CONNECTING QTLS TO THE G-MATRIX OF EVOLUTIONARY QUANTITATIVE GENETICS

Evolutionary quantitative genetics has recently advanced in two distinct streams. Many biologists address evolutionary questions by estimating phenotypic selection and genetic (co)variances ( G matrices). Simultaneously, an increasing number of studies have applied quantitative trait locus (QTL) mapping methods to dissect variation. Both conceptual and practical difficulties have isolated these two foci of quantitative genetics. A conceptual integration follows from the recognition that QTL allele frequencies are the essential variables relating the G -matrix to marker-based mapping experiments. Breeding designs initiated from randomly selected parental genotypes can be used to estimate QTL-specific genetic (co)variances. These statistics appropriately distill allelic variation and provide an explicit population context for QTL mapping estimates. Within this framework, one can parse the G -matrix into a set of mutually exclusive genomic components and ask whether these parts are similar or dissimilar in their respective features, for example the magnitude of phenotypic effects and the extent and nature of pleiotropy. As these features are critical determinants of sustained response to selection, the integration of QTL mapping methods into G -matrix estimation can provide a concrete, genetically based experimental program to investigate the evolutionary potential of natural populations.     

THE ONTOGENETIC TRAJECTORY OF THE PHENOTYPIC COVARIANCE MATRIX, WITH EXAMPLES FROM CRANIOFACIAL SHAPE IN RATS AND HUMANS

Many classic quantitative genetic theories assume the covariance structure among adult phenotypic traits to be relatively static during evolution. But the cross-sectional covariance matrix arises from the joint variation of a large range of developmental processes and hence is not constant over the period during which a population of developing organisms is actually exposed to selection. To examine how development shapes the phenotypic covariance structure, we ordinate the age-specific covariance matrices of shape coordinates for craniofacial growth in rats and humans. The metric that we use for this purpose is given by the square root of the summed squared log relative eigenvalues. This is the natural metric on the space of positive-definite symmetric matrices, which we introduce and justify in a biometric context. In both species, the covariance matrices appear to change continually throughout the full period of postnatal development. The resulting ontogenetic trajectories alter their direction at major changes of the developmental programs whereas they are fairly straight in between. Consequently, phenotypic covariance matrices—and thus also response to selection—should be expected to vary both over ontogenetic and phylogenetic time scales as different phenotypes are necessarily produced by different developmental pathways.     

EVOLUTION OF GENETIC INTEGRATION BETWEEN DISPERSAL AND COLONIZATION ABILITY IN A BIRD

Discrete behavioral strategies comprise a suite of traits closely integrated in their expression with consistent natural selection for such coexpression leading to developmental and genetic integration of their components. However, behavioral traits are often also selected to respond rapidly to changing environments, which should both favor their context-dependent expression and inhibit evolution of genetic integration with other, less flexible traits. Here we use a multigeneration pedigree and long-term data on lifetime fitness to test whether behaviors comprising distinct dispersal strategies of western bluebirds—a species in which the propensity to disperse is functionally integrated with aggressive behavior—are genetically correlated. We further investigated whether selection favors flexibility in the expression of aggression in relation to current social context. We found a significant genetic correlation between aggression and dispersal that is concordant with consistent selection for coexpression of these behaviors. To a limited extent, individuals modified their aggression to match their mate; however, we found no fitness consequences on such adjustments. These results introduce a novel way of viewing behavioral strategies, where flexibility of behavior, while often aiding an organism's fit in its current environment, may be limited and thereby enable integration with less flexible traits.     

Seasonal controls on interannual variability in carbon dioxide exchange of a near-end-of rotation Douglas-fir stand in the Pacific Northwest, 1997–2006

This study analyzes 9 years of eddy‐covariance (EC) data carried out in a Pacific Northwest Douglas‐fir (Pseudotsuga menzesii) forest (58‐year old in 2007) on the east coast of Vancouver Island, Canada, and characterizes the seasonal and interannual variability in net ecosystem productivity (NEP), gross primary productivity (GPP), and ecosystem respiration (R_e) and primary climatic controls on these fluxes. The annual values (± SD) of NEP, GPP and R_e were 357 ± 51, 2124 ± 125, and 1767 ± 146 g C m^?2 yr^?1, respectively, with ranges of 267–410, 1592–2338, and 1642–2071 g C m^?2 yr^?1, respectively. Spring to early summer (March–June) accounted for more than 80% of annual NEP while late spring to early autumn (May–August) was mainly responsible for its interannual variability (～80%). The major drivers of interannual variability in annual carbon (C) fluxes were annual and spring mean air temperatures (T_a) and water deficiency during late summer and autumn (July–October) when this Douglas‐fir forest growth was often water‐limited. Photosynthetically active radiation (Q), and the combination of Q and soil water content (θ) explained 85% and 91% of the variance of monthly GPP, respectively; and 91% and 96% of the variance of monthly R_e was explained by T_a and the combination of T_a and θ, respectively. Annual net C sequestration was high during optimally warm and normal precipitation years, but low in unusually warm or severely dry years. Excluding 1998 and 1999, the 2 years strongly affected by an El Niño/La Niña cycle, annual NEP significantly decreased with increasing annual mean T_a. Annual NEP will likely decrease whereas both annual GPP and R_e will likely increase if the future climate at the site follows a trend similar to that of the past 40 years.     

