Changes In Trait Expression Research Articles

Parental developmental experience affects vocal learning in offspring

Cultural and genetic inheritance combine to enable rapid changes in trait expression, but their relative importance in determining trait expression across generations is not clear. Birdsong is a socially learned cognitive trait that is subject to both cultural and genetic inheritance, as well as being affected by early developmental conditions. We sought to test whether early-life conditions in one generation can affect song acquisition in the next generation. We exposed one generation (F1) of nestlings to elevated corticosterone (CORT) levels, allowed them to breed freely as adults, and quantified their son’s (F2) ability to copy the song of their social father. We also quantified the neurogenetic response to song playback through immediate early gene (IEG) expression in the auditory forebrain. F2 males with only one corticosterone-treated parent copied their social father’s song less accurately than males with two control parents. Expression of ARC in caudomedial nidopallium (NCM) correlated with father-son song similarity, and patterns of expression levels of several IEGs in caudomedial mesopallium (CMM) in response to father song playback differed between control F2 sons and those with a CORT-treated father only. This is the first study to demonstrate that developmental conditions can affect social learning and neurogenetic responses in a subsequent generation.

The Indirect Genetic Effect Interaction Coefficient ψ: Theoretically Essential and Empirically Neglected.

The interaction effect coefficient ψ has been a much-discussed, fundamental parameter of indirect genetic effect (IGE) models since its formal mathematical description in 1997. The coefficient simultaneously describes the form of changes in trait expression caused by genes in the social environment and predicts the evolutionary consequences of those IGEs. Here, we report a striking mismatch between theoretical emphasis on ψ and its usage in empirical studies. Surveying all IGE research, we find that the coefficient ψ has not been equivalently conceptualized across studies. Several issues related to its proper empirical measurement have recently been raised, and these may severely distort interpretations about the evolutionary consequences of IGEs. We provide practical advice on avoiding such pitfalls. The majority of empirical IGE studies use an alternative variance-partitioning approach rooted in well-established statistical quantitative genetics, but several hundred estimates of ψ (from 15 studies) have been published. A significant majority are positive. In addition, IGEs with feedback, that is, involving the same trait in both interacting partners, are far more likely to be positive and of greater magnitude. Although potentially challenging to measure without bias, ψ has critically-developed theoretical underpinnings that provide unique advantages for empirical work. We advocate for a shift in perspective for empirical work, from ψ as a description of IGEs, to ψ as a robust predictor of evolutionary change. Approaches that “run evolution forward” can take advantage of ψ to provide falsifiable predictions about specific trait interactions, providing much-needed insight into the evolutionary consequences of IGEs.

Sex and morph differences in age-dependent trait changes in a polymorphic songbird.

There is growing evidence of important variation in how animals age, in particular in how the expression of traits changes with age among different species and populations. However, less is known about variation within populations, which may include variation in ageing patterns between different types of individuals (e.g. sexes or distinct polymorphisms) and between different types of traits (e.g. general traits versus those used in social signalling contexts). We used 6years of longitudinal data to examine age-related changes in trait expression in a captive population of Gouldian finches (Erythrura gouldiae), a socially monogamous songbird with genetically determined colour morphs that differ in behaviour and physiology. We contrasted ageing patterns of different types of traits (social signalling vs. size-related) in both sexes and in two colour morphs, using a mixed model approach to account for both within- and between-individual effects. We found pronounced sex differences in how social signalling traits change with age, showing a quadratic pattern in males, but not changing with age in females. In contrast, we observed no sex-specific ageing patterns in size traits. We also found subtle morph differences in how size-related traits changed with age, with black morphs stable or increasing with age while red morphs showing a decline with age. Finally, we found an interesting sex by morph interaction in one important social signal (headband width). These results highlight the importance of using within-individual approaches to understand ageing patterns across types of individuals (sex, morph, etc.) and the need for further research on the ageing patterns of traits that may experience different selective pressures.

Bumblebee Behavior on Flowers, but Not Initial Attraction, Is Altered by Short-Term Drought Stress.

