders diapause late and store as much as pure capital breeders when exposed to low risk, but behave as income breeders and diapause early when mortality is high. Our model shows that the degree of capital breeding impacts phenology of diapause in a risk-dependent manner. This prediction should impact how diapause timing is thought of across a wide range of taxa, including the much studied marine copepods. Timing of diapause, including triggers and cues, can only be understood when the diversity of reproductive strategies and the adaptive value of storage is taken into account.     

Soil pH determines fungal diversity along an elevation gradient in Southwestern China

Fungi play important roles in ecosystem processes, and the elevational pattern of fungal diversity is still unclear. Here, we examined the diversity of fungi along a 1,000 m elevation gradient on Mount Nadu, Southwestern China. We used MiSeq sequencing to obtain fungal sequences that were clustered into operational taxonomic units(OTUs) and to measure the fungal composition and diversity. Though the species richness and phylogenetic diversity of the fungal community did not exhibit significant trends with increasing altitude, they were significantly lower at mid-altitudinal sites than at the base. The Bray-Curtis distance clustering also showed that the fungal communities varied significantly with altitude. A distance-based linear model multivariate analysis(DistLM) identified that soil pH dominated the explanatory power of the species richness(23.72%),phylogenetic diversity(24.25%) and beta diversity(28.10%) of the fungal community. Moreover, the species richness and phylogenetic diversity of the fungal community increased linearly with increasing soil pH(P0.05). Our study provides evidence that pH is an important predictor of soil fungal diversity along elevation gradients in Southwestern China.     

Landscape, flux, correlation, resonance, coherence, stability, and key network wirings of stochastic circadian oscillation

Circadian rhythms with a period of ∼24 h, are natural timing machines. They are broadly distributed in living organisms, such as Neurospora, Drosophila, and mammals. The underlying natures of the rhythmic behavior have been explored by experimental and theoretical approaches. However, the global and physical natures of the oscillation under fluctuations are still not very clear. We developed a landscape and flux framework to explore the global stability and robustness of a circadian oscillation system. The potential landscape of the network is uncovered and has a global Mexican-hat shape. The height of the Mexican-hat provides a quantitative measure to evaluate the robustness and coherence of the oscillation. We found that in nonequilibrium dynamic systems, not only the potential landscape but also the probability flux are important to the dynamics of the system under intrinsic noise. Landscape attracts the systems down to the oscillation ring while flux drives the coherent oscillation on the ring. We also investigated the phase coherence and the entropy production rate of the system at different fluctuations and found that dissipations are less and the coherence is higher for larger number of molecules. We also found that the power spectrum of autocorrelation functions show resonance peak at the frequency of coherent oscillations. The peak is less prominent for smaller number of molecules and less barrier height and therefore can be used as another measure of stability of oscillations. As a consequence of nonzero probability flux, we show that the three-point correlations from the time traces show irreversibility, providing a possible way to explore the flux from the observations. Furthermore, we explored the escape time from the oscillation ring to outside at different molecular number. We found that when barrier height is higher, escape time is longer and phase coherence of oscillation is higher. Finally, we performed the global sensitivity analysis of the underlying parameters to find the key network wirings responsible for the stability of the oscillation system.     

我国农业生态效率的时空差异

农业生态效率是按照定量化的方式反映区域农业发展可持续发展水平,可以作为决策者制定政策的一个抓手。利用基于机会成本的经济核算方法对我国2003—2010年的农业生态效率进行总体分析与评价,并利用回归模型分析农业生态效率的影响因素。结果表明:我国农业生态效率总体水平比较低,但呈逐年好转的趋势,其中劳动力资源和COD环境要素在不同时期对生态价值增长起到关键性作用;农业生态效率空间分布特征显著,秦岭-淮河以北的省市区和传统粮食主产区的农业生态效率相对较低;区域资源环境禀赋条件有助于农业生态效率的提高,但是农资投入和农业政策支持与农业生态效率呈显著负相关,未来进一步提升农业生产资源与环境要素合理配置是保障农业生产可持续的必然选择。