Evidence For Constraints Research Articles

Evidence for constraints on probing dynamics: A case study of adjectival concord

AbstractBased on facts of adjectival concord in Standard Arabic, this article offers evidence that upward probing (i.e., the goal c-commands the probe) is permitted only if downward probing (i.e., where the probe c-commands the goal) does not result in valuing the probe's uninterpretable feature. Such a constraint on upward probing allows us to account for several intriguing observations in Arabic grammar, including the fact that an adjective can agree in number and gender with one nominal, but in definiteness with another nominal. Hence, on the one hand, this article lends support to Agree proposals according to which absence of a match in the c-command domain of an unvalued feature (uF) is not fatal to the derivation. On the other hand, it speaks against Agree proposals that do not license downward probing or view it as parasitic on upward probing.

Rapid-cycling Brassica rapa evolves even earlier flowering under experimental drought.

Changes in climate can impose selection on populations and may lead to rapid evolution. One such climatic stress is drought, which plant populations may respond to with escape (rapid growth and early flowering) or avoidance (slow growth and efficient water-use). However, it is unclear if drought escape would be a viable strategy for populations that already flower early from prior selection. In an experimental evolution study, we subjected rapid-cycling Brassica rapa (RCBr), which was previously selected for early flowering, to four generations of experimental drought or watered conditions. We then grew ancestral and descendant populations concurrently under drought and watered conditions to assess evolution, plasticity, and adaptation. The RCBr populations that evolved under drought had earlier flowering and lower water-use efficiency than the populations that evolved under watered conditions, indicating evolutionary divergence. The drought descendants also had a trend of earlier flowering compared to ancestors, indicating evolution. Evolution of earlier flowering under drought followed the direction of selection and increased fitness and was consistent with studies in natural and experimental populations of this species, suggesting adaptive evolution. We found rapid adaptive evolution of drought escape in RCBr and little evidence for constraints on flowering time evolution, even though RCBr already flowers extremely early. Our results suggest that some populations may harbor sufficient genetic variation for evolution even after strong selection has occurred. Our study also illustrates the utility of combining artificial selection, experimental evolution, and the resurrection approach to study the evolution of functional traits.

No Evidence of Robust Noun-Referent Associations in German-Learning 6- to 14-Month-Olds.

Work with the looking-while-listening (LWL-) paradigm suggested that 6-month-old English-learning infants associated several labels for common nouns with pictures of their referents: While one distractor picture was present, infants systematically fixated the named target picture. However, recent work revealed constraints of infants' noun comprehension. The age at which these abilities can be obtained appears to relate to the infants' familiarity with the talker, the target language, and word frequency differences in target-distractor pairs. Here, we present further data to this newly established field of research. We tested 42 monolingual German-learning infants aged 6–14 months by means of the LWL-paradigm. Infants saw two pictures side-by-side on a screen, whilst an unfamiliar male talker named one of both. Overall, infants did not fixate the target picture more than the distractor picture. In line with previous results, infants' performance on the task was higher when target and distractor differed within their word frequency—as operationalized by the parental rating of word exposure. Together, our results add further evidence for constraints on early word learning. They point to cross-linguistic differences in early word learning and strengthen the view that infants might use extra-linguistic cues within the stimulus pairing, such as frequency imbalance, to disambiguate between two potential referents.

Parallelism in Flower Evolution and Development

Flower evolution is characterized by widespread repetition, with adaptations to pollinator environment evolving in parallel. Recent studies have expanded our understanding of the developmental basis of adaptive floral novelties—petal fusion, bilateral symmetry, heterostyly, and floral dimensions. In this article, we describe patterns of trait evolution and review developmental genetic mechanisms underlying floral novelties. We discuss the diversity of mechanisms for parallel adaptation, the evidence for constraints on these mechanisms, and how constraints help explain observed macroevolutionary patterns. We describe parallel evolution resulting from similarities at multiple hierarchical levels—genetic, developmental, morphological, functional—which indicate general principles in floral evolution, including the central role of hormone signaling. An emerging pattern is mutational bias that may contribute to rapid patterns of parallel evolution, especially if the derived trait can result from simple degenerative mutations. We argue that such mutational bias may be less likely to govern the evolution of novelties patterned by complex developmental pathways.