Climate change is leading to increasing drought and higher temperatures, both of which reduce soil water levels and consequently water availability for plants. This reduction often induces physiological stress in plants, which in turn can affect floral development and production inducing phenotypic alterations in flowers. Because flower visitors notice and respond to small differences in floral phenotypes, changes in trait expression can alter trait-mediated flower visitor behavior. Temperature is also known to affect floral scent emission and foraging behavior and, therefore, might modulate trait-mediated flower visitor behavior. However, the link between changes in flower visitor behavior and floral traits in the context of increasing drought and temperature is still not fully understood. In a wind-tunnel experiment, we tested the behavior of 66 Bombus terrestris individuals in response to watered and drought-stressed Sinapis arvensis plants and determined whether these responses were modulated by air temperature. Further, we explored whether floral traits and drought treatment were correlated with bumblebee behavior. The initial attractiveness of drought and watered plants did not differ, as the time to first visit was similar. However, bumblebees visited watered plants more often, their visitation rate to flowers was higher on watered plants, and bumblebees stayed for longer, indicating that watered plants were more attractive for foraging. Bumblebee behavior differed between floral trait expressions, mostly independently of treatment, with larger inflorescences and flowers leading to a decrease in the time until the first flower visit and an increase in the number of visits and the flower visitation rate. Temperature modulated bumblebee activity, which was highest at 25°C; the interaction of drought/water treatment and temperature led to higher visitation rate on watered plants at 20°C, possibly as a result of higher nectar production. Thus, bumblebee behavior is influenced by the watered status of plants, and bumblebees can recognize differences in intraspecific phenotypes involving morphological traits and scent emission, despite overall morphological traits and scent emission not being clearly separated between treatments. Our results indicate that plants are able to buffer floral trait expressions against short-term drought events, potentially to maintain pollinator attraction.

Secular change in morphological cranial and mandibular trait frequencies in European Americans born 1824-1987.

Secular change in cranial and postcranial morphometrics and morphological traits has been documented in several studies. However, to date, few studies have addressed temporal changes occurring in the expression of cranial morphological traits commonly used in ancestry estimation. This study examines secular change in the expression of 23 cranial and mandibular morphological traits; accounting for age-at-death, sex, and year-of-birth. Data were collected on 23 morphological cranial and mandibular traits for European American individuals (19-97 years of age) from the Hamann-Todd Skeletal Collection (n = 518) and the William M. Bass Donated Skeletal Collection (n = 602). Individuals were divided into six birth-year cohorts: 1824-1849 (Cohort 1), 1850-1874 (Cohort 2), 1875-1899 (Cohort 3), 1900-1924 (Cohort 4), 1925-1949 (Cohort 5), and 1950-1987 (Cohort 6). Statistical analyses, including Pearson's chi-square, correspondence analysis, and ordinal regression, demonstrate that secular changes have occurred in 11 traits, including: anterior nasal spine (ANS); malar tubercle (MT); nasal bone contour (NBC); postbregmatic depression (PBD); supranasal suture (SPS); transverse palatine suture (TPS); zygomaticomaxillary suture (ZS); ascending ramus shape (ARS); gonial angle flare (GAF); mandibular tori (MDT); and posterior ramus edge inversion (PREI), with changes occurring in both sexes for ANS, MT, TPS, ZS, GAF, MDT, and PREI. Significant changes in trait expression were found predominately between Cohorts 3 and 4, and Cohorts 4 and 5. While the sex of an individual affected the expression of ANS, MT, NBC, PBD, SPS, ZS, ARS, GAF, and PREI, age-at-death only affected MT and PREI. This study demonstrates that secular change in morphological cranial and mandibular traits has occurred over the last two centuries in European Americans, with the most considerable change appearing at the turn of the twentieth century. Changes in morphological trait expression over a relatively short period of time correspond with changes seen in craniometric analyses and correlate with the industrialization of society and environmental and cultural changes, such as medical advancements, nutrition, and population health/stress.

