The phenomenology of niche evolution via quantitative traits in a 'black-hole' sink

Previous studies of adaptive evolution in sink habitats (in which isolated populations of a species cannot persist deterministically) have highlighted the importance of demographic constraints in slowing such evolution, and of immigration in facilitating adaptation. These studies have relied upon either single-locus models or deterministic quantitative genetic formulations. We use individual-based simulations to examine adaptive evolution in a 'black-hole' sink environment where fitness is governed by a polygenic character. The simulations track both the number of individuals and their multi-locus genotypes, and incorporate, in a natural manner, both demographic and genetic stochastic processes. In agreement with previous studies, our findings reveal the central parts played by demographic constraints and immigration in adaptation within a sink (adaptation is more difficult in environments with low absolute fitness, and higher immigration can accelerate adaptation). A novel finding is that there is a 'punctuational' pattern in adaptive evolution in sink environments. Populations typically stay maladapted for a long time, and then rapidly shift into a relatively adapted state, in which persistence no longer depends upon recurrent immigration.     

Phenotypic plasticity can facilitate adaptive evolution in gene regulatory circuits

Background  

Many important evolutionary adaptations originate in the modification of gene regulatory circuits to produce new gene activity phenotypes. How do evolving populations sift through an astronomical number of circuits to find circuits with new adaptive phenotypes? The answer may often involve phenotypic plasticity. Phenotypic plasticity allows a genotype to produce different - alternative - phenotypes after non-genetic perturbations that include gene expression noise, environmental change, or epigenetic modification.     

Cellulolytic bacteria from soils in harsh environments

It is believed that the exposure of organisms to harsh climate conditions may select for differential enzymatic activities, making the surviving organisms a very promising source for bioprospecting. Soil bacteria play an important role in degradation of organic matter, which is mostly due to their ability to decompose cellulose-based materials. This work focuses on the isolation and identification of cellulolytic bacteria from soil found in two environments with stressful climate conditions (Antarctica and the Brazilian semi-arid caatinga). Cellulolytic bacteria were selected using enrichments at high and low temperatures (4 or 60°C) in liquid media (trypic soy broth—TSB and minimum salt medium—MM) supplemented with cellulose (1%). Many of the isolates (119 out of 254—46.9%) displayed the ability to degrade carboxymethyl-cellulose, indicating the presence of endoglucolytic activity, while only a minority of these isolates (23 out of 254—9.1%) showed exoglucolytic activity (degradation of avicel). The obtained isolates revealed a preferential endoglucolytic activity according to the temperature of enrichments. Also, the identification of some isolates by partial sequencing of the 16S rRNA gene indicated that the Bacteroidetes (e.g., Pedobacter, Chryseobacterium and Flavobacterium) were the main phylum of cellulolytic bacteria isolated from soil in Antarctica; the Firmicutes (e.g., Bacillus) were more commonly isolated from samples from the caatinga; and Actinobacteria were found in both types of soil (e.g., Microbacterium and Arthrobacter). In conclusion, this work reports the isolation of bacteria able to degrade cellulose-based material from soil at very low or very high temperatures, a finding that should be further explored in the search for cellulolytic enzymes to be used in the bioenergy industry.     

Behavioural environments and niche construction: the evolution of dim-light foraging in bees

Most bees forage for floral resources during the day, but temporal patterns of foraging activity vary extensively, and foraging in dim-light environments has evolved repeatedly. Facultative dim-light foraging behaviour is known in five of nine families of bees, while obligate behaviour is known in four families and evolved independently at least 19 times. The light intensity under which bees forage varies by a factor of 10⁸, and therefore the evolution of dim-light foraging represents the invasion of a new, extreme niche. The repeated evolution of dim-light foraging behaviour in bees allows tests of the hypothesis that behaviour acts as an evolutionary pacemaker. With the exception of one species of Apis , facultative dim-light foragers show no external structural traits that are thought to enable visually mediated flight behaviour in low-light environments. By contrast, most obligate dim-light foragers show a suite of convergent optical traits such as enlarged ocelli and compound eyes. In one intensively studied species ( Megalopta genalis ) these optical changes are associated with neurobiological changes to enhance photon capture. The available ecological evidence suggests that an escape from competition for pollen and nectar resources and avoidance of natural enemies are driving factors in the evolution of obligate dim-light foraging.     