Non-Linguistic Grammar Learning by 12-Month-Old Infants: Evidence for Constraints on Learning

ABSTRACTInfants acquiring their native language are adept at discovering grammatical patterns. However, it remains unknown whether these learning abilities are limited to language, or available more generally for sequenced input. The current study is a conceptual replication of a prior language study, and was designed to ask whether infants can track phrase structure-like patterns from nonlinguistic auditory materials (sequences of computer alert sounds). One group of 12-month-olds was familiarized with an artificial grammar including predictive dependencies between sounds concatenated into strings, simulating the basic structure of phrases in natural languages. A second group of infants was familiarized with a grammar that lacked predictive dependencies. All infants were tested on the same set of familiar strings vs. novel (grammar-inconsistent) strings. Only infants exposed to the materials containing predictive dependencies showed successful discrimination between the test sentences, replicating the results from linguistic materials, and suggesting that predictive dependencies facilitate learning from nonlinguistic input.

Bank Lending and Asset Prices in the Euro Area

We examine the dynamics of bank lending to the private sector for countries of the Euro area by applying a Markov switching error correction model. We identify for Belgium, Germany, Ireland and Portugal stable, mean reverting regimes and unstable regimes with no tendency to return to the long term credit demand equation, whereas for some other countries there is only weak evidence. Furthermore, for these as well as for other countries we detect in the less stable regimes a strong co-movement with the development of the stock market. We interpret this as evidence for constraints in bank lending. In contrast, the banks' capital seems to have only marginal impact on the lending behaviour.

Additive genetic variation in resistance traits of an exotic pine species: little evidence for constraints on evolution of resistance against native herbivores.

The apparent failure of invasions by alien pines in Europe has been explained by the co-occurrence of native pine congeners supporting herbivores that might easily recognize the new plants as hosts. Previous studies have reported that exotic pines show reduced tolerance and capacity to induce resistance to those native herbivores. We hypothesize that limited genetic variation in resistance to native herbivores and the existence of evolutionary trade-offs between growth and resistance could represent additional potential constraints on the evolution of invasiveness of exotic pines outside their natural range. In this paper, we examined genetic variation for constitutive and induced chemical defences (measured as non-volatile resin in the stem and total phenolics in the needles) and resistance to two major native generalist herbivores of pines in cafeteria bioassays (the phloem-feeder Hylobius abietis and the defoliator Thaumetopoea pityocampa) using half-sib families drawn from a sample of the population of Pinus radiata introduced to Spain in the mid-19th century. We found (i) significant genetic variation, with moderate-to-high narrow-sense heritabilities for both the production of constitutive non-volatile resin and induced total phenolics, and for constitutive resistance against T. pityocampa in bioassays, (ii) no evolutionary trade-offs between plant resistance and growth traits or between the production of different quantitative chemical defences and (iii) a positive genetic correlation between constitutive resistance to the two studied herbivores. Overall, results of our study indicate that the exotic pine P. radiata has limited genetic constraints on the evolution of resistance against herbivores in its introduced range, suggesting that, at least in terms of interactions with these enemies, this pine species has potential to become invasive in the future.

Factors affecting cork oak growth under dry conditions: local adaptation and contrasting additive genetic variance within populations

Increased drought severity is expected in the Mediterranean Basin over the twenty-first century, but our understanding of the potential of most forest tree species to cope with it remains uncertain. In this study, (1) we examined the potential effects of long-term selection and the capacity to respond to future changes in selective pressures in three populations of cork oak (Quercus suber L.). For this purpose, we evaluated the response to dry conditions of 45 open-pollinated trees originating from populations in Morocco, Portugal, and Spain. Growth, leaf size, specific leaf area (SLA), carbon isotope discrimination (Δ13C), leaf nitrogen content (Nmass), and total chlorophyll content (Chlmass) were measured in 9-year-old plants. (2) We also investigated the relationships between functional traits and aboveground growth by regression models. Plants presenting larger and more sclerophyllous leaves (low SLA and high leaf thickness) exhibited higher growths, with results suggesting that these traits are subjected to divergent selection in this species. Heritability estimates were moderately high for Δ13C (0.43 ± 0.25–0.83 ± 0.31) and stem diameter (0.40 ± 0.15–0.71 ± 0.28) for the tree populations. For the rest of the traits (except for annual growth), heritability values varied among populations, particularly for height, leaf size, leaf thickness, and Nmass. Our results suggest that natural selection has led to local adaptations and has also affected the genetic variance intrapopulation in these cork oak populations, although studies with a higher number of populations should be carried out across different years. Additionally, the absence of significant genetic correlations and the fact that correlated traits did not undergo opposing selection provided little evidence for constraints on evolution caused by genetic correlations.