Adaptive phenotypic divergence in an annual grass differs across biotic contexts*

Climate is a powerful force shaping adaptation within species, yet adaptation to climate occurs against a biotic background: species interactions can filter fitness consequences of genetic variation by altering phenotypic expression of genotypes. We investigated this process using populations of teosinte, a wild annual grass related to maize (Zea mays ssp. mexicana), sampling plants from 10 sites along an elevational gradient as well as rhizosphere biota from three of those sites. We grew half-sibling teosinte families in each biota to test whether trait divergence among teosinte populations reflects adaptation or drift, and whether rhizosphere biota affect expression of diverged traits. We further assayed the influence of rhizosphere biota on contemporary additive genetic variation. We found that adaptation across environment shaped divergence of some traits, particularly flowering time and root biomass. We also observed that different rhizosphere biota shifted expressed values of these traits within teosinte populations and families and altered within-population genetic variance and covariance. In sum, our results imply that changes in trait expression and covariance elicited by rhizosphere communities could have played a historical role in teosinte adaptation to environments and that they are likely to play a role in the response to futureselection.

The reciprocal relationship between competition and intraspecific trait variation

Summary Trait differences among plants are expected to influence the outcome of competition; competition should be strongest between similar species (or individuals) under limiting similarity, and between dissimilar species within competitive hierarchies. These hypotheses are often used to infer competitive dynamics from trait patterns within communities. However, plant traits are frequently plastic in response to competition. This variation is poorly accounted for in trait‐based studies of competition and community assembly. To explore the relationship between trait responses and competitive outcomes, we grew 15 species alone, in monoculture and in mixture. We measured traits relating to leaf and root tissue morphology as well as biomass allocation and related competition‐induced changes in these traits to intra‐ and interspecific competition using multi‐model inference. Additionally, we tested how traits from different competitive environments influenced potential community assembly inferences. The competitive environment had large effects on species’ traits, although many effects were species specific. Differences among species in how competition affected trait expression were linked to both intra‐ and interspecific competition, frequently affecting competitive hierarchies. Intraspecific competition was lower for species that limited competition‐induced increases in root allocation and had less variability in this trait overall. Interspecific competition was lower for species with larger leaves and lower specific leaf area than their neighbours. Switching to more stress‐tolerant strategies by increasing root diameter and leaf tissue density also reduced competition. However, dissimilarity in root tissue density also minimized competition, consistent with limiting similarity affecting competitive outcomes. Moreover, changes in these traits were linked to changes in functional diversity, suggesting that competition affects functional diversity by affecting trait expression. Synthesis. Both trait hierarchies and trait dissimilarity affect the outcome of competition by acting on different traits, although competition‐induced changes in trait expression can alter competitive outcomes. Moreover, the magnitude of these trait changes suggests that the source environment where plant traits are collected can affect the inferences drawn from trait patterns within communities. Combined, our results suggest that considering the effect of competition on trait expression is critical to understanding the relationship between traits and community assembly.

Simultaneous age-dependent and age-independent sexual selection in the lekking black grouse (Lyrurus tetrix).

Individuals' reproductive success is often strongly associated with their age, with typical patterns of early-life reproductive improvement and late-life senescence. These age-related patterns are due to the inherent trade-offs between life-history traits competing for a limited amount of resources available to the organisms. In males, such trade-offs are exacerbated by the resource requirements associated with the expression of costly sexual traits, leading to dynamic changes in trait expression throughout their life span. Due to the age dependency of male phenotypes, the relationship between the expression of male traits and mating success can also vary with male age. Hence, using longitudinal data in a lekking species with strong sexual selection - the black grouse Lyrurus tetrix - we quantified the effects of age, life span and age of first lek attendance (AFL) on male annual mating success (AMS) to separate the effects of within-individual improvement and senescence on AMS from selective (dis)appearance of certain phenotypes. Then, we used male AMS to quantify univariate and multivariate sexual selection gradients on male morphological and behavioural traits with and without accounting for age and age-related effects of other traits. Male AMS increased with age, and there was no significant reproductive senescence. Most males never copulated, and of the ones that did, the majority had only one successful year. Life span was unrelated to AMS, but early AFL tended to lead to higher AMS at ages 1-3. AMS was related to most morphological and behavioural traits when male age was ignored. Accounting for age and age-specific trait effects (i.e. the interaction between a trait and age) reduced the magnitude of the selection gradients and revealed that behavioural traits are under consistent sexual selection, while sexual selection on morphological traits is stronger in old males. Therefore, sexual selection in black grouse operates primarily on male behaviour and morphological traits may act as additional cues to supplement female choice. These results demonstrate the multifaceted influence of age on both fitness and sexual traits and highlight the importance of accounting for such effects when quantifying sexual selection.