Quantifying energy dissipation by grazing animals in harsh environments

Grazing systems in harsh environments are common throughout the world, and animal production is the mainstay of the livelihoods of many resource-poor farmers. The energy cost of the various activities involved in the process of harvesting the pasture to transform it into animal product can be estimated through an energy balance. This cost would be the difference between the metabolizable energy intake (MEI) and the energy expenditures for maintenance (MEm), temperature regulation (MEtr), and the energy for production (MEp). Each of the ME has its own net energy (NE) and its associated efficiency (K). When MEI>MEm+MEtr+MEp, the difference is attributable to the energy dissipated during grazing. The efficiency of converting the energy consumed into animal products depends on the magnitude of the dissipation. The inefficiency is associated with the energy spent in locomotion and the stress produced when there is low availability of energy in the pasture. This paper presents a method to quantify the dissipation of energy by grazing animals by considering it as a function of available energy. Such an understanding is required in order to develop management strategies to increase conversion efficiency.     

Temporal evolution of the European forest sector carbon sink from 1950 to 1999

Estimates of the role of the European terrestrial biosphere in the global carbon cycle still vary by a factor 10. This is due to differences in methods and assumptions employed, but also due to difference in reference periods of the studies. The magnitude of the sink varies between years because of inter‐annual variation of short‐term climate, but also due to long‐term trends in development of the vegetation and its management. For this purpose, we present the results of an application of a carbon bookkeeping model to the forest sector of the European forests from 1950 to 1999. The analysis includes the compartments trees, soils, and wood products. The model uses statistics on European (30 countries excl. CIS) stemwood volume increment, forest area change, fellings, wood products and their international trade, and natural disturbances, supplemented with conversion coefficients, soil parameters and information on management. An (almost uninterrupted) increasing sink (Net Biome Production) in the European forest sector was found, increasing from 0.03 Pg C year ^?1 in the 1950s to 0.14 Pg C year ^?1 in the 1990s (for resp. 132 million hectares and 140 million hectares of forest). The sink in the tree and the soil compartment were approximately of the same size until 1970. After the 1970s the size of the sink in the tree biomass increases quickly, causing the tree biomass to account for some two thirds of the total sink in the 1990s. The results as presented here have to be regarded with caution especially with regard to the early decades of the analysis and with regard to the soil compartment.     

Facilitated variation: how evolution learns from past environments to generalize to new environments

One of the striking features of evolution is the appearance of novel structures in organisms. Recently, Kirschner and Gerhart have integrated discoveries in evolution, genetics, and developmental biology to form a theory of facilitated variation (FV). The key observation is that organisms are designed such that random genetic changes are channeled in phenotypic directions that are potentially useful. An open question is how FV spontaneously emerges during evolution. Here, we address this by means of computer simulations of two well-studied model systems, logic circuits and RNA secondary structure. We find that evolution of FV is enhanced in environments that change from time to time in a systematic way: the varying environments are made of the same set of subgoals but in different combinations. We find that organisms that evolve under such varying goals not only remember their history but also generalize to future environments, exhibiting high adaptability to novel goals. Rapid adaptation is seen to goals composed of the same subgoals in novel combinations, and to goals where one of the subgoals was never seen in the history of the organism. The mechanisms for such enhanced generation of novelty (generalization) are analyzed, as is the way that organisms store information in their genomes about their past environments. Elements of facilitated variation theory, such as weak regulatory linkage, modularity, and reduced pleiotropy of mutations, evolve spontaneously under these conditions. Thus, environments that change in a systematic, modular fashion seem to promote facilitated variation and allow evolution to generalize to novel conditions.     

Clonal splitters and integrators in harsh environments of the Trans-Himalaya

Individuals of clonal plants consist of physically and physiologically connected ramets. In splitters, they are integrated for a time shorter than ramet generation time (i.e. the time it takes to produce the first offspring ramet), whereas in integrators connections between ramets persist for a longer time. It has been predicted that integrators should prevail in stressful environments, such as habitats poor in nutrients, whereas splitters are expected to dominate in benign habitats, such as fertile areas with a moderate climate. I tested these predictions in four dry mountain areas of the Trans-Himalaya, in high altitudes subjected to multiple stresses. In accordance with the expectations I found that clonal plants with integrated ramets reach higher mean and maximum altitudes than splitters. Integrators were over-represented in nutrient-poor habitats, such as dry semi-deserts, sandy steppes and in subnival habitats, whereas splitters preferentially colonised mesic habitats, saline sites and wetlands. While there was no difference in the representation of splitters and integrators in habitats with an unstable surface, such as screes, dunes and water bodies, fully integrated clonal plants preferred very stable environments, such as banks of streams covered by closed-canopy vegetation. Most relationships between clonal integration and environmental factors were explainable by the phylogenetic relationship between the species, only the significant preference of splitters for shaded environments persisted in phylogenetically corrected analysis. The results indicate that clonal integration belongs to a set of evolutionarily conservative plant traits, usually shared by related species. Consequently, the adaptive value of clonal integration in individual habitats remains questionable.     