Correlation of microsynteny conservation and disease gene distribution in mammalian genomes

BackgroundWith the completion of the whole genome sequence for many organisms, investigations into genomic structure have revealed that gene distribution is variable, and that genes with similar function or expression are located within clusters. This clustering suggests that there are evolutionary constraints that determine genome architecture. However, as most of the evidence for constraints on genome evolution comes from studies on yeast, it is unclear how much of this prior work can be extrapolated to mammalian genomes. Therefore, in this work we wished to examine the constraints on regions of the mammalian genome containing conserved gene clusters.ResultsWe first identified regions of the mouse genome with microsynteny conservation by comparing gene arrangement in the mouse genome to the human, rat, and dog genomes. We then asked if any particular gene types were found preferentially in conserved regions. We found a significant correlation between conserved microsynteny and the density of mouse orthologs of human disease genes, suggesting that disease genes are clustered in genomic regions of increased microsynteny conservation.ConclusionThe correlation between microsynteny conservation and disease gene locations indicates that regions of the mouse genome with microsynteny conservation may contain undiscovered human disease genes. This study not only demonstrates that gene function constrains mammalian genome organization, but also identifies regions of the mouse genome that can be experimentally examined to produce mouse models of human disease.

Hierarchy and the reconstruction of evolutionary trends: evidence for constraints on the evolution of body size in terrestrial caniform carnivorans (Mammalia)

Body mass is an important organism-level variable in mammalian biology, correlated with physiology, life history, and ecology. To analyze the dynamics of body size evolution, increases and decreases in body mass were tallied for ancestor-descendant (AD) species pairs for 519 terrestrial caniform taxa. To account for uncertainty phylogeny, a bootstrapping routine shuffled hypothesized AD pairs, and average proportions of increases were binned as a function of ancestral body mass. A set of models relating the rate of body size increase were evaluated with the Akaike Information Criterion (AIC). AIC selected three models of the candidate set as equivalent in support by the observed body mass data. These three models propose body size increase for small AD pairs and body size decrease for large AD pairs, although they differ in their treatment of taxa at intermediate sizes.These results demonstrate the presence of constraints bounding the caniform distribution at large and small body sizes, stabilizing the distribution through time, which stands in contrast to a broader mammalian pattern. At a finer phylogenetic scale, subclades within intermediate size classes display proportions that are significantly different from unbiased, with several clades previously cited as examples of “Cope's Rule” showing biased increases in size, and basal mustelids (badgers, and allied genera), Mephitidae (skunks), and Vulpini (“foxes”) exhibiting biased decreases. The caniform pattern is therefore the result of superimposed, clade-specific trajectories, demonstrating that the inferred dynamics of body size evolution and even the direction of trends in body size evolution within the Caniformia, and for mammals in general, depend on the hierarchical scale of the analysis.

On the evolution of clonal plant life histories

Clonal plant life histories are special in at least four respects: (1) Clonal plants can also reproduce vegetatively, (2) vegetative reproduction can be realised with short or long spacers, (3) and it may allow to plastically place vegetative offspring in benign patches. (4) Moreover, ramets of clonal plants may remain physically and physiologically integrated. Because of the apparent utility of such traits and because ecological patterns of distribution of clonal and non-clonal plants differ, adaptation is a tempting explanation of observed clonal life-history variation. However, adaptive evolution requires (1) heritable genetic variation and (2) a trait effect on fitness, and (3) it may be constrained if other evolutionary forces are overriding selection or by constraints, costs and trade-offs. (1) The few studies undertaken so far reported broad-sense heritability for clonal traits. Variation in selectively neutral genetic markers appears as pronounced in populations of clonal as non-clonal plants. However, neutral markers may not reflect heritable variation of life-history traits. Moreover, clonal plants may have been sampled at larger spatial scales. Empirical information on the contribution of somatic mutations to heritable variation is lacking. (2) Clonal life-history traits were found to affect fitness. However, much of this evidence stems from artificial rather than natural environments. (3) The relative importance of gene flow, inbreeding, and genetic drift, compared with selection, in the evolution of clonal life histories is hardly explored. Benefits of clonal life-history traits were frequently studied and found. However, there is also evidence for constraints, trade-offs, and costs. In conclusion, though it is very likely, that clonal life-history traits are adaptive, it is neither clear to which degree this is the case, nor which clonal life-history traits constitute adaptations to which environmental factors. Moreover, evolutionary interactions among clonal life-history traits and between clonal and non-clonal ones, such as the mating system, are not well explored. There remains much interesting work to be done in this field – which will be particularly interesting if it is done in the field.

Ontogenetic constraints on neural and behavioral plasticity: evidence from imprinting and face processing.