Change in CO2 balance under a series of forestry activities in a cool-temperate mixed forest with dense undergrowth

To evaluate the effects on CO₂ exchange of clearcutting a mixed forest and replacing it with a plantation, 4.5 years of continuous eddy covariance measurements of CO₂ fluxes and soil respiration measurements were conducted in a conifer-broadleaf mixed forest in Hokkaido, Japan. The mixed forest was a weak carbon sink (net ecosystem exchange, −44 g C m⁻² yr⁻¹), and it became a large carbon source (569 g C m⁻² yr⁻¹) after clearcutting. However, the large emission in the harvest year rapidly decreased in the following 2 years (495 and 153 g C m⁻² yr⁻¹, respectively) as the gross primary production (GPP) increased, while the total ecosystem respiration (RE) remained relatively stable. The rapid increase in GPP was attributed to an increase in biomass and photosynthetic activity of Sasa dwarf bamboo, an understory species. Soil respiration increased in the 3 years following clearcutting, in the first year mainly owing to the change in the gap ratio of the forest, and in the following years because of increased root respiration by the bamboo. The ratio of soil respiration to RE increased from 44% in the forest to nearly 100% after clearcutting, and aboveground parts of the vegetation contributed little to the RE although the respiration chamber measurements showed heterogeneous soil condition after clearcutting.     

Strong links between teleconnections and ecosystem exchange found at a Pacific Northwest old-growth forest from flux tower and MODIS EVI data

Variability in three Pacific teleconnection patterns are examined to see if net carbon exchange at a low‐elevation, old‐growth forest is affected by climatic changes associated with these periodicities. Examined are the Pacific Decadal Oscillation (PDO), Pacific/North American Oscillation (PNA) and El Niño‐Southern Oscillation (ENSO). We use 9 years of eddy covariance CO₂, H₂O and energy fluxes measured at the Wind River AmeriFlux site, Washington, USA and 8 years of tower‐pixel remote sensing data from the Moderate Resolution Imaging Spectroradiometer (MODIS) to address this question. We compute a new Composite Climate Index (CCI) based on the three Pacific Oscillations to divide the measurement period into positive‐ (2003 and 2005), negative‐ (1999 and 2000) and neutral‐phase climate years (2001, 2002, 2004, 2006 and 2007). The forest transitioned from an annual net carbon sink (NEP=+217 g C m^?2 yr^?1, 1999) to a source (NEP=?100 g C m^?2 yr^?1, 2003) during two dominant teleconnection patterns. Net ecosystem productivity (NEP), water use efficiency (WUE) and light use efficiency (LUE) were significantly different (P<0.01) during positive (NEP=?0.27 g C m^?2 day^?1, WUE=4.1 mg C g^?1 H₂O, LUE=0.94 g C MJ^?1) and negative (NEP=+0.37 g C m^?2 day^?1, WUE=3.4 mg C g^?1 H₂O, LUE=0.83 g C MJ^?1) climate phases. The CCI was linked to variability in the MODIS Enhanced Vegetation Index (EVI) but not to MODIS Fraction of absorbed Photosynthetically Active Radiation (FPAR). EVI was highest during negative climate phases (1999 and 2000) and was positively correlated with NEP and showed potential for using MODIS to estimate teleconnection‐driven anomalies in ecosystem CO₂ exchange in old‐growth forests. This work suggests that any increase in the strength or frequency of ENSO coinciding with in‐phase, low frequency Pacific oscillations (PDO and PNA) will likely increase CO₂ uptake variability in Pacific Northwest conifer forests.     

Patterns and controls of the variability of radiation use efficiency and primary productivity across terrestrial ecosystems

Aim The controls of gross radiation use efficiency (RUE), the ratio between gross primary productivity (GPP) and the radiation intercepted by terrestrial vegetation, and its spatial and temporal variation are not yet fully understood. Our objectives were to analyse and synthesize the spatial variability of GPP and the spatial and temporal variability of RUE and its climatic controls for a wide range of vegetation types. Location A global range of sites from tundra to rain forest. Methods We analysed a global dataset on photosynthetic uptake and climatic variables from 35 eddy covariance (EC) flux sites spanning between 100 and 2200 mm mean annual rainfall and between ?13 and 26°C mean annual temperature. RUE was calculated from the data provided by EC flux sites and remote sensing (MODIS). Results Rainfall and actual evapotranspiration (AET) positively influenced the spatial variation of annual GPP, whereas temperature only influenced the GPP of forests. Annual and maximum RUE were also positively controlled primarily by annual rainfall. The main control parameters of the growth season variation of gross RUE varied for each ecosystem type. Overall, the ratio between actual and potential evapotranspiration and a surrogate for the energy balance explained a greater proportion of the seasonal variation of RUE than the vapour pressure deficit (VPD), AET and precipitation. Temperature was important for determining the intra‐annual variability of the RUE at the coldest energy‐limited sites. Main conclusions Our analysis supports the idea that the annual functioning of vegetation that is adapted to its local environment is more constrained by water availability than by temperature. The spatial variability of annual and maximum RUE can be largely explained by annual precipitation, more than by vegetation type. The intra‐annual variation of RUE was mainly linked to the energy balance and water availability along the climatic gradient. Furthermore, we showed that intra‐annual variation of gross RUE is only weakly influenced by VPD and temperature, contrary to what is frequently assumed. Our results provide a better understanding of the spatial and temporal controls of the RUE and thus could lead to a better estimation of ecosystem carbon fixation and better modelling.     

Seasonal and interannual variations in carbon dioxide exchange over a cropland in the North China Plain

In China, croplands account for a relatively large form of vegetation cover. Quantifying carbon dioxide exchange and understanding the environmental controls on carbon fluxes over croplands are critical in understanding regional carbon budgets and ecosystem behaviors. In this study, the net ecosystem exchange (NEE) at a winter wheat/summer maize rotation cropping site, representative of the main cropping system in the North China Plain, was continuously measured using the eddy covariance technique from 2005 to 2009. In order to interpret the abiotic factors regulating NEE, NEE was partitioned into gross primary production (GPP) and ecosystem respiration (R_eco). Daytime R_eco was extrapolated from the relationship between nighttime NEE and soil temperature under high turbulent conditions. GPP was then estimated by subtracting daytime NEE from the daytime estimates of R_eco. Results show that the seasonal patterns of the temperature responses of R_eco and light‐response parameters are closely related to the crop phenology. Daily R_eco was highly dependent on both daily GPP and air temperature. Interannual variability showed that GPP and R_eco were mainly controlled by temperature. Water availability also exerted a limit on R_eco. The annual NEE was ?585 and ?533 g C m^?2 for two seasons of 2006–2007 and 2007–2008, respectively, and the wheat field absorbed more carbon than the maize field. Thus, we concluded that this cropland was a strong carbon sink. However, when the grain harvest was taken into account, the wheat field was diminished into a weak carbon sink, whereas the maize field was converted into a weak carbon source. The observations showed that severe drought occurring during winter did not reduce wheat yield (or integrated NEE) when sufficient irrigation was carried out during spring.     

The European carbon balance. Part 3: forests

We present a new synthesis, based on a suite of complementary approaches, of the primary production and carbon sink in forests of the 25 member states of the European Union (EU‐25) during 1990–2005. Upscaled terrestrial observations and model‐based approaches agree within 25% on the mean net primary production (NPP) of forests, i.e. 520±75 g C m^?2 yr^?1 over a forest area of 1.32 × 10⁶ km² to 1.55 × 10⁶ km² (EU‐25). New estimates of the mean long‐term carbon forest sink (net biome production, NBP) of EU‐25 forests amounts 75±20 g C m^?2 yr^?1. The ratio of NBP to NPP is 0.15±0.05. Estimates of the fate of the carbon inputs via NPP in wood harvests, forest fires, losses to lakes and rivers and heterotrophic respiration remain uncertain, which explains the considerable uncertainty of NBP. Inventory‐based assessments and assumptions suggest that 29±15% of the NBP (i.e., 22 g C m^?2 yr^?1) is sequestered in the forest soil, but large uncertainty remains concerning the drivers and future of the soil organic carbon. The remaining 71±15% of the NBP (i.e., 53 g C m^?2 yr^?1) is realized as woody biomass increments. In the EU‐25, the relatively large forest NBP is thought to be the result of a sustained difference between NPP, which increased during the past decades, and carbon losses primarily by harvest and heterotrophic respiration, which increased less over the same period.     

Estimation of gross primary production of irrigated maize using Landsat-8 imagery and Eddy Covariance data

A study was conducted to understand the potential of Landsat-8 in the estimation of gross primary production (GPP) and to quantify the productivity of maize crop cultivated under hyper-arid conditions of Saudi Arabia. The GPP of maize crop was estimated by using the Vegetation Photosynthesis Model (VPM) utilizing remote sensing data from Landsat-8 reflectance (GPP_VPM) as well as the meteorological data provided by Eddy Covariance (EC) system (GPP_EC), for the period from August to November 2015. Results revealed that the cumulative GPP_EC for the entire growth period of maize crop was 1871 g C m⁻². However, the cumulative GPP determined as a function of the enhanced vegetation index – EVI (GPP_EVI) was 1979 g C m⁻², and that determined as a function of the normalized difference vegetation index – NDVI (GPP_NDVI) was 1754 g C m⁻². These results indicated that the GPP_EVI was significantly higher than the GPP_EC (R² = 0.96, P = 0.0241 and RMSE = 12.6%). While, the GPP_NDVI was significantly lower than the GPP_EC (R² = 0.93, P = 0.0384 and RMSE = 19.7%). However, the recorded relative error between the GPP_EC and both the GPP_EVI and the GPP_NDVI was −6.22% and 5.76%, respectively. These results demonstrated the potential of the landsat-8 driven VPM model for the estimation of GPP, which is relevant to the productivity and carbon fluxes.