The physiological basis of adaptation in goats to harsh environments

Goat living in harsh environments represents a climax in the capacity of domestic ruminants to adjust to such areas. This ability is multifactorial: low body mass, and low metabolic requirements of goats can be regarded as an important asset to them for it minimise their maintenance and water requirements, in areas where water sources are widely distributed and food sources are limited by their quantity and quality. An ability to reduce metabolism allows goats to survive even after prolonged periods of severe limited food availability. A skillful grazing behaviour and efficient digestive system enable goats to attain maximal food intake and maximal food utilisation in a given condition. There is a positive interaction between the better recycling rate of urea and a better digestion of such food in desert goats. The rumen plays an important role in the evolved adaptations by serving as a huge fermentation vat and water reservoir. The water stored in the rumen is utilised during dehydration, and the rumen serves as a container, which accommodates the ingested water upon rehydration. The rumen, the salivary glands and the kidney coordinately function in the regulation of water intake and water distribution following acute dehydration and rapid rehydration. Goats in the tropics, when possible, eat a diet composed of tree-leaves and shrubs (browse), which ensure a reliable and steady supply of food all year around, albeit, from a low to medium quality food. Some of the physiological features of ruminants defined as intermediate feeders like large salivary gland, the large absorptive area of their rumen epithelium, and the capacity to change rapidly the volume of the foregut in response to environmental changes are most likely responsible for the goat’s superior digestion capacity.     

Sociosexual environments can drive the evolution of plasticity in mating behavior*

How do male mating behaviors evolve in response to a competitive social environment? Using an experimental evolution approach, Dore et al. demonstrated that sociosexual environments can lead to the evolution of novel plastic male mating behaviors in Drosophila melanogaster, with both mating latency and mating duration extended in male‐biased populations after exposure to male rivals.     

Epiphytic bryophytes in harsh environments: the Juniperus thurifera forests

Abstract

The forests of Juniperus thurifera are peculiar ecosystems that typically grow on mountains and highplateaux of the western Mediterranean basin with dry and continental climates. Some previous surveys suggested that these forests house a rather distinctive epiphytic bryophyte flora. Epiphytic bryophyte communities were systematically sampled in 19 representative juniper forests, for the first time spanning all the distribution area of this conifer. The flora consists of 44 species (32 acrocarpous mosses, 10 pleurocarpous mosses and 2 liverworts). Orthotrichum species are the most frequent and abundant in most of the sampled localities, including some uncommon taxa such as Orthotrichum vittii which shows a clear association with J. thurifera. Epiphytic bryophyte communities on this species were highly homogeneous, although they can be arranged into different groups in response to environmental conditions. Typically, the epiphytic communities of the Mediterranean juniper forests comprise a distinct combination of xerophytic taxa that enhances the interest of these ecosystems and provides new fields for their research and conservation.     

Spatially structured intraspecific trait variation can foster biodiversity in disturbed,heterogeneous environments

Trait variation within populations is an important area of research for empirical and theoretical ecologists. While differences between individuals are doubtlessly ubiquitous, their role for species coexistence is much less clear and highly debated. Both unstructured (random) and structured (linked to space, time or inheritance) intraspecific trait variation (ITV) may modify species interactions with nontrivial consequences for emerging community compositions. In many ecosystems, these compositions are further driven by prevalent disturbance regimes. I therefore explored the effects of unstructured as well as spatially structured ITV under disturbances in a generic ecological model of competing sessile species. Using spatially explicit, individual‐based simulations, I studied how intraspecific variation in life history traits together with interspecific tradeoffs and disturbance regimes shape long‐term community composition. I found that 1) unstructured ITV does not affect species coexistence in the given context, 2) spatially structured ITV may considerably increase coexistence, but 3) spatially clumped disturbances reduce this effect of spatially structured ITV, especially if interspecific tradeoffs involve dispersal distance. The findings suggest that spatially structured ITV with individual trait responses to local habitat conditions differing among species may create or expand humps in disturbance–diversity relationships. Hence, if present, these forms of spatially structured ITV should be included in ecological models and will be important for reliably assessing community responses to environmental heterogeneity and change.     