This paper addresses the extent and limits on brain plasticity during development through the detailed study of imprinting in the domestic chick and the development of face processing in human infants. In both of these systems, evidence for constraints on plasticity is reviewed. The first source of constraint comes from the basic architecture of learning mechanisms that support plasticity. With regard to the chick, a specific "Hebbian" model based on the known neural circuitry of the region of the brain involved is presented and discussed. In human infants, a more abstract model inspired by cortical circuitry is mentioned. The second source of constraint comes from biases on the nature of the stimuli selected for attention by the young organism. Both in the chick and the human there is evidence for a subcortical brain system which orients their attention toward conspecifics, and particularly to their faces. It is argued that these systems tutor, or bias the input to, the more plastic learning systems.

GENETIC CONSTRAINTS AND THE PHYLOGENY OF INSECT-PLANT ASSOCIATIONS: RESPONSES OF OPHRAELLA COMMUNA (COLEOPTERA: CHRYSOMELIDAE) TO HOST PLANTS OF ITS CONGENERS.

We ask whether patterns of genetic variation in a phytophagous insect's responses to potential host plants shed light on the phylogenetic history of host association. Ophraella communa feeds chiefly, and in eastern North America exclusively, on Ambrosia (Asteraceae: Ambrosiinae). Using mostly half-sib breeding designs, we screened for genetic variation in feeding responses to and larval survival on its own host and on seven other plants that are hosts (or, on one case, closely related to the host) of other species of Ophraella. We found evidence for genetic variation in feeding responses to five of the seven test plants, other than the natural host. We found no evidence of genetic variation in feeding responses to two plant species, nor in capacity for larval survival on six. These results imply constraints on the availability of genetic variation; however, little evidence for constraints in the form of negative genetic correlations was found. These results are interpreted in the context of a provisional phylogeny of, and a history of host shifts within, the genus. Ophraella communa does not present evidence of genetic variation in its ability to feed and/or survive on Solidago, even though it is probably descended from a lineage that fed on Solidago or related plants, possibly as recently as 1.9 million years ago. Genetic variation in performance on this plant may have been lost. Based on evidence for genetic variation and on mean performance, by far the greatest potentiality for adaptation to a congener's host was evinced in responses to Iva frutescens, which not only is related and chemically similar to Ambrosia, but also is the host of a closely related species of Ophraella that may have been derived from an Ambrosia-associated ancestor. Genetic variation in O. communa's capacity to feed and/or survive on its congeners' hosts is less evident for plants that do not represent historically realized host shifts (with one exception) than for those that may (but see Note Added in Proof). The results offer some support for the hypothesis that the evolution of host shifts has been guided in part by constrained genetic variation.

Temperature regulation of young coypus (Myocastor coypus) in air and water

Coypus (Myocastor coypus Molina) have recently colonized temperate regions of Europe in large numbers, originating from populations indigenous to subtropical regions of South America. Observations of coypu behavior in the wild and in enclosures under a temperate climate showed a preference for close proximity to water and frequent swimming bouts. Little change was observed in the semiaquatic habit in winter, despite high mortality during this period, particularly among young and adult male coypus. Evidence for constraints on thermal adaptation was sought from measurements in a thermostatic chamber of the resting metabolic rate (RMR) of young coypus caught from wild populations in France and of the body (Tb) and skin (Tsk) temperatures as a function of ambient temperature (Ta). A light heterothermic response with deep Tb falling by 0.8 degree C and stabilizing at a new level of 36.7 degrees C was observed in water and air at Ta below the lower critical temperatures. In common with other semiaquatic mammals, RMR (2.94 W/kg) was high relative to terrestrial mammals of equivalent body mass. Minimal thermal conductance was 0.10 W.kg-1.degree C-1 in air for Ta between -10 and +20 degrees C, and conductance varied in water from 0.24 to 1.18 W.kg-1.degrees C-1 for Ta between 5 and 35 degrees C. The physiological adaptations to cold and particularly the high insulative value of the fur (80-90% of the insulation) help to explain the successful extension of the coypu's range to temperate regions. Energetic costs were nevertheless high in cold water, suggesting that social factors may be implicated in severe winter mortality, particularly dominance hierarchy regulating the persistent use of water bodies even under freezing conditions.

Altered VH gene segment utilization in the response to phosphorylcholine by aged mice.

Newly generated bone marrow B cell precursors of aged BALB/c mice, stimulated in splenic fragment cultures, display a markedly increased frequency of phosphorylcholine (PC)-responsive cells. This increased frequency is found for both precursors that utilize VHS107, a phenotype common to essentially all PC-specific B cells of young mice, and, surprisingly, for precursors that utilize VH genes other than VHS107. PC-specific hybridomas derived from bone marrow cells of aged mice utilize members of at least three VH gene segment families that have never been observed in PC responses of young mice. The ability of aged but not young mice to generate these unique PC-specific clonotypes may be evidence for constraints on V region utilization during repertoire development in young adults and has important implications for aging-associated changes in immune responsiveness.