Secular Change in Morphological Pelvic Traits used for Sex Estimation.

This research evaluates secular change in Phenice's (Am J Phys Anthropol, 30, 1969 and 297) three morphological traits of the pubis, as described by Klales et al. (Am J Phys Anthropol, 149, 2012 and 104): medial aspect of the ischio-pubic ramus, subpubic contour, and ventral arc. Ordinal scores were collected for these traits and compared between a sample of innominates from the historical Hamann-Todd Collection (n = 170) and modern Bass Donated Collection (n = 129). Using the Freeman-Halton test, significant differences between temporal sample score frequencies were found for all traits in females and for the subpubic contour and ventral arc in males. Despite these findings, classification accuracy using logistic regression between the temporal periods remained low (68.7%). These results suggest that secular changes in trait expression are occurring; however, sex estimation methods using these traits and created with historical samples are still applicable to modern forensic cases. In fact, the secular changes occurring in these traits contribute to better classification accuracy between sexes in modern populations.

Shade affects responses to drought and flooding - acclimation to multiple stresses in bittersweet (Solanum dulcamara L.).

Plants exposed to environmental stress often respond by a change in their phenotypic traits. These changes in trait expression may alleviate the negative effect of such stress factors. However, if multiple stresses are present, responses are likely to be less predictable and hence do not necessarily correlate to plant performance. This study tested if this expectation was true, by subjecting Solanum dulcamara plants to various simultaneous stress factors. Plants were grown in well-watered conditions, drought or flooding, and exposed to either full light or shade for 4 weeks. Shoot and root biomass, stem morphological parameters, such as height, number of nodes and length of stem internodes, and leaf traits like length, specific leaf area, chlorophyll content and stomatal conductance were determined. Both variation in light and in water availability typically caused slower growth, and resulted in distinct phenotypic changes in stem, leaf and root traits. However, effects of stresses on the expression of traits were not always additive. Instead, some combined stress responses (e.g. leaf size) appeared to be limited by physical or physiological constraints, whereas other responses were opposite to each other (e.g. root:shoot ratio), resulting in an intermediate phenotype in the combined stress treatment. These data suggest that in natural conditions, where combined stress factors are likely to be present, the optimal phenotype may not necessarily be expressed. Responses of plants to multiple stress factors may therefore not be associated with immediate advantages in terms of increased performance.

DIFFERENTIAL AGING OF BITE AND JUMP PERFORMANCE IN VIRGIN AND MATED TELEOGRYLLUS COMMODUS CRICKETS

Evolutionary theories of aging state that the force of natural selection declines with age, resulting in trait senescence. However, sexual selection theory predicts that costly traits that signal mate value should increase in expression as survival prospects decline. Mortality rates and fertility tend to show strong signatures of senescence, whereas sexual signaling traits increase with age, but how the expression of traits such as whole-organism performance measures that are subject to both sexual and nonsexual selection should change with age is unclear. We examined the effects of both a key life-history event (mating) and diet quality (male and female optimal diets) on aging in two whole-organism performance traits (bite force and jump take-off velocity) in male and female Teleogryllus commodus crickets. We found no evidence for diet effects on any of the measured traits. Aging effects were more evident in females than in males for both jumping and biting, and constitute a mix of senescence and terminal investment patterns depending on sex/mating class. Sex and mating therefore have important implications for resource allocation to performance traits, and hence for aging of those traits, and interactions between these two factors can result in complex changes in trait expression over individual lifetimes.

Variation in flooding-induced morphological traits in natural populations of white clover (Trifolium repens) and their effects on plant performance during soil flooding.

Soil flooding leads to low soil oxygen concentrations and thereby negatively affects plant growth. Differences in flooding tolerance have been explained by the variation among species in the extent to which traits related to acclimation were expressed. However, our knowledge of variation within natural species (i.e. among individual genotypes) in traits related to flooding tolerance is very limited. Such data could tell us on which traits selection might have taken place, and will take place in future. The aim of the present study was to show that variation in flooding-tolerance-related traits is present among genotypes of the same species, and that both the constitutive variation and the plastic variation in flooding-induced changes in trait expression affect the performance of genotypes during soil flooding. Clones of Trifolium repens originating from a river foreland were subjected to either drained, control conditions or to soil flooding. Constitutive expression of morphological traits was recorded on control plants, and flooding-induced changes in expression were compared with these constitutive expression levels. Moreover, the effect of both constitutive and flooding-induced trait expression on plant performance was determined. Constitutive and plastic variation of several morphological traits significantly affected plant performance. Even relatively small increases in root porosity and petiole length contributed to better performance during soil flooding. High specific leaf area, by contrast, was negatively correlated with performance during flooding. The data show that different genotypes responded differently to soil flooding, which could be linked to variation in morphological trait expression. As flooded and drained conditions exerted different selection pressures on trait expression, the optimal value for constitutive and plastic traits will depend on the frequency and duration of flooding. These data will help us understanding the mechanisms affecting short- and long-term dynamics in flooding-prone ecosystems.

Does the differential seedling mortality caused by slugs alter the foliar traits and subsequent susceptibility of hybrid willows to a generalist herbivore?

Abstract1. Many Salicaceae species naturally form hybrid swarms with parental and hybrid taxa that differ in secondary chemical profile and in resistance to herbivores. Theoretically, the differential mortality in the seedling stage can lead to changes in trait expression and alter subsequent interactions between plants and herbivores. This study examines whether herbivory by the generalist slugArion subfuscus, which causes extensive mortality in young willow seedlings, causes shifts in (a) the foliar chemistry of F2 willow hybrids (Salix sericeaandSalix eriocephala), and (b) the subsequent susceptibility to Japanese Beetles,Popillia japonica.2. In 2001, two populations of F2 seedlings were generated: those that survived slug herbivory (80–90% of seedlings placed in the field were killed by the slugs) were designated as S‐plants, whereas C‐plants (controls) experienced no mortality.3. Common garden experiments with cuttings from these populations, in 2001 and 2002, revealed extensive variation in the phenolic chemistry of F2 hybrids, but revealed no significant difference between S‐ and C‐plants, although the levels of foliar nutrients, proteins and nitrogen tended to be higher in S‐plants.4. Concentrations of salicortin and 2′‐cinnamoylsalicortin explained 55 and 38% of the the variation in leaf damage caused by Japanese beetles, and secondary chemistry was highly correlated within replicate clones (salicortinR2= 0.85, 2‐cinnamoylsalicortinR2= 0.77, condensed tanninsR2= 0.68).5. Interestingly, Japanese beetle damage and condensed tannins were positively correlated within the S‐plants, but not in the C‐plants, suggesting that slugs had selected for plants with a positive relationship between tannins andP. japonicadamage. This is unlikely to be a consequence of a preference for tannins, but is suggested to be related to the elevated nutrient levels in the S‐plants, perhaps in combination with the complex‐binding properties of tannins.6. The damage was highly correlated within replicate clones and a model choice analysis suggested that Japanese beetle damage may be explained by four factors: concentrations of salicortin, condensed tannins, and nitrogen, as well as the specific leaf area (thick leaves were damaged less).

Kittiwakes strategically reduce investment in replacement clutches

Many life-history traits are expressed interactively in life, but to a varying extent on different occasions. Changes in trait expression can be accounted for by differences in the quality of the environment ('environmental constraint' hypothesis) or by strategic adjustments, if the relative contribution of the trait to fitness varies with time ('strategic allocation' hypothesis). In birds, egg production is lower in replacement clutches than in first clutches, but it is unknown whether this reduction results from an environmental constraint (e.g. food being less available at the time when the replacement clutch is produced) or from a strategic allocation of resources between the two breeding attempts. To distinguish between these two hypotheses, we performed an experiment with black-legged kittiwakes (Rissa tridactyla). Pairs were either food-supplemented or not before the first clutch was laid onwards and we induced them to produce a replacement clutch by removing eggs once when the first clutch was complete. As predicted by the 'strategic allocation' hypothesis, egg production of food-supplemented and non-food-supplemented birds decreased between first and replacement clutches. This suggests that kittiwakes strategically reduce investment in egg production for their replacement clutches compared to first clutches.