Temporal variation and the evolution of a parasitoid foraging cue

This work details theory in which selection favors generalists in a more variable environment. Specifically, in a two-host-one-parasitoid model, temporal variation in host abundances alters the optimal searching strategy and leads to the evolution of more generalist parasitoid strategies. Consistent with empirical observations, parasitoids learn host/plant odors, and use them as a cue to search for oviposition sites. The amount of unsuccessful search time required before a parasitoid alters its searching cues (the "giving-up time") is modeled in order to understand the evolutionarily optimal giving-up times under a variety of conditions. When host abundances vary across time, a generalist parasitoid strategy evolves with short giving-up times as it is likely that the host initially favored by a parasitoid will now have a low abundance. In contrast, when populations reach stable dynamics across time, giving-up times typically evolve to longer times, i.e. parasitoids remain specialized longer. The effect of temporal fluctuations is consistent across variation caused by endogenous population interactions and, to some degree, by environmental stochasticity. The conclusions are robust in that there is a strong degree of concordance between the results of a stochastic, individual-based model and a deterministic, numerical model. As an extension, spatial variation in hosts that leads to unequal tradeoffs between generalist parasitoids and specialist parasitoids may also result in the evolution of reduced giving-up times.     

Learning capabilities enhanced in harsh environments: a common garden approach

Previous studies have suggested that the ability to inhabit harsh environments may be linked to advanced learning traits. However, it is not clear if individuals express such traits as a consequence of experiencing challenging environments or if these traits are inherited. To assess the influence of differential selection pressures on variation in aspects of cognition, we used a common garden approach to examine the response to novelty and problem-solving abilities of two populations of black-capped chickadees (Poecile atricapillus). These populations originated from the latitudinal extremes of the species''s range, where we had previously demonstrated significant differences in memory and brain morphology in a multi-population study. We found that birds from the harsh northern population, where selection for cognitive abilities is expected to be high, significantly outperformed conspecifics from the mild southern population. Our results imply differences in cognitive abilities that may be inherited, as individuals from both populations were raised in and had experienced identical environmental conditions from 10 days of age. Although our data suggest an effect independent of experience, we cannot rule out maternal effects or experiences within the nest prior to day 10 with our design. Nevertheless, our results support the idea that environmental severity may be an important factor in shaping certain aspects of cognition.     

The evolution of conformist social learning can cause population collapse in realistically variable environments

Why do societies collapse? We use an individual-based evolutionary model to show that, in environmental conditions dominated by low-frequency variation (“red noise”), extirpation may be an outcome of the evolution of cultural capacity. Previous analytical models predicted an equilibrium between individual learners and social learners, or a contingent strategy in which individuals learn socially or individually depending on the circumstances. However, in red noise environments, whose main signature is that variation is concentrated in relatively large, relatively rare excursions, individual learning may be selected from the population. If the social learning system comes to lack sufficient individual learning or cognitively costly adaptive biases, behavior ceases tracking environmental variation. Then, when the environment does change, fitness declines and the population may collapse or even be extirpated. The modeled scenario broadly fits some human population collapses and might also explain nonhuman extirpations. Varying model parameters showed that the fixation of social learning is less likely when individual learning is less costly, when the environment is less red or more variable, with larger population sizes, and when learning is not conformist or is from parents rather than from the general population. Once social learning is fixed, extirpation is likely except when social learning is biased towards successful models. Thus, the risk of population collapse may be reduced by promoting individual learning and innovation over cultural conformity, or by preferential selection of relatively fit individuals as models for social learning.     

Temporal decision making in complex environments

The basic hypothesis of the author is that under the influence of technological development and market pressure, situations take on temporal characteristics that are more and more difficult for the operator to control. The temporal strategies traditionally installed by the operator disappear, are transferred or transformed. Far from counterbalancing these phenomena, the displays, as they are designed in the workplace, obliterate the temporal dimension. The errors that are seen to appear are the product of a mismatch between the characteristics of the situation and the operator's resources. Four mechanisms of time estimation are discussed. Field study results on temporal strategies, such as anticipation, assessment of a process evolution and planning adjustment are developed.     

Evolution of a single niche specialist in variable environments

The pattern (space versus time) and scale (relative to the lifetime of individuals) of environmental variation is thought to play a central role in governing the evolution of the ecological niche and the maintenance of genetic variance in fitness. To evaluate this idea, we serially propagated an initially genetically uniform population of the bacterium Pseudomonas fluorescens for a few hundred generations in environments that differed in both the pattern and scale at which two highly contrasted carbon substrates were experienced. We found that, contrary to expectations, populations often evolved into a single niche specialist adapted to the less-productive substrate in variable environments and that the genetic variance in fitness across different components of the environment was not generally higher in variable environments when compared with constant environments. We provide evidence to suggest that our results reflect a novel constraint on niche evolution imposed by the supply of beneficial mutations available to selection in variable